Endangered and Threatened Wildlife and Plants; Endangered Status for 49 Species From the Hawaiian Islands, 67786-67860 [2016-23112]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R1–ES–2015–0125;
4500030113]
RIN 1018–BB07
Endangered and Threatened Wildlife
and Plants; Endangered Status for 49
Species From the Hawaiian Islands
Fish and Wildlife Service,
Interior.
ACTION: Final rule.
AGENCY:
We, the U.S. Fish and
Wildlife Service (Service), determine
endangered status under the
Endangered Species Act of 1973 (Act),
as amended, for 10 animal species,
including the Hawaii DPS of the bandrumped storm-petrel (Oceanodroma
castro), the orangeblack Hawaiian
damselfly (Megalagrion xanthomelas),
the anchialine pool shrimp (Procaris
hawaiana), and seven yellow-faced bees
(Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana), and for 39
plant species from the Hawaiian Islands.
This rule adds these species to the
Federal Lists of Endangered and
Threatened Wildlife and Plants.
DATES: This rule is effective October 31,
2016.
ADDRESSES: This final rule is available
on the Internet at https://
www.regulations.gov and at https://
www.fws.gov/pacificislands. Comments
and materials we received, as well as
supporting documentation we used in
preparing this rule, are available for
public inspection at https://
www.regulations.gov, or, by
appointment, during normal business
hours at: U.S. Fish and Wildlife Service,
Pacific Islands Fish and Wildlife Office,
300 Ala Moana Boulevard, Honolulu, HI
96850; telephone 808–792–9400; or
facsimile 808–792–9581.
FOR FURTHER INFORMATION CONTACT:
Mary M. Abrams, Ph.D., Field
Supervisor, Pacific Islands Fish and
Wildlife Office, 300 Ala Moana
Boulevard, Honolulu, HI 96850;
telephone 808–792–9400; or facsimile
808–792–9581. Persons who use a
telecommunications device for the deaf
(TDD) may call the Federal Information
Relay Service (FIRS) at 800–877–8339.
SUPPLEMENTARY INFORMATION:
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SUMMARY:
Executive Summary
Why we need to publish a rule. Under
the Endangered Species Act (Act), a
species may warrant protection through
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listing if it is endangered or threatened
throughout all or a significant portion of
its range. Listing a species as an
endangered or threatened species can
only be completed by issuing a rule.
Critical habitat is to be designated, to
the maximum extent prudent and
determinable, for any species
determined to be an endangered or
threatened species under the Act.
This rule makes final the listing of 10
animal species (the Hawaii DPS of the
band-rumped storm-petrel
(Oceanodroma castro), the orangeblack
Hawaiian damselfly (Megalagrion
xanthomelas), the anchialine pool
shrimp (Procaris hawaiana), and seven
yellow-faced bees (Hylaeus anthracinus,
H. assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana)),
and 39 plant species (Asplenium
diellaciniatum (no common name,
NCN), Calamagrostis expansa (Maui
reedgrass), Cyanea kauaulaensis (NCN),
Cyclosorus boydiae (kupukupu makalii),
Cyperus neokunthianus (NCN),
Cyrtandra hematos (haiwale), Deparia
kaalaana (NCN), Dryopteris glabra var.
pusilla (hohiu), Exocarpos menziesii
(heau), Festuca hawaiiensis (NCN),
Gardenia remyi (nanu), Huperzia
stemmermanniae (NCN), Hypolepis
hawaiiensis var. mauiensis (olua),
Joinvillea ascendens ssp. ascendens
(ohe), Kadua fluviatilis (kamapuaa),
Kadua haupuensis (NCN), Labordia
lorenciana (NCN), Lepidium orbiculare
(anaunau), Microlepia strigosa var.
mauiensis (NCN), Myrsine fosbergii
(kolea), Nothocestrum latifolium (aiea),
Ochrosia haleakalae (holei),
Phyllostegia brevidens (NCN),
Phyllostegia helleri (NCN), Phyllostegia
stachyoides (NCN), Portulaca villosa
(ihi), Pritchardia bakeri (Baker’s loulu),
Pseudognaphalium sandwicensium var.
molokaiense (enaena), Ranunculus
hawaiensis (makou), Ranunculus
mauiensis (makou), Sanicula
sandwicensis (NCN), Santalum
involutum (iliahi), Schiedea diffusa ssp.
diffusa (NCN), Schiedea pubescens
(maolioli), Sicyos lanceoloideus
(anunu), Sicyos macrophyllus (anunu),
Solanum nelsonii (popolo), Stenogyne
kaalae ssp. sherffii (NCN), and
Wikstroemia skottsbergiana (akia), as
endangered species.
Delineation of critical habitat requires
identification of the physical or
biological features essential to the
species’ conservation. A careful
assessment of the biological needs of the
species and the areas that may have the
physical or biological features essential
for the conservation of the species and
that may require special management
considerations or protections, and thus
qualify for designation as critical
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habitat, is required. We require
additional time to analyze the best
available scientific data in order to
identify specific areas appropriate for
critical habitat designation and to
analyze the impacts of designating such
areas as critical habitat. Accordingly, we
find designation of critical habitat to be
‘‘not determinable’’ at this time.
The basis for our action. Under the
Act, we can determine that a species is
an endangered or threatened species
based on any of five factors: (A) The
present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; or (E)
other natural or manmade factors
affecting its continued existence. We
have determined that these 49 species
are experiencing population-level
impacts as the result of the following
current and ongoing threats:
• Habitat loss and degradation due to
urbanization; nonnative feral ungulates
(hoofed mammals, e.g., pigs, goats, axis
deer, black-tailed deer, mouflon, and
cattle); nonnative plants; wildfire; and
water extraction.
• Predation or herbivory by nonnative
feral ungulates, rats, slugs, bullfrogs,
Jackson’s chameleons, ants, and wasps.
• Stochastic events such as
landslides, flooding, drought, tsunami,
and hurricanes.
• Human activities such as
recreational use of anchialine pools,
dumping of nonnative fish and trash
into anchialine pools, and manmade
structures and artificial lighting.
• Vulnerability to extinction due to
small numbers of individuals and
occurrences and lack of regeneration.
• Competition with nonnative plants
and nonnative invertebrates.
Existing regulatory mechanisms and
conservation efforts are not adequate to
ameliorate the impacts of these threats
on any of the 49 species such that listing
is not warranted. Environmental effects
from climate change are likely to
exacerbate the impacts of these threats.
Peer review and public comment. We
sought comments from independent
specialists to ensure that our
designation is based on scientifically
sound data, assumptions, and analyses.
We invited these peer reviewers to
comment on our listing proposal. We
also considered all comments and
information we received during two
comment periods, including at one
public hearing.
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Previous Federal Actions
Please refer to the proposed listing
rule for the 49 species from the
Hawaiian Islands (80 FR 58820;
September 30, 2015) for a detailed
description of previous Federal actions
concerning these species.
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Summary of Comments and
Recommendations
On September 30, 2015, we published
a proposed rule to list 49 species (39
plants and 9 animals) from the
Hawaiian Islands as endangered
throughout their ranges and the Hawaii
population (distinct population segment
(DPS)) of the band-rumped storm-petrel
as endangered (80 FR 58820). The
comment period for the proposed rule
lasted 60 days, ending November 30,
2015 We published a public notice of
the proposed rule in the local Honolulu
Star Advertiser, West Hawaii Today,
Hawaii Tribune-Herald, Molokai
Dispatch, The Maui News, and The
Garden Island newspapers at the
beginning of the comment period. We
received two requests for a public
hearing. On January 22, 2016 (81 FR
3767), we reopened the comment period
for an additional 30 days, ending on
February 22, 2016, and we announced a
public meeting and public hearing for
the proposed rule. We again published
a public notice in local newspapers and
provided the public notice to local
media. For both comment periods, we
requested that all interested parties
submit comments or information
concerning the proposed listing of the
49 species. We contacted all appropriate
State and Federal agencies, county
governments, elected officials, scientific
organizations, and other interested
parties and invited them to comment.
The public meeting and hearing were
held in Hilo, Hawaii, on February 9,
2016.
During the comment periods, we
received a total of 41 unique public
comment letters (including comments
received at the public hearing) on the
proposed listing of the 49 species. Of
the 41 commenters, 21 were peer
reviewers, 3 were Federal agencies
(Hawaii Volcanoes National Park,
Haleakala National Park, and KalokoHonokohau and Puuhonua o Honaunau
National Historical Parks (NHPs)), 4
were State of Hawaii agencies (Hawaii
Department of Health, Hawaii
Department of Land and Natural
Resources Division of Aquatic
Resources, Hawaii Division of Forestry
and Wildlife, and Hawaii Department of
Hawaiian Home Lands), and 13 were
nongovernmental organizations or
individuals (including those who
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provided comments or testimony at the
public hearing). The National Park
Service (NPS) provided new
information about the numbers and
range of species in this rule that occur
on NPS lands, and about graduate
research on the orangeblack Hawaiian
damselfly. We appreciate the time and
effort taken by all commenters to submit
their views and information, and we
have incorporated all substantive new
information, e.g., from the National Park
Service, into this final rule. However,
we received some comments from the
public on the possible future
designation of critical habitat and on a
variety of other topics. To the extent
that comments do not pertain to the
proposed listing rule, we do not address
them in this final rule. In this final rule,
we address only those comments
relevant to the listing of the 49 species
from the Hawaiian Islands.
All substantive information related to
the listing action provided during the
comment periods has either been
incorporated directly into this final rule,
or is addressed below. For readers’
convenience, we have combined similar
comments into a single comment and
response.
Peer Review
In accordance with our peer review
policy published in the Federal Register
on July 1, 1994 (59 FR 34270), we
solicited expert opinions from 29
knowledgeable individuals with
scientific expertise on one or more of
the 49 Hawaiian Islands species, which
include 39 plants, a seabird, a
damselfly, an anchialine pool shrimp,
and seven yellow-faced bees, and their
habitats. This expertise also included
familiarity with the geographic region in
which these species occur and
conservation biology principles. We
received responses from 21 of these
individuals. We reviewed all comments
we received from the peer reviewers for
substantive issues and new information
regarding the 49 species. Of these 21
peer reviewers, 18 provided comments
or new information on one or more of
the 49 species. Ten peer reviewers
stated support for the proposed listing,
and 11 were neutral regarding the
proposed listing. These peer reviewers
generally supported our methodology
and conclusions. Peer reviewer
comments are either addressed below or
are incorporated into this final rule as
appropriate.
(1) Comment: One peer reviewer
stated that sea-level rise and coastal
inundation collectively are also
potential future threats to the welfare of
Procaris hawaiana, because they may
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cause further loss of anchialine pool
habitat.
Our Response: We have added sealevel rise and coastal inundation as
threats to P. hawaiana and its habitat
under the discussion in this rule titled
‘‘Climate Change’’ (Factor E. Other
Natural or Manmade Factors Affecting
Their Continued Existence).
(2) Comment: One peer reviewer
stated that because sea-level rise could
increase surface connectivity between
currently isolated anchialine pools,
invasion by nonnative fish would be
exacerbated.
Our Response: In this rule, we have
added surface connectivity to our
summary description of the status and
stressors to P. hawaiana as a factor
likely to exacerbate the threat posed by
nonnative fish to this species and its
anchialine pool habitat (see Anchialine
pool shrimp (Procaris hawaiana), under
Summary of Biological Status of the 49
Hawaiian Islands Species).
(3) Comment: One peer reviewer
recommended that the island of Lanai,
and coastal habitat, be included as
habitat for the band-rumped stormpetrel, as birds were observed during
the breeding season transiting this
habitat, which is conducive to nesting
where crevices and ledges are numerous
and can provide some protection from
feral cats (Felis catus), goats (Capra
hircus), and mouflon (Ovis gmelini
musimon).
Our Response: We have added coastal
habitat on Lanai in our description of
habitat for the band-rumped stormpetrel in this final listing rule.
(4) Comment: One peer reviewer
recommended that coastal habitat on
leeward east Maui be included for the
band-rumped storm-petrel, as remains
of a chick were found there in 1999.
Our Response: We understand that
coastal habitat on east Maui may be part
of the species’ historical range, but we
have not added coastal areas on leeward
east Maui as currently occupied habitat
for the band-rumped storm-petrel in this
final rule. Unlike coastal Lanai, in
coastal areas on leeward east Maui, no
indication of the species’ presence or
use of this habitat has been observed for
17 years.
(5) Comment: One peer reviewer
stated that predation by bullfrogs
(Lithobates catesbeianus) should be
included as a threat to the orangeblack
Hawaiian damselfly, and that impacts of
backswimmers (Notonectidae family)
and caddisflies (Trichoptera order) on
the damselfly are speculative.
Our Response: We have included in
this final rule that bullfrogs are a threat
to the orangeblack Hawaiian damselfly,
and clarified that the effects of
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predation by backswimmers and
caddisflies are not well understood.
Comments From State Agencies
(6) Comment: The Hawaii Department
of Land and Natural Resources’ Division
of Forestry and Wildlife did not
comment in support of, or in opposition
to, the proposed listing of the 49 species
from the Hawaiian Islands. District
botanists from Kauai, Oahu, Maui, and
Hawaii Island provided plant species
occurrence updates by island.
Our Response: We appreciate the
information provided regarding the 49
plant species from the Hawaiian Islands,
and have incorporated it into the
Summary of Biological Status of the 49
Hawaiian Islands Species for the
appropriate species in this final rule.
(7) Comment: The Hawaii Department
of Health acknowledged that protecting
wildlife and plants can often be
important for human and environmental
health. They further commented that
managing and controlling wild
ungulates is necessary for 95 percent of
these proposed plant species, the
orangeblack Hawaiian damselfly
(Megalagrion xanthomelas), and the
yellow-faced bees (Hylaeus spp.), but
that it is also essential to preventing
erosion, and, therefore, protecting water
quality. Fire is a natural process that is
now unnaturally frequent, intense, and
destructive to the Hawaiian Islands, in
part due to invasive grasses. Mitigating
wildfires is essential to caring for 38
percent of the plant species, the
damselfly, and yellow-faced bees, but it
also limits the release of air pollutants
that are known to be harmful to human
health. Protection of coastal and
wetland habitat such as that populated
by the anchialine pool shrimp (Procaris
hawaiana) limits further human
pressures on our sensitive coastlines
and aquatic environments.
Our Response: We agree that
managing and controlling ungulates
would provide significant conservation
benefits to listed plant and animal
species, and would also prevent erosion
and protect water quality of the islands
and near shore reefs. We also
acknowledge that nonnative grasses
contribute to the increase in numbers
and intensity of wildfires in Hawaii.
Protection of coastal habitat (through
nonnative plant and ungulate control,
and prevention of wildfires) would
provide a conservation benefit to the
anchialine pool shrimp, and to other
species that depend on coastal habitat.
(8) Comment: The Hawaii Department
of Land and Natural Resources Division
of Aquatic Resources concurred that the
information in the proposed rule for the
anchialine pool shrimp, Procaris
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hawaiana, is the most accurate and upto-date information available, and
supported listing the species as
endangered under the Act (16 U.S.C.
1531 et seq.).
Our Response: We appreciate this
support for the proposed listing of the
anchialine pool shrimp, Procaris
hawaiana.
(9) Comment: The Department of
Hawaiian Home Lands (DHHL) asked
that the Secretary of the Interior
consider the effects of designation of
endangered species that may potentially
have critical habitat on Hawaiian Home
Lands in a similar manner to the effects
such designation has on tribal lands,
including the impact on tribal
sovereignty. DHHL is aware that
Secretarial Order 3206, issued in June
1997, establishes guidelines for the
Service when dealing with Indian tribes
relating to endangered species.
Secretarial Order 3206 recognizes that,
in order to respect the cultural and
social aspects of Indian tribes, some
environmental restrictions on Indian
tribal lands are not appropriate, and it
calls on the Service to preserve
endangered species while respecting
tribal authority over their own lands.
While native Hawaiians are not an
‘‘Indian tribe’’ under the Order, DHHL’s
mission, to place native Hawaiians on
its lands for residential, agricultural,
and pastoral homesteading purposes, is
analogous to the circumstances of
Indian tribes. The Department also
recommends that the Secretaries of the
Interior and Commerce, in determining
endangered species and critical habitat
designations, consult directly with the
Hawaiian Homes Commission, DHHL,
Office of Native Hawaiian Relations,
and beneficiaries of the Hawaiian
Homes Commission Act to include
native intelligence and knowledge on
species, habitat, and place-based
management and protection.
Our Response: In accordance with the
President’s memorandum of April 29,
1994 (Government-to-Government
Relations With Native American Tribal
Governments; 59 FR 22951), Executive
Order 13175 (Consultation and
Coordination With Indian Tribal
Governments), and the Department of
the Interior’s manual at 512 DM 2, we
readily acknowledge our responsibility
to communicate meaningfully with
recognized Federal Tribes on a
government-to-government basis. In
accordance with Secretarial Order 3206
of June 5, 1997 (American Indian Tribal
Rights, Federal-Tribal Trust
Responsibilities, and the Endangered
Species Act), we readily acknowledge
our responsibilities to work directly
with tribes in developing programs for
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healthy ecosystems; to incorporate
native intelligence and knowledge of
species, habitat, and place-based
management and protection; to
acknowledge that tribal lands are not
subject to the same controls as Federal
public lands; to remain sensitive to
Indian culture; and to make information
available to tribes. In addition, a 2004
consolidated appropriations bill (Pub. L.
108–199, see section 148) established
the Office of Native Hawaiian Relations
within the Secretary of the Interior’s
Office, and its duties include
effectuating and implementing the
special legal relationship between the
Native Hawaiian people and the United
States, and fully integrating the
principle and practice of meaningful,
regular, and appropriate consultation
with the Native Hawaiian people by
assuring timely notification of and prior
consultation with the Native Hawaiian
people before any Federal agency takes
any actions that may have the potential
to significantly affect Native Hawaiian
resources, rights, or lands. A 2011
memorandum of understanding (MOU)
signed by the Department of the Interior
states that ‘‘Federal agencies are
required to consult with Native
Hawaiian organizations before taking
any action that may have the potential
to significantly affect Native Hawaiian
resources, rights, or lands.’’ Although
native Hawaiians are not technically a
‘‘recognized Federal tribe’’ as referenced
in the above Executive and Secretarial
Orders, we endeavor to fully engage and
work directly with native Hawaiians as
much as possible. At the time we
published our proposed rule (80 FR
58820; September 30, 2015), we notified
several Hawaiian organizations
including the DHHL, Kamehameha
Schools, the Office of Hawaiian Affairs,
the Kahoolawe Island Reserve
Commission (KIRC), and Kahea-The
Hawaiian-Environmental Alliance. We
contacted the Department of the
Interior’s Office of Native Hawaiian
Relations on September 28, 2015, to
inform them of our proposed listing
action. We also conducted in-person
meetings with staff of the Department of
Hawaiian Home Lands, Kamehameha
Schools, and KIRC. We considered all
comments and recommendations
provided by these organizations in
developing this final listing rule. At the
time we prepare a proposed critical
habitat rule for these species, we will
notify these groups and organizations,
and carefully consider any comments
and new information they provide
regarding habitat for these species.
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Public Comments
Seven public commenters supported
listing of all 49 Hawaiian Islands
species. Seven public commenters
opposed the listing of the 49 Hawaiian
Islands species, and one of these
commenters supported the intent of
listing but opposed designation of
critical habitat on their lands.
(10) Comment: One commenter
supported this rule because of the facts
and analysis stated in the proposed rule.
Two commenters stated that humans
need to be a voice for plants and
animals, and that this listing will
positively impact the conservation of
many animals and positively lead other
conversations in the right direction.
Our Response: We appreciate the
comments and believe that listing status
will help provide conservation benefits
to the species and their habitats.
(11) Comment: One commenter stated
that the 49 species also play a pivotal
role in promoting tourism and building
the economy of Hawaii and that they
deserve to be put onto the Lists of
Endangered and Threatened Wildlife
and Plants. Two commenters stated that
listing these species will attract wildlife
enthusiasts and nature lovers from all
around the world, and their spending
and tourism helps to build and maintain
sources of revenue in Hawaii; most
markets within the islands depend on
the tourism dollars that wildlife attracts.
Our Response: We do not consider
economic consequences in our
decisions to list or not list species as
endangered or threatened under the Act.
Section 4(b)(1)(a) of the Act specifies
that listing determinations be made
‘‘solely on the basis of the best scientific
and commercial data available.’’
(12) Comment: One commenter stated
that the potential negative impacts of
listing to landowners is very small, as
the vast majority of the habitat for these
rare species occurs on State and Federal
lands, or in private lands devoted to
conservation.
Our Response: We agree that many of
the 49 species occur or were known
from State and Federal lands, or in
undeveloped areas already dedicated to
conservation. However, listing a species
as endangered or threatened is based on
the species’ biological status; the
development of a proposed rule for
critical habitat for these species will be
completed in a separate rule, and the
effects of critical habitat on landowners
will be analyzed upon preparation of
that proposed rule.
(13) Comment: One commenter stated
that island residents have entirely lost
historical and cultural opportunities
and rights as a result of species
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protection enforcement and that those
in the field of endangered species
protection have a single focus, with
little or no concern for cultural and
historical values. Another commenter
stated that this listing would cause a
further loss for the public of cultural,
historical, and economic resources. A
third commenter stated that native
Hawaiian society believes they should
be able to manage their people, land,
and resources autonomously.
Our Response: Listing a species as
endangered or threatened does not
cause loss of historical and cultural
opportunities; in fact, it highlights the
need to protect the characteristics that
are unique to the Hawaiian Islands. We
acknowledge that some economic
impacts are a possible consequence of
listing a species under the Act; for
example, there may be costs to the
landowner associated with the
development of a habitat conservation
plan (HCP). In other cases, if the
landowner does not acquire a permit for
incidental take (for animals), the
landowner may choose to forego certain
activities on their property to avoid
violating the Act, resulting in potential
lost income. However, the Act does not
provide for the consideration of such
impacts when making a listing decision.
Section 4(b)(1)(a) of the Act specifies
that listing determinations be made
‘‘solely on the basis of the best scientific
and commercial data available.’’ The
language provided by Congress in the
Act thus precludes such costs from
consideration in association with a
listing determination. We work
collaboratively with private landowners,
and strongly encourage those with listed
species on their property to work with
us to develop incentive-based measures
such as strategic habitat areas (SHAs)
and HCPs, which have the potential to
provide conservation measures that
effect positive results for the species and
their habitat while providing regulatory
relief for landowners. The conservation
and recovery of endangered and
threatened species, especially of those
in Hawaii that occur nowhere else in the
world, and the ecosystems upon which
they depend, is the ultimate objective of
the Act, and the Service recognizes the
vital importance of voluntary,
nonregulatory conservation measures
that provide incentives for landowners
in achieving that objective. In regards to
land management by native Hawaiians,
see our response to Comment (9), above.
The Act does provide for the
consideration of potential economic
impacts in the course of designating
critical habitat (limited to activities that
are funded, authorized, or carried out by
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a Federal agency), and that analysis will
be conducted as we prepare a rule
proposing critical habitat for the multiisland species.
(14) Comment: Four commenters were
concerned that listing a species would
entail removal of nonnative species with
cultural significance, or removal of
those used for food and sport hunting,
and that control of nonnative ungulates
would not be conducted humanely.
Our Response: Habitat destruction
and modification by ungulates is a
threat to 37 of the 39 plants, and to 9
of the 10 animals proposed for listing.
Herbivory by ungulates is a threat to 27
of the 39 plants proposed for listing.
Hawaii was inhabited as early as the
2nd century; therefore, hunting of game
mammals is a relatively recent activity
(Tomich 1986, p. 1). The first
Polynesian settlers brought domestic
pigs of southeast Asia (Sus scrofa or a
species derived from Sus scrofa vittatus)
with them that were small in size,
domesticated, and allowed to run freely
around habitations (Tomich 1986, p.
120). Cook brought English pigs on his
first voyage to Hawaii and landed a boar
and sow on Niihau in 1778 (Tomich
1986, p. 121). Goats and European boars
were introduced and released (on
Niihau in 1778) by ship captains with
the intent of establishing feral
populations of these animals to be an
available food source in future visits to
the islands. Cattle (Bos taurus) and
domestic sheep (Ovis aries) were
released in 1794, by Vancouver. Deer
were released later; first, axis deer in
1867, and then mule deer (black-tailed
deer) in 1961 (Tomich 1986, pp. 127,
133, 141, 150, 158). These ungulates
multiplied rapidly, with immense
negative impacts to native vegetation
(Loope 1988, pp. 274–276). The need for
control of feral cattle was recognized as
early as 1918, by C.S. Judd (Tomich
1986, p. 146). The commenter may be
referring to the Federal court order
mandating the removal of sheep and
goats for protection of the palila
(Loxioides bailleui), an endangered bird
endemic to Hawaii. Aerial hunting is an
efficient control method and was chosen
by the State to comply with this order.
Carcasses taken during hunts (in both
2014 and 2015) were available to the
permitted public for salvage (DLNR
2014, in litt.; DLNR 2015, in litt.). Aerial
hunting is not conducted by the Service
in Hawaii.
(15) Comment: One commenter stated
that once species are listed for
protection under the Act, there is no
public recourse.
Our Response: There is public
recourse after a rulemaking is published
in the Federal Register. Under the Act,
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an interested person may petition to add
a species to, or to remove a species
from, either of the Lists of Endangered
and Threatened Wildlife and Plants.
Within 12 months of the petition, the
Secretary will make a finding as to
whether the petition presents
substantial scientific or commercial
information indicating that the
petitioned action may be warranted.
Persons may also petition to designate
or revise a critical habitat designation.
Our petition regulations are set forth at
50 CFR 424.14.
(16) Comment: Two commenters
expressed concern that the magnitude of
the proposed listing rule and the
subsequent designation of critical
habitat will have negative effects on
Hawaii’s economy, property values, and
land use.
Our Response: We understand there is
confusion and concern about the effects
of listing the 49 multi-island species.
Listing provides certain protections to
the species under the Act. Section 7 of
the Act states that each Federal agency
(through consultation) shall insure that
any action authorized, funded, or
carried out by the agency is not likely
to jeopardize the continued existence of
any endangered or threatened species.
For endangered species of fish or
wildlife, section 9 of the Act prohibits
any person subject to the jurisdiction of
the United States to import or export;
‘‘take’’ (defined as harass, harm, pursue,
hunt, shoot, wound, kill, trap, capture,
collect, or attempt any of these actions)
within the United States or the
territorial sea of the United States; take
upon the high seas; deliver, receive,
carry, transport, or ship in interstate or
foreign commerce in the course of a
commercial activity; or sell or offer for
sale in interstate or foreign commerce.
For endangered plants, section 9 of the
Act prohibits any person subject to the
jurisdiction of the United States to
import or export; deliver, receive, carry,
transport, or ship in interstate or foreign
commerce in the course of a commercial
activity; sell or offer for sale in interstate
or foreign commerce; remove and
reduce the species to possession from
areas under Federal jurisdiction;
maliciously damage or destroy any such
species on areas under Federal
jurisdiction; or remove, cut, dig up, or
damage or destroy any species species
in knowing violation of any State law or
regulations or in the course of any
violation of a State criminal trespass
law. Section 10 of the Act provides for
permitting of actions that may enhance
the propagation or survival of the
species, or that may ‘‘take’’ a species.
We acknowledge that some economic
impacts are a possible consequence of
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listing a species under the Act; for
example, there may be costs to the
landowner associated with the
development of an HCP. In other cases,
if the landowner does not acquire a
permit for incidental take, the
landowner may choose to forego certain
activities on their property to avoid
violating the Act, resulting in potential
lost income. However, the statute does
not provide for the consideration of
such impacts when making a listing
decision. Listing determinations are
made ‘‘solely on the basis of the best
scientific and commercial data
available.’’ This rule only lists the 49
species from the Hawaiian Islands; it
does not designate critical habitat.
(17) Comment: Two commenters
stated that listing species and
designating critical habitat on private
property in Hawaii will alienate
ranchers, a group that can help with
species and habitat conservation. The
commenters state that conservation can
best be achieved by cooperation and
coordination with private landowners.
Our Response: This rule only
addresses the listing of 49 species from
the Hawaiian Islands and does not
designate critical habitat. We agree that
partnerships can provide benefits for
listed species and their habitat through
development of conservation plans and
implementation of management actions.
(18) Comment: One commenter stated
that the Service should include the
public now, not after designating critical
habitat, with outreach, public forums,
presentations, and meetings on every
island for community groups, industry
and business groups, the Soil and Water
Conservation Districts, the Farm Bureau,
Hawaii Cattlemen’s Council, and
schools.
Our Response: As described above,
the publication of the proposed listing
rule did not include a critical habitat
proposal. We opened a 60-day comment
period on the proposed listing rule,
obtained extensive peer review,
published notices in numerous local
newspapers, reopened the comment
period, and held a public hearing and
information meeting. We considered all
comments we received in preparing this
final listing rule, and this rule
incorporates new, substantive
information provided to us by
commenters.
Summary of Changes From the
Proposed Rule
In preparing this final rule, we
reviewed and fully considered
comments from the public and peer
reviewers on the proposed rule, and
incorporated the following substantive
changes into this final rule. None of the
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new information we received changed
our evaluation of the threats to these
species or our determinations in this
final rule that they are endangered.
(1) We made revisions to the
demographic status or distribution of 31
species of plants, based on comments
from peer reviewers, by correcting
current locations or numbers of
individuals for: Asplenium
diellaciniatum, Calamagrostis expansa,
Cyanea kauaulaensis, Cyclosorus
boydiae, Cyrtandra hematos, Dryopteris
glabra var. pusilla, Exocarpos menziesii,
Gardenia remyi, Huperzia
stemmermanniae, Joinvillea ascendens
ssp. ascendens, Kadua fluviatilis,
Microlepia strigosa var. mauiensis,
Myrsine fosbergii, Nothocestrum
latifolium, Ochrosia haleakalae,
Phyllostegia brevidens, P. helleri, P.
stachyoides, Portulaca villosa,
Pritchardia bakeri, Pseudognaphalium
sandwicensium var. molokaiense,
Ranunculus hawaiensis, R. mauiensis,
Sanicula sandwicensis, Santalum
involutum, Schiedea diffusa ssp.
diffusa, S. pubescens, Sicyos
lanceoloideus, S. macrophyllus,
Stenogyne kaalae ssp. sherffii, and
Wikstroemia skottsbergiana.
(2) We made revisions to specific
threats to 31 plant species, based on
comments from peer reviewers,
including: Asplenium diellaciniatum,
Calamagrostis expansa, Cyanea
kauaulaensis, Cyclosorus boydiae,
Cyperus neokunthianus, Cyrtandra
hematos, Deparia kaalaana, Dryopteris
glabra var. pusilla, Exocarpos menziesii,
Huperzia stemmermanniae, Hypolepis
hawaiiensis var. mauiensis, Joinvillea
ascendens ssp. ascendens, Kadua
fluviatilis, K. haupuensis, Labordia
lorenciana, Lepidium orbiculare,
Microlepia strigosa var. mauiensis,
Myrsine fosbergii, Nothocestrum
latifolium, Ochrosia haleakalae,
Phyllostegia brevidens, P. helleri, P.
stachyoides, Portulaca villosa, Sanicula
sandwicensis, Santalum involutum,
Schiedea diffusa ssp. diffusa, S.
pubescens, Sicyos lanceoloideus,
Solanum nelsonii, and Wikstroemia
skottsbergiana.
(3) We corrected the taxonomy for the
nonnative plant, California grass, from
Brachiaria mutica to Urochloa mutica.
(4) We added further references
concerning genetic research that
supports differences in populations of
the band-rumped storm-petrel breeding
in different oceans and archipelagos.
(5) We added additional information
on current nesting sites of the bandrumped storm-petrel on Lehua Island,
Kauai, Molokai (coastal), Lanai (coastal),
Hawaii Island (Hawaii Volcanoes
National Park), and subalpine habitat
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(Hawaii Island), based on comments
regarding audio detections.
(6) We added information regarding
additional populations of the
orangeblack Hawaiian damselfly on
Hawaii Island.
(7) We added information on
predation of the orangeblack Hawaiian
damselfly by Jackson’s chameleons,
backswimmers, and bullfrogs as a threat,
and predation by the black twig borer as
a threat to Labordia lorenciana and
Nothocestrum latifolium.
(8) We added competition with
caddisflies for resources, prey, and
space as a potential threat to the
orangeblack Hawaiian damselfly.
(9) We made revisions to the
demographic status or distribution of
the yellow-faced bees Hylaeus
anthracinus, H. facilis, and H.
longiceps.
(10) We added tsunami as a threat to
the yellow-faced bees that occur in
coastal areas (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H.
longiceps), and to Solanum nelsonii,
also in coastal areas.
(11) We changed ‘‘Australian colletid’’
to ‘‘alien Hylaeus’’ bees, and included
competition with sweat bees
(Lasioglossum spp.) as a threat to the
yellow-faced bees.
(12) We noted that transmission of
diseases carried by nonnative insects
through shared food sources could be a
threat to the yellow-faced bees, but we
have no specific evidence of this type of
disease transmission.
(13) We added drought as a potential
threat to all seven yellow-faced bees.
(14) We added infiltration of waste
water, fertilizers, or pesticides resulting
from development activities as a
potential threat to the anchialine pool
shrimp.
(15) We added sea-level rise and
coastal inundation as a potential threat
to Solanum nelsonii, as occurrences in
low-lying coastal areas are at risk, and
to the anchialine pool shrimp, as these
events could increase connectivity of
anchialine pools leading to further
incursion by nonnative fish from one
pool to another.
67791
Background
Please refer to the proposed listing
rule for the 49 species from the
Hawaiian Islands (80 FR 58820;
September 30, 2015), available at https://
www.regulations.gov (see ADDRESSES),
for the following information:
• For background information on the
Hawaii Islands, see ‘‘The Hawaiian
Islands’’ under Background;
• For ecosystem descriptions, see An
Ecosystem-Based Approach To
Assessing the Conservation Status of the
49 Species in the Hawaiian Islands;
• For detailed descriptions of the
species and their taxonomy, see
Description of the 49 Hawaiian Islands
Species.
Hawaiian Islands Species Addressed in
This Final Rule
Table 1A (plants) and Table 1B
(animals), below, provide the common
name, scientific name, and range (by
Hawaiian Island) for the 49 species
addressed in this final rule.
TABLE 1A—PLANT SPECIES LISTED AS ENDANGERED
Common name
Hawaiian Island
Plants
Asplenium diellaciniatum ..................................................
Calamagrostis expansa ....................................................
Cyanea kauaulaensis .......................................................
Cyclosorus boydiae ..........................................................
Cyperus neokunthianus ....................................................
Cyrtandra hematos ...........................................................
Deparia kaalaana ..............................................................
Dryopteris glabra var. pusilla ............................................
Exocarpos menziesii .........................................................
Festuca hawaiiensis .........................................................
Gardenia remyi .................................................................
Huperzia stemmermanniae ...............................................
Hypolepis hawaiiensis var. mauiensis ..............................
Joinvillea ascendens ssp. ascendens ..............................
Kadua fluviatilis .................................................................
Kadua haupuensis ............................................................
Labordia lorenciana ..........................................................
Lepidium orbiculare ..........................................................
Microlepia strigosa var. mauiensis ...................................
Myrsine fosbergii ...............................................................
Nothocestrum latifolium ....................................................
Ochrosia haleakalae .........................................................
Phyllostegia brevidens ......................................................
Phyllostegia helleri ............................................................
Phyllostegia stachyoides ..................................................
Portulaca villosa ................................................................
asabaliauskas on DSK3SPTVN1PROD with RULES
Scientific name
No common name (NCN) ..
Maui reedgrass ..................
NCN ....................................
kupukupu makalii ...............
NCN ....................................
haiwale ...............................
NCN ....................................
hohiu ...................................
heau ...................................
NCN ....................................
nanu ...................................
NCN ....................................
olua .....................................
ohe .....................................
kamapuaa ...........................
NCN ....................................
NCN ....................................
anaunau .............................
NCN ....................................
kolea ...................................
aiea .....................................
holei ....................................
NCN ....................................
NCN ....................................
NCN ....................................
ihi ........................................
Pritchardia bakeri ..............................................................
Pseudognaphalium sandwicensium var. molokaiense .....
Ranunculus hawaiensis ....................................................
Ranunculus mauiensis ......................................................
Sanicula sandwicensis ......................................................
Santalum involutum ..........................................................
Schiedea diffusa ssp. diffusa ............................................
Schiedea pubescens ........................................................
Sicyos lanceoloideus ........................................................
Sicyos macrophyllus .........................................................
Solanum nelsonii ..............................................................
Baker’s loulu .......................
enaena ...............................
makou .................................
makou .................................
NCN ....................................
iliahi ....................................
NCN ....................................
maolioli ...............................
anunu .................................
anunu .................................
popolo .................................
Stenogyne kaalae ssp. sherffii .........................................
NCN ....................................
Kauai.
Hawaii, Maui.
Maui.
Hawaii (H), Maui, Oahu.
Maui (H).
Molokai.
Hawaii (H), Maui, Kauai (H).
Kauai.
Hawaii, Lanai (H).
Hawaii, Maui (H).
Hawaii, Maui, Molokai, Kauai.
Hawaii, Maui (H).
Maui.
Hawaii, Maui, Molokai, Oahu, Kauai.
Oahu, Kauai.
Kauai (H).
Kauai.
Kauai.
Hawaii, Maui, Oahu.
Oahu, Kauai.
Maui, Lanai (H), Molokai, Oahu, Kauai (H).
Hawaii, Maui.
Hawaii, Maui.
Kauai.
Hawaii (H), Maui, Molokai.
Hawaii, Maui, Kahoolawe, Lanai (H), Molokai, Oahu
(H), Kaula (H), Lehua (H), Nihoa (H).
Oahu.
Maui, Lanai (H), Molokai, Oahu (H).
Hawaii, Maui (H).
Hawaii (H), Maui, Molokai (H), Oahu (H), Kauai.
Hawaii, Maui.
Kauai.
Maui, Molokai (H).
Maui, Lanai (H), Molokai.
Oahu, Kauai.
Hawaii, Maui (H).
Hawaii, Maui (H), Molokai, Niihau (H), Pearl & Hermes,
Kure, Midway, Laysan, Nihoa.
Oahu (H).
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TABLE 1A—PLANT SPECIES LISTED AS ENDANGERED—Continued
Scientific name
Common name
Hawaiian Island
Wikstroemia skottsbergiana ..............................................
akia .....................................
Kauai.
(H) = historically known from island, but not observed in the past 20 years.
TABLE 1B—ANIMAL SPECIES LISTED AS ENDANGERED
Common name
Scientific name
Hawaiian Island
Animals
Band-rumped storm-petrel ..............
Oceanodroma castro .....................
Yellow-faced bee .............................
Yellow-faced bee .............................
Yellow-faced bee .............................
Yellow-faced bee .............................
Yellow-faced bee .............................
Yellow-faced bee .............................
Yellow-faced bee .............................
Orangeblack Hawaiian damselfly ....
Anchialine pool shrimp ....................
Hylaeus anthracinus ......................
Hylaeus assimulans .......................
Hylaeus facilis ................................
Hylaeus hilaris ...............................
Hylaeus kuakea .............................
Hylaeus longiceps .........................
Hylaeus mana ................................
Megalagrion xanthomelas .............
Procaris hawaiana .........................
Hawaii, Maui, Kahoolawe, Lanai, Molokai (H), Oahu (H), Kauai,
Lehua.
Hawaii, Maui, Kahoolawe, Lanai (H), Molokai, Oahu.
Maui, Kahoolawe, Lanai, Oahu (H).
Maui (H), Lanai (H), Molokai, Oahu.
Maui (H), Lanai (H), Molokai.
Oahu.
Maui, Lanai, Molokai, Oahu.
Oahu.
Hawaii, Maui, Lanai, Molokai, Oahu, Kauai (H).
Hawaii, Maui.
(H) = Historically known from the island, but not observed in the last 20 years.
asabaliauskas on DSK3SPTVN1PROD with RULES
Summary of Biological Status of the 49
Hawaiian Islands Species
The Act directs us to determine
whether any species is an endangered
species or a threatened species because
of any one of the factors listed in section
4(a)(1). We summarize, below, the
biological condition of, and factors
affecting, each of the 49 species and
determine whether each species is
endangered or threatened. The
summaries below include only brief
lists of factors affecting each species.
Each of these factors is fully considered,
in detail, in the subsequent section,
Summary of Factors Affecting the 49
Species From the Hawaiian Islands.
Climate Change Vulnerability
Assessment for the Hawaiian Plants
Twenty-seven of the plant species
described below were evaluated for
their vulnerability to climate change as
part of a comprehensive vulnerability
analysis of native Hawaiian plants, as
indicated in Table 2 (Fortini et al. 2013,
134 pp.). This analysis used ‘‘climate
envelopes’’ (geographic ranges
encompassing suitable climate for each
species, as defined by temperature and
moisture (Fortini et al. 2013, p. 17))
developed from field records by Price et
al. (2012) to project each species’
potential range in the year 2100. The
location and spatial extent of these
future ranges, and their overlap with
current ranges, allows calculation of a
vulnerability score. Estimates of
vulnerability based on climate-envelope
modeling are conservative in that they
do not take into account potential
changes in interspecific interactions
such as predation, disease, pollination,
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or competition. This study provides a
landscape- or island-scale picture of
potential climate-change vulnerability
of Hawaiian plants; the results are less
clear at finer spatial scales (Fortini et al.
2013, p. 42). However, all 27 of these
plant species scored moderately or
extremely vulnerable in the analysis
because of their relative inability to
exhibit the possible responses necessary
for persistence under projected climate
change (Fortini et al. 2013, 134 pp.).
These responses include the migration
response (dispersal and establishment
in new areas beyond their current
distribution), the microrefugia response
(persistence in topographically complex
areas that are less exposed),
evolutionary adaptation response
(morphological changes in response to
the changing environment), and
toleration response (adaptation to
environmental changes through
phenotypic plasticity). In the study,
response probabilities ranged from 0
(not vulnerable at all) to 1.0 (extremely
vulnerable; species likely to disappear
or ‘‘wink out’’ by the year 2100) (Fortini
et al. 2013, pp. 6–7). Many species
found to be moderately vulnerable in
this study, with scores of 0.5 or greater,
already are listed as endangered; some
already are extinct (Fortini et al. 2013,
pp. 24, 93). Therefore, because the
species in this rule were found by the
Fortini et al. (2013) study to be
moderately (0.5) to extremely (1.0)
vulnerable, we deem the likelihood of
their persistence to be low with the
impacts of climate change in addition to
other threats these species face. The
environmental changes associated with
climate change are likely to exacerbate
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these ongoing threats and further reduce
the likelihood that these species will
persist in the future.
Plants
Asplenium diellaciniatum (no
common name, NCN), a terrestrial or
epipetric (growing on rocks) fern in the
spleenwort family (Aspleniaceae), is
endemic to Kauai (Palmer 2003, p. 117).
Little is known of the historical
distribution of this species. It was
described from a collection from
‘‘Halemanu,’’ the Knudsen homestead
area on western Kauai. Currently, this
fern is found in montane mesic forest at
Kawaiiki and Kaluahaulu Ridge (Palmer
2003, p. 117; HBMP 2010; Lorence et al.
2013, p. 167) in 3 occurrences, totaling
approximately 100 individuals, 30 of
which are in an ungulate exclosure
(TNCH 2007; HBMP 2010; Lorence et al.
2013, p. 167; Wood 2013, in litt.; Plant
Extinction Prevention Program (PEPP)
2014, pp. 33, 59; Kishida 2015, in litt.;
Williams 2015, in litt.).
Feral pigs (Sus scrofa), goats (Capra
hircus), and black-tailed deer
(Odocoileus hemionus columbianus)
modify and destroy the habitat of
Asplenium diellaciniatum on Kauai,
with evidence of the activities of these
animals reported in the areas where A.
diellaciniatum occurs (Service 1999, p.
72; HBMP 2010). Feral pigs, goats, and
black-tailed deer also forage on A.
diellaciniatum. Ungulates are managed
in Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt; Hawaii Administrative
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Rule-Hawaii Department of Land and
Natural Resources (HAR–DLNR) 2010,
in litt.). Nonnative plants, such as
Adiantum hispidulum (rough
maidenhair fern), Blechnum
appendiculatum (no common name),
Erigeron karvinskianus (daisy fleabane),
and Rubus argutus (prickly Florida
blackberry), compete with A.
diellaciniatum, modify and destroy
native habitat, and displace native plant
species by competing for water,
nutrients, light, and space; they may
also produce chemicals that inhibit
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74;
Williams 2015, in litt.). Additionally,
the small number of individuals of A.
diellaciniatum limits this species’
ability to adapt to environmental
change.
The remaining occurrences of
Asplenium diellaciniatum are at risk; A.
diellaciniatum numbers are decreasing
on Kauai, and both the species and its
habitat continue to be negatively
affected by destruction and modification
by ungulates and by direct competition
by nonnative plants, combined with
herbivory by nonnative ungulates.
Because of the threats described above,
we find that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Calamagrostis expansa (Maui
reedgrass), a perennial in the grass
family (Poaceae), is known from the
islands of Maui and Hawaii (O’Connor
1999, p. 1509; Wagner and Herbst 2003,
p. 59). Historically, C. expansa was
known from wet forest, open bogs, and
bog margins on Maui at 17 locations on
east Maui, and in a large occurrence
covering nearly the entire summit on
west Maui, and was discovered in 7
occurrences totaling approximately 750
individuals on the island of Hawaii in
1995 (O’Connor 1999, p. 1509; HBMP
2010; Smithsonian National Museum of
Natural History (NMNH) Botany
Collections 2014, in litt.; Vetter 2015, in
litt.). Currently, this species is known
from 13 to 33 occurrences totaling fewer
than 750 individuals. This species is
rhizomatous (growing from
underground stems), making it difficult
to determine exact numbers of distinct
individuals and populations, and
botanists’ estimations vary. On the
island of Maui, there are 2 occurrences
in the west Maui Mountains
(approximately 100 individuals) and
from 7 to as many as 40 occurrences in
the east Maui Mountains (totaling at
least 200 individuals), often along ridges
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above 6,000 feet (ft) (1,830 meters (m)),
or on raised hummocks in wet forest
and bogs, in the montane wet ecosystem
(Wood 2005a, in litt.; TNCH 2007;
Welton 2008 and 2010, in litt.; Fay
2010, in litt.; HBMP 2010; Oppenheimer
2010, in litt.; Agorastos 2011, in litt.;
Vetter 2015, in litt.). Most of the east
Maui occurrences are in exclosures
(Duvall 2015, in litt.). On the island of
Hawaii, there are 3 occurrences in the
Kohala Mountains (totaling several
hundred individuals) and 1 occurrence
of 6 individuals last observed in 2004 in
Upper Waiakea Forest Reserve, in the
montane wet ecosystem (Perry 2006, in
litt; TNCH 2007; HBMP 2010; Perry
2015, in litt.).
Feral pigs modify and destroy the
habitat of Calamagrostis expansa on
Maui and Hawaii, with evidence of the
activities of feral pigs reported in the
areas where C. expansa occurs on east
Maui, and on Hawaii Island in the
Kohala Mountains and in the Waiakea
Forest Reserve (Hobdy 1996, in litt.;
Perlman 1996, in litt.; Wood 1996, in
litt.; Perry 2006, in litt.; HBMP 2010).
Some occurrences on east and west
Maui are currently fenced; however,
ungulate and weed control activities
must be maintained to provide
continued protection (Duvall 2015, in
litt.). Ungulates are managed in Hawaii
as game animals, but public hunting
does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Rats have been noted by biologists
as a threat to C. expansa at
Laupahoehoe Natural Area Reserve
(NAR) on Hawaii Island, by consuming
seeds (HBMP 2010). Nonnative plants
compete with this species and modify
and destroy native habitat, negatively
affecting C. expansa on east and west
Maui and Hawaii Island. Additionally,
the small number of individuals limits
this species’ ability to adapt to
environmental change. Fortini et al.
(2013, p. 68) found that, as
environmental conditions are altered by
climate change, C. expansa is unlikely
to tolerate or adapt to projected changes
in temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to C.
expansa described above (see ‘‘Climate
Change’’ under Factor E. Other Natural
or Manmade Factors Affecting Their
Continued Existence, below).
The remaining occurrences of
Calamagrostis expansa are at risk; C.
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expansa populations are decreasing on
Maui and Hawaii Island, and this
species continues to be negatively
affected by habitat modification and
destruction by feral pigs, and by direct
competition from nonnative plants,
combined with herbivory by feral pigs
and rats. This species is vulnerable to
the effects of climate change, and the
likelihood of its persistence with the
impacts of climate change, exacerbated
by the ongoing threats, is low. We find
that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Cyanea kauaulaensis (NCN), a shrub
in the bellflower family
(Campanulaceae), is endemic to Maui
(Oppenheimer and Lorence 2012, p. 15).
Cyanea kauaulaensis occurs on leeward
west Maui, on talus or basalt boulderstrewn slopes along perennial streams
from 2,400 to 3,000 ft (730 to 900 m),
in the lowland wet ecosystem (TNCH
2007; HBMP 2010; Oppenheimer and
Lorence 2012, pp. 17–18). This species
was first collected during a botanical
survey in 1989. Further surveys (in
2008, 2009, and 2011) revealed more
individuals, and study of the collections
indicated that it was a new species of
Cyanea. Currently, C. kauaulaensis is
known from Kauaula Valley
(approximately 100 individuals)
(Oppenheimer and Lorence 2012, pp.
15–16, 20; Duvall 2015, in litt.;
Oppenheimer 2015, in litt.).
The greatest threats to this species
currently are the low numbers of
occurrences and individuals, its limited
range, poor seedling recruitment, and
loss of pollinators and dispersal agents
(Oppenheimer and Lorence 2012, pp.
20–21; Duvall 2015, in litt.). Rats and
slugs are noted as a threat to Cyanea
kauaulaensis because of their herbivory
and seed predation. Additionally,
nonnative plants modify and destroy
native habitat and outcompete native
species, negatively affecting C.
kauaulaensis and its habitat. Although
feral ungulates are present on west
Maui, the known occurrences of C.
kauaulaensis may be less at risk from
this particular threat because of their
location in extremely steep and rugged
terrain; however, erosion, landslides,
flooding, and drying due to climate
change affect this species because of the
terrain where it occurs (Oppenheimer
and Lorence 2012, pp. 20–21; Duvall
2015, in litt.). The remaining occurrence
of Cyanea kauaulaensis is at risk.
Because of the threats described above,
we find that this species is endangered
throughout all of its range, and,
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therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Cyclosorus boydiae (previously
Christella boydiae) (kupukupu makalii)
is a small to medium-sized member of
the thelypteroid fern family
(Thelypteridaceae) (Pukui and Elbert
1986, p. 186; Palmer 2003, pp. 87–88).
Typical habitat for C. boydiae is
exposed, rocky, or moss-covered banks
of stream courses in dense-wet
Metrosideros-Acacia (ohia-koa) forest,
from 2,300 to 4,400 ft (700 to 1,350 m),
with other native ferns, grasses, and
dwarfed woody species, in the lowland
wet and montane wet ecosystems
(Hillebrand 1888, p. 572; Medeiros et al.
1993, p. 87; Wagner (W.H.) et al. 1999,
p. 156; TNCH 2007; HBMP 2010; Gates
2015, in litt.). Historically, this fern was
known from near sea level to 4,400 ft
(1,350 m) on Oahu, Maui, and Hawaii
Island (Hillebrand 1888, p. 572;
Medeiros et al. 1993, pp. 86–87; Palmer
2003, pp. 87–88). Currently, C. boydiae
is found on Oahu and east Maui, in 13
occurrences totaling approximately 400
individuals (Palmer 2003, pp. 87–88;
Oppenheimer 2008, in litt.; Fay 2010, in
litt.; HBMP 2010; Welton 2010, in litt.).
On east Maui, there are at least 11
occurrences (over 1,000 individuals) in
the lowland wet and montane wet
ecosystems, and on Oahu there are 2
occurrences in the Koolau Mountains in
the montane wet ecosystem, totaling 40
individuals, and one historic occurrence
in Kaluanui Drainage, but the status of
the species at this location is currently
unknown (Palmer 2003, pp. 87–88;
Wood 2007a, in litt.; Kam 2008, in litt.;
Oppenheimer 2008 and 2010, in litt.;
HBMP 2010; Welton 2010, in litt.; Ching
2011, in litt.; Ching Harbin 2015, in litt.;
Oppenheimer 2015, in litt.). The
historical occurrence of C. boydiae on
the island of Hawaii was found in the
lowland wet ecosystem (HBMP 2010).
Feral pigs modify and destroy the
habitat of Cyclosorus boydiae on Maui
and Oahu, with evidence of their
activities reported at three occurrences
of C. boydiae on east Maui and at two
occurrences on Oahu. However, on east
Maui, two of the five occurrences are
provided protection in Haleakala
National Park (Wood 2007a, in litt.;
HBMP 2010; Kawelo 2011, in litt.).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Historical occurrences of C.
boydiae on Oahu have dramatically
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declined in numbers or disappeared as
a result of habitat modification and
destruction, landslides and flooding,
invasion of lower elevation stream
courses by nonnative plants, and
manmade stream diversions (Medeiros
et al. 1993, p. 88; Palmer 2003, p. 88).
Nonnative plants, such as Tibouchina
herbacea (glorybush), modify and
destroy native habitat of. C. boydiae and
outcompete this and other native
species for water, nutrients, light, and
space (Smith 1985, pp. 180–250;
Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74). Herbivory by feral
pigs negatively impacts this species
(HBMP 2010). This species occurs on
stream banks at or just above water
level, and flash floods or drought can
damage and destroy it (Ching Harbin
2015, in litt.). Fortini et al. (2013, p. 72)
found that, as environmental conditions
are altered by climate change, C.
boydiae is unlikely to tolerate or adapt
to projected changes in temperature and
moisture, and is unlikely to be able to
move to areas with more suitable
climatic conditions. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to C. boydiae
described above.
The remaining occurrences of
Cyclosorus boydiae are at risk; C.
boydiae populations are decreasing on
Oahu and Maui, and the species
continues to be negatively affected by
habitat loss and destruction by
ungulates, direct competition with
nonnative plants, and herbivory by
ungulates. Flash floods and drought can
damage and destroy this species. The
effects of climate change are likely to
further exacerbate these threats. Because
of the threats describe above, we find
that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Cyperus neokunthianus (NCN) is a
perennial plant in the sedge family
(Cyperaceae) (Koyama 1999, p. 1420).
Cyperus neokunthianus occurs in
riparian areas of the lowland wet
ecosystem on west Maui (Koyama 1999,
p. 1420; TNCH 2007; HBMP 2010).
Historically, this species was known
from Honokohau Falls and Waihee
Valley (HBMP 2010; Global Biodiversity
Information Facility (GBIF) database
2014, in litt.). This species was last
observed in 1996. Currently, there are
no known individuals in the wild;
however, Waihee Valley and Maui
County lands have been suggested as
potential habitat for further surveys
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(PEPP 2013, p. 32; PEPP 2014, p. 59;
Duvall 2015, in litt.).
Feral pigs modify and destroy the
habitat of Cyperus neokunthianus on
west Maui, with evidence of the
activities of feral pigs reported in the
area where this species was last
observed (HBMP 2010). Habitat
modifications resulting from activities
of feral pigs that affect C. neokunthianus
include direct destruction of this
species and other native plants,
disruption of topsoil leading to erosion,
and establishment and spread of
nonnative plants. Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Additionally,
nonnative plants modify and destroy
native habitat and outcompete native
species, also negatively affecting habitat
of C. neokunthianus on west Maui.
Currently, there are no known extant
individuals, and low numbers makes
this species more vulnerable to
extinction because of the higher risks
from genetic bottlenecks, random
demographic fluctuations, and localized
catastrophes.
Cyperus neokunthianus is at risk and
continues to be negatively affected by
modification and destruction by
nonnative animals and plants (Duvall
2015, in litt.). Because of the threats
described above, we find that this
species is endangered throughout all of
its range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Cyrtandra hematos (haiwale), a shrub
in the African violet family
(Gesneriaceae), is endemic to Molokai
(Wagner et al. 1999, pp. 760, 762).
Cyrtandra hematos occurs in wet forest
from 3,400 to 3,800 ft (1,030 to 1,150 m)
on eastern Molokai, in the montane wet
ecosystem (Wagner et al. 1999, pp. 760,
762; HBMP 2010; TNCH 2007).
Historically, this species was known
from four locations on Molokai (Wagner
et al. 1999, pp. 760, 762). Currently,
there are fewer than 100 individuals at
two locations on Molokai (Duvall 2015,
in litt.; Oppenheimer 2015, in litt.).
Feral pigs and goats modify and
destroy the habitat of Cyrtandra
hematos on Molokai, with evidence of
the activities of these animals reported
in the areas where this species occurs
(Service 2015, in litt.). Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
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destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Additionally,
nonnative plants modify and destroy
native habitat and outcompete this and
other native species for water, nutrients,
light, and space, or a nonnative plant
may produce chemicals that inhibit
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; Service
2015, in litt.). This species experiences
reduced reproductive vigor due to low
numbers and lack of regeneration,
leading to diminished capacity to adapt
to environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
This species hybridizes with C. grayana
(Oppenheimer 2015, in litt.). Fortini et
al. (2013, p. 72) found that, as
environmental conditions are altered by
climate change, C. hematos is unlikely
to tolerate or adapt to projected changes
in temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to C.
hematos described above.
The remaining occurrences of
Cyrtandra hematos are at risk. The
known individuals are restricted to a
small area on Molokai and continue to
be negatively affected by habitat
modification and destruction by
ungulates and nonnative plants, and by
direct competition with nonnative
plants. The low number of remaining
individuals limits this species’ ability to
adapt to environmental changes.
Hybridization results in a reduction of
the numbers of C. hematos. The effects
of climate change are likely to further
exacerbate these threats. We find that
this species is endangered throughout
all of its range, and, therefore, find that
it is unnecessary to analyze whether it
is endangered or threatened in a
significant portion of its range.
Deparia kaalaana (NCN), a small,
terrestrial fern in the ladyfern family
(Athyriaceae), is recognized as a distinct
taxon by Palmer (2003, pp. 109–111)
and Christenhusz et al. (2012, p. 16).
This fern is historically known from the
islands of Kauai, Maui, and Hawaii, on
rocky stream banks and in wet forest, in
the lowland mesic and lowland wet
ecosystems (Palmer 2003, pp. 109–111;
TNCH 2007; HBMP 2010; Oppenheimer
and Bustamente 2014, p. 103; PEPP
2014, p. 95). Deparia kaalaana was
presumed extinct on all three islands
where it previously occurred until one
individual was discovered on east Maui,
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growing along a perennial stream on the
western side of a small pool with other
native ferns and herbaceous plants
(Oppenheimer and Bustamente 2014,
pp. 103–107; PEPP 2014, p. 95).
Feral pigs modify and destroy habitat
of Deparia kaalaana by facilitating the
spread of nonnative plants, which
converts vegetation communities from
native to nonnative (Cuddihy and Stone
1990, p. 63; Oppenheimer and
Bustamente 2014, p. 106). Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt;
HAR–DLNR 2010, in litt.). Nonnative
plants, such as Blechnum
appendiculatum (NCN), Clidemia hirta
(Koster’s curse), Hedychium
gardnerianum (kahili ginger), Prunella
vulgaris (selfheal), and Rubus argutus,
are capable of displacing all of the
riparian habitat elements, including
native plants, in the area where D.
kaalaana occurs. Nonnative slugs such
as Derocerus laeve and Limax maximus
are common in the area and can
consume young plants (Joe and Daehler
2008, pp. 252–253). Flash floods and
drought can damage and destroy this
species at its only known location. A
single catastrophic event may result in
extirpation of the remaining individual.
The remaining occurrence of Deparia
kaalaana is at risk, and both the species
and its habitat on Hawaii, Maui, and
Kauai continues to be negatively
affected by modification and destruction
by nonnative ungulates, and by direct
competition with nonnative plants,
combined with herbivory by nonnative
ungulates and slugs. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to D. kaalaana
described above. We find that this
species is endangered throughout all of
its range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Dryopteris glabra var. pusilla (hohiu)
is a small, terrestrial fern in the wood
fern family (Dryopteridaceae) (Palmer
2003, p. 144). Habitat for D. glabra var.
pusilla is deep shade on rocky, mossy
streambanks in wet forest at about 4,000
ft (1,200 m), in the montane wet
ecosystem on Kauai (Palmer 2003, p.
144; TNCH 2007; HBMP 2010).
Historically, D. glabra var. pusilla was
known from the Kawaikoi stream area
(HBMP 2010). Currently, this variety is
known from fewer than 250 individuals
in the Alakai Wilderness Preserve on
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Kauai (National Tropical Botanical
Garden (NTBG) Herbarium Database
1995, in litt.; HBMP 2010; Wood 2015,
in litt.).
Dryopteris glabra var. pusilla is at risk
from habitat modification and
destruction by nonnative plants, feral
pigs, and black-tailed deer (Wood 2015,
in litt.). Most individuals occur in the
Alakai Wilderness Preserve; however,
only portions of the Preserve are fenced
to prevent ungulate incursion.
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants modify and
destroy native habitat and outcompete
this and other native species for water,
nutrients, light, and space, or a
nonnative plant may produce chemicals
that inhibit growth of other plants, also
negatively affecting habitat of D. glabra
var. pusilla (Smith 1985, pp. 180–250;
Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74). Herbivory by rats
and slugs is a threat to D. glabra var.
pusilla (Wood 2015, in litt.). In addition,
the limited number of occurrences and
few individuals lead to a diminished
capacity to adapt to environmental
changes, thereby lessening the
probability of long-term persistence, and
a single catastrophic event may result in
extirpation of remaining occurrences.
Landslides along streambanks have been
known to destroy populations of this
fern (Wood 2015, in litt.).
Fortini et al. (2013, p. 74) found that,
as environmental conditions are altered
by climate change, D. glabra var. pusilla
is unlikely to tolerate or adapt to
projected changes in temperature and
moisture, and is unlikely to be able to
move to areas with more suitable
climatic conditions. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to D. glabra var.
pusilla described above. Because of
these threats, we find that this variety is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Exocarpos menziesii (heau) is a shrub
in the sandalwood family (Santalaceae)
(Wagner et al. 1999, p. 1218). This
species occurs in Metrosideros
shrubland or drier forest areas, and on
lava flows with sparse vegetation, from
4,600 to 6,900 ft (1,400 to 2,100 m), in
the montane dry ecosystem on the
island of Hawaii (Wagner et al. 1999, p.
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1218; TNCH 2007), and historically
occurred in the lowland mesic (Lanai
and Hawaii Island) and montane mesic
ecosystems (Hawaii Island) (TNCH
2007; Bishop Museum 2014). Exocarpos
menziesii is historically known from the
island of Lanai and was wide-spread on
the island of Hawaii (Wagner et al. 1999,
p. 1218; TNCH 2007; Bishop Museum
2014). Currently, there are seven
scattered occurrences on Hawaii Island,
six of which consist of only a few
individuals, the seventh totals an
estimated 1,800 individuals (PEPP 2013,
pp. 10, 33; Thomas 2014, in litt.; Evans
2015a, in litt.; Orlando 2015, in litt.;
Perry 2015, in litt.). There are no
currently known occurrences of this
species on Lanai.
Feral goats, mouflon, and sheep
modify and destroy the habitat of
Exocarpos menziesii on Hawaii Island,
and may forage on this species, with
evidence of the activities of these
animals reported in the areas where this
species occurs (Service 2015, in litt.).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt; HAR–DLNR 2010, in
litt.). Feral ungulate management is
incorporated into the U.S. Army’s
Pohakuloa Training Area (PTA)
management plan, and plants at PTA
may be provided some protection
within fenced management units in the
training area (Evans 2015a, in litt.);
however, it is reported that feral goats
are still being removed from within the
fenced area (Nadig 2015, in litt.). Any
individuals of E. menziesii outside of
fenced exclosures or outside of the
managed area are at risk. Additionally,
nonnative plants modify and destroy
native habitat and outcompete this and
other native species for water, nutrients,
light, and space, or a nonnative plant
may produce chemicals that inhibit
growth of other plants, also negatively
affecting habitat of E. menziesii (Smith
1985, pp. 180–250; Vitousek et al. 1987
in Cuddihy and Stone 1990, p. 74).
Occurrences and numbers of
individuals have declined on the island
of Hawaii (HBMP 2010; Thomas 2014,
in litt.), where E. menziesii was once
widely distributed from the south to the
west sides of the island, and are now
restricted to seven locations.
Consequently, E. menziesii experiences
reduced reproductive vigor due to
reduced levels of genetic variability,
leading to diminished capacity to adapt
to environmental changes, thereby
reducing the probability of long-term
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persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fire is a likely threat to this species;
although the U.S. Army has constructed
firebreaks and has standard operating
procedures (SOPs) in place for
prevention and suppression of wildfires
at the PTA, wildfires may encroach from
other areas (U.S. Army Garrison 2013, in
litt.). The small number of individuals
outside the larger occurrence at the PTA
limits this species’ ability to adapt to
environmental changes. Fortini et al.
(2013, p. 76) found that, as
environmental conditions are altered by
climate change, E. menziesii is unlikely
to tolerate or adapt to projected changes
in temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to E.
menziesii described above.
The remaining occurrences of
Exocarpos menziesii are at risk from
modification and destruction by feral
goats, mouflon, and sheep; from
herbivory by these ungulates; and by the
small number of remaining occurrences.
Fire is a likely threat to this species. The
effects of climate change are likely to
exacerbate these threats. Because of
these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Festuca hawaiiensis (NCN) is a
cespitose (growing in tufts or clumps)
annual in the grass family (Poaceae)
(O’Connor 1999, p. 1547). Typical
habitat for this species is dry forest at
6,500 ft (2,000 m), in the montane dry
ecosystem (O’Connor 1999, p. 1547).
Historically, F. hawaiiensis occurred at
Hualalai and Puu Huluhulu on the
island of Hawaii, and possibly at
Ulupalakua on Maui; however, it is no
longer found at these sites (O’Connor
1999, p. 1547). Currently, F. hawaiiensis
is only known from the U.S. Army’s
PTA on the island of Hawaii (HBMP
2010). These remaining four
occurrences are within an area of less
than 10 square miles (mi) (26 square
kilometers (km)) and total
approximately 1,500 individuals (U.S.
Army Garrison 2013, in litt.; Evans
2015a, in litt.).
Habitat destruction and modification
by feral goats and sheep is a threat to
Festuca hawaiiensis. These ungulates
also browse on native plants such as
grasses, including F. hawaiiensis.
Ungulates are managed in Hawaii as
game animals, but public hunting does
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not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Feral ungulate management is
incorporated into the U.S. Army’s PTA
management plan, and these plants are
provided some protection within fenced
management units in the training area
(Evans 2015a, in litt.); however, feral
goats are still being removed from inside
the fenced area (Nadig 2015, in litt.). In
addition, any individuals of F.
hawaiiensis outside of fenced
exclosures or outside of the managed
area are at risk. Nonnative plants, such
as Cenchrus setaceus (Pennisetum
setaceum; fountain grass), are
naturalized in the area and outcompete
F. hawaiiensis and other native plants.
Occurrences and numbers of
individuals are declining on the island
of Hawaii, and F. hawaiiensis
experiences reduced reproductive vigor
due to reduced levels of genetic
variability, leading to diminished
capacity to adapt to environmental
changes, thereby reducing the
probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman
and Pilson 1997, p. 361; HBMP 2010).
Fire is a likely threat to this species,
especially because of the ingress of
nonnative grass species. Although the
U.S. Army has constructed firebreaks
and has SOPs in place for prevention
and suppression of wildfires at the PTA,
fires may encroach from other areas,
exacerbated by fuel loads provided by
nonnative grasses (U.S. Army Garrison
2013, in litt.). Fortini et al. (2013, p. 76)
found that, as environmental conditions
are altered by climate change, F.
hawaiiensis is unlikely to tolerate or
adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to F.
hawaiiensis described above.
The remaining occurrences of Festuca
hawaiiensis are at risk; F. hawaiiensis
occurrences have decreased on Hawaii
Island, as it no longer occurs at Hualalai
and Puu Huluhulu, and the species may
be extirpated from Maui. This species
continues to be negatively affected by
habitat modification and destruction by
ungulates and by direct competition
with nonnative plants, combined with
herbivory by ungulates. Fire is a likely
threat to the species and its habitat. The
effects of climate change are likely to
further exacerbate these threats. Because
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of the threats described above, we find
that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Gardenia remyi (nanu) is a tree in the
coffee family (Rubiaceae) (Wagner et al.
1999, p. 1133). Typical habitat for G.
remyi is mesic to wet forest from 190 to
3,000 ft (60 to 760 m), in the lowland
mesic (Kauai, Molokai, and Hawaii
Island) and lowland wet ecosystems
(Kauai, Molokai, Maui, and Hawaii
Island) (Wagner et al. 1999, p. 1133;
TNCH 2007; HBMP 2010; Oppenheimer
2015, in litt.). Historically, this species
was found on the island of Hawaii at
Wao Kele O Puna NAR, Waiakea Forest
Reserve, Pahoa, and Hakalau Nui. On
Maui, this species was known from
Wailuaiki and Waikamoi in the Koolau
Forest Reserve, and from Papaaea and
Kipahulu. On Molokai, this species was
known from Keopukaloa, Pukoo,
Honomuni, Halawa, and Kaluaaha
(HBMP 2010). On Kauai, this species
ranged across the island, and was
known from Halelea, Kealia, Moloaa,
and Lihue-Koloa Forest Reserves,
including Hanakapiai Valley,
Mahaulepu, and east Wahiawa Bog.
Currently, G. remyi is known from 16
occurrences totaling approximately 90
individuals on the islands of Hawaii,
Maui, Molokai, and Kauai (Wood 2005b,
in litt.; Oppenheimer 2006, in litt; Perry
2006, in litt.; Welton 2008, in litt.;
Agorastos 2010, in litt.; HBMP 2010;
Perlman 2010, in litt.). An occurrence
on east Maui has been observed to
decline from 14 individuals in 1992, to
only 1 individual by 2015 (Duvall 2015,
in litt.).
Habitat modification and destruction
by feral pigs, goats, and axis deer
negatively affects Gardenia remyi and
areas suitable for its reintroduction
(Perry, in litt. 2006; PEPP 2008, p. 102;
HBMP 2010). Feral pigs and signs of
their activities have been reported at
occurrences of G. remyi on the island of
Hawaii, on Kauai, on east and west
Maui, and on Molokai. Goats and signs
of their activities are reported at the
occurrences G. remyi on Kauai and
Molokai. Axis deer and signs of their
activities are reported at the occurrences
of G. remyi on Molokai (HBMP 2010).
Herbivory by these ungulates is a threat
to G. remyi, as they browse on leaves
and other parts of almost any woody or
fleshy plant species. Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
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animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Nonnative
plants, such as Clidemia hirta,
Hedychium gardnerianum, Psidium
cattleianum (strawberry guava), and
Tibouchina herbacea on Hawaii Island
(Perry 2006, in litt.); Lantana camara
(lantana), Psidium guajava, and Rubus
argutus on Kauai (Wood 2004, in litt.);
Ageratina adenophora (Maui
pamakani), Rubus rosifolius
(thimbleberry), and T. herbacea on Maui
(HBMP 2010); and C. hirta and P.
cattleianum on Molokai (HBMP 2010),
modify and destroy native habitat of G.
remyi and outcompete this and other
native plants for water, nutrients, light,
and space in areas where G. remyi
occurs on these islands. Landslides are
a threat to occurrences and habitat of G.
remyi on Hawaii Island (Perry 2006, in
litt.). Lack of pollination was suggested
as the cause for abortion of immature
fruits that were seen among plants on
Hawaii Island (PEPP 2010, p. 73).
Similarly, Agorastos (2011, in litt.)
reported no viable seed production in
the wild or within ex situ collections
and no recruitment in the wild among
the 14 individuals observed on the
island of Hawaii, Maui, and Molokai, for
unknown reasons (Duvall 2015, in litt.;
Oppenheimer 2015, in litt.). Some
species of Gardenia are dioecious (male
and female flowers on separate plants)
and although the breeding system of G.
remyi is currently unknown, this may be
a cause of failure to produce viable seed
in isolated individuals (Lorence 2015, in
litt.). Predation of seeds by rats is
reported as a threat to individuals on
Kauai (NTBG 2008, in litt.). Fortini et al.
(2013, p. 76) found that, as
environmental conditions are altered by
climate change, G. remyi is unlikely to
tolerate or adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to G.
remyi described above.
The remaining occurrences of
Gardenia remyi are at risk. Gardenia
remyi continues to be negatively
affected by habitat modification and
destruction by ungulates, and by direct
competition from nonnative plants,
combined with herbivory by ungulates
and seed predation by rats. Natural
events such as landslides are a threat to
occurrences on the island of Hawaii.
Pollination and seed production are
observed to be limited. Low numbers of
individuals (90 total individuals
distributed across 4 islands) makes this
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species more vulnerable to extinction
because of the higher risks from genetic
bottlenecks, random demographic
fluctuations, and localized catastrophes.
The effects of climate change are likely
to exacerbate these threats. Because of
the threats, we find that this species is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Huperzia stemmermanniae (NCN) is
an epiphytic hanging fir-moss (a fern
ally) in the club moss family
(Lycopodiaceae) (Palmer 2003, pp. 257–
259). This species is epiphytic on rough
bark of living trees or fallen logs in
Metrosideros polymorpha-Acacia koa
forest on the island of Hawaii, from
3,200 to 3,800 ft (975 to 1,160 m), in the
montane wet ecosystem (Medeiros et al.
1996b, p. 93; Palmer 2003, pp. 257, 259;
TNCH 2007; HBMP 2010). There is little
information available on the historical
range of this species. Huperzia
stemmermanniae was first collected in
1981, from two occurrences totaling 10
individuals in Laupahoehoe NAR on the
island of Hawaii, and was mistakenly
identified as H. mannii (Medeiros et al.
1996b, p. 93; HBMP 2010). One
individual occurred in Kaapahu Valley
on east Maui, but has not been relocated
since 1995 (Perry 2006, in litt.; Welton
2008, in litt.; HBMP 2010; Conry 2012,
in litt.; Perry 2015, in litt.). In 2006,
there were estimated to be as many as
20 individuals in Laupahoehoe (Perry
2006, in litt.). Currently, there are only
a few individuals remaining due to
prolonged drought conditions (Perry
2015, in litt.).
Feral pigs, goats, axis deer, and cattle
modify and destroy the habitat of
Huperzia stemmermanniae on Maui,
and feral pigs modify and destroy the
habitat of this species on Hawaii Island
(Medeiros et al. 1996b, p. 96; Wood
2003, in litt.; HBMP 2010). Herbivory by
these ungulates is a threat to H.
stemmermanniae. Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Nonnative
plants, such as Clidemia hirta, Miconia
calvescens, Psidium cattleianum, and
Cyathea cooperi (Australian tree fern),
modify and destroy the forest habitat
that supports the native species upon
which this epiphytic plant grows, and
drought also negatively affects this
species and its habitat (Medeiros et al.
1996b, p. 96; Perry 2006, in litt.; HBMP
2010). Huperzia stemmermanniae
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experiences reduced reproductive vigor
due to reduced levels of genetic
variability, leading to diminished
capacity to adapt to environmental
changes, thereby lessening the
probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman
and Pilson 1997, p. 361; HBMP 2010).
Fortini et al. (2013, p. 77) found that, as
environmental conditions are altered by
climate change, H. stemmermanniae is
unlikely to tolerate or adapt to projected
changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to H. stemmermanniae
described above.
The remaining occurrences of
Huperzia stemmermanniae are at risk.
The known individuals are restricted to
a small area on Hawaii Island, and this
species continues to be negatively
affected by habitat modification and
destruction by ungulates. The low
numbers of individuals H.
stemmermanniae reduces the
probability of its long-term persistence.
The effects of climate change are likely
to further exacerbate these threats.
Because of the threats described above,
we find that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Hypolepis hawaiiensis var. mauiensis
(olua) is a small terrestrial member of
the bracken fern family
(Dennstaedtiaceae), and is recognized as
a distinct taxon by Palmer (2003, pp.
168–169). Hypolepis hawaiiensis var.
mauiensis occurs in wet forest,
predominately in the montane wet
ecosystem (Palmer 2003, pp. 168–170;
Oppenheimer 2015, in litt.). This variety
is historically known from west Maui
(Palmer 2003, pp. 168–170). Currently,
5 to 10 individuals are known from
openings between bogs on west Maui,
and a few individuals are known from
east Maui (Maui Nui Task Force (MNTF)
2010, in litt.).
Nonnative plants, such as Tibouchina
herbacea, modify and destroy the
habitat of Hypolepis hawaiiensis var.
mauiensis on east and west Maui
(HBMP 2010; MNTF 2010, in litt.).
Nonnative plants also displace this and
other native plant species by competing
for water, nutrients, light, and space, or
they may produce chemicals that inhibit
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stones 1990, p. 74; MNTF
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2010, in litt.). Herbivory by slugs is a
threat (Oppenheimer 2015, in litt.). This
fern experiences reduced reproductive
vigor due to low numbers of
individuals, leading to diminished
capacity to adapt to environmental
changes, and thereby lessening the
probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman
and Pilson 1997, p. 361). Fortini et al.
(2013, p. 78) found that, as
environmental conditions are altered by
climate change, H. hawaiiensis var.
mauiensis is unlikely to tolerate or
adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to H.
hawaiiensis var. mauiensis described
above.
The remaining occurrences of
Hypolepis hawaiiensis var. mauiensis
are at risk. Nonnative plants modify and
destroy native habitat, and also
outcompete native plants, and this plant
is threatened by herbivory by slugs. This
fern is also vulnerable to the impacts of
climate change, and the small number of
remaining individuals limits its ability
to adapt to environmental change.
Because of these threats, we find that
this variety is endangered throughout all
of its range, and, therefore, find that it
is unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Joinvillea ascendens ssp. ascendens
(ohe) is an erect perennial herb in the
joinvillea family (Joinvilleaceae)
(Wagner et al. 1999, p. 1450). Joinvillea
ascendens ssp. ascendens occurs in wet
to mesic Metrosideros polymorphaAcacia koa lowland and montane forest,
and along intermittent streams, from
1,000 to 4,300 ft (305 to 1,300 m); in the
lowland mesic (Kauai and Oahu),
lowland wet (Oahu, Molokai, Maui, and
Hawaii Island), montane wet (Kauai,
Oahu, Molokai, Maui, and Hawaii
Island), and montane mesic ecosystems
(Kauai) (TNCH 2007; HBMP 2010).
Historically, this subspecies was found
in widely distributed occurrences on the
islands of Kauai, Oahu, Molokai, Maui,
and Hawaii Island (HBMP 2010). On
Kauai, this subspecies was wide-ranging
across the mountains and into coastal
areas (HBMP 2010). On Oahu, this
subspecies was known from the summit
area of the Waianae Mountains, and
ranged along the entire length of the
Koolau Mountain range. On Molokai,
this subspecies was known from the
eastern half of the island ranging from
Pelekunu Preserve and east to Halawa
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Valley. On west Maui, this subspecies
occurred in the summit area, and on the
northeastern side of east Maui it ranged
from the Koolau FR to Kaapahu (Gates
2015, in litt.). On Hawaii Island, it
occurred almost island-wide. Currently,
J. ascendens ssp. ascendens is still
found on the same islands, in only 56
occurrences totaling approximately 200
individuals (HBMP 2010; Conry 2012,
in litt.).
Nonnative ungulates modify and
destroy habitat on all of the islands
where Joinvillea ascendens ssp.
ascendens occurs (Moses 2006, in litt.;
Oppenheimer 2006, in litt.; Welton and
Haus 2008, p. 16; HBMP 2010; Perlman
2010, in litt.). Herbivory by feral pigs,
goats, axis deer, black-tailed deer, and
rats is a threat to this subspecies (HBMP
2010; Williams 2015, in litt.). Ungulates
are managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Many
nonnative plant species, such as
Passiflora tarminiana (banana poka),
Rubus ellipticus (yellow Himalayan
raspberry), and Setaria palmifolia
(palmgrass) on Hawaii Island; Clidemia
hirta, Psidium cattleianum, and P.
guajava on Kauai; C. hirta and
Tibouchina herbacea on Maui; Juncus
effusus (Japanese mat rush) on Molokai;
and C. hirta and P. cattleianum on
Oahu, modify and destroy habitat and
outcompete this subspecies (HBMP
2010). Randomly occurring natural
events, such as landslides, are a threat
to the occurrences of J. ascendens ssp.
ascendens on Kauai and Molokai
(HBMP 2010). Fire is likely to be a
threat to this subspecies in the drier
areas of the Waianae Mountains of Oahu
(HBMP 2010). This subspecies is
usually found as widely separated
individuals. Seedlings have rarely been
observed in the wild, and, although
mature seeds germinate in cultivation,
these seedlings also rarely survive to
maturity. It is uncertain if this rarity of
reproduction is typical, or if it is related
to habitat disturbance, or possibly a lack
of soil mycorrhizae (symbiotic
relationship between fungi and plants)
required for successful establishment
(Wagner et al. 1999, p. 1451;
Oppenheimer 2015, in litt.). Fortini et
al. (2013, p. 76) found that, as
environmental conditions are altered by
climate change, J. ascendens ssp.
ascendens is unlikely to tolerate or
adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
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more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to J.
ascendens ssp. ascendens described
above.
The remaining occurrences of
Joinvillea ascendens ssp. ascendens are
at risk. The known individuals continue
to be negatively affected by habitat
modification and destruction by
ungulates, compounded with herbivory
by ungulates and rats. The small
number of remaining individuals,
smaller distribution, and poor
recruitment in the wild limits this
subspecies’ ability to adapt to
environmental changes. Destruction by
fire, landslides, rockfalls, and floods can
occur at any time. The effects of climate
change are likely to further exacerbate
these threats. Because of these threats,
we find that this subspecies is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Kadua fluviatilis (previously Hedyotis
fluviatilis) (kamapuaa) is a climbing
shrub in the coffee family (Rubiaceae)
family (Wagner et al. 1999, pp. 1142–
1144). Typical habitat for this species on
Kauai is mixed native shrubland and
Metrosideros forest from 750 to 2,200 ft
(230 to 680 m), in the lowland mesic
ecosystem (TNCH 2007; HBMP 2010);
and in open shrubland with sparse tree
cover in the lowland mesic ecosystem
(Wood 1998, in litt.; TNCH 2007). On
Oahu, K. fluviatilis occurs along rocky
streambanks in wet Metrosideros forest
from 820 to 1,990 ft (250 to 607 m) in
the lowland wet ecosystem (TNCH
2007; HBMP 2010).
Historically, Kadua fluviatilis was
known from the island of Kauai in at
least 5 occurrences ranging from the
north coast across the central plateau to
the south coast, and from the island of
Oahu in at least 11 occurrences in the
northern Koolau Mountains (HBMP
2010; Williams 2015, in litt.). Currently,
during surveys on Oahu in 2013, only
20 to 25 individuals were observed in
one occurrence (Wood 2005b, in litt.,
NTBG 2009, in litt.; HBMP 2010; Ching
Harbin 2015, in litt.). On Kauai, K.
fluviatilis is known from two
occurrences totaling approximately 500
individuals (HBMP 2010).
Feral pigs and goats modify and
destroy habitat of Kadua fluviatilis
(HBMP 2010). Evidence of the activities
of feral pigs has been reported at the
occurrences on Kauai and Oahu (Wood
1998, in litt.; HBMP 2010). Feral goats
and evidence of their activities have
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been observed at one location on Kauai
(HBMP 2010). Herbivory by feral pigs
and goats is a threat to K. fluviatilis.
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plant species, such as
Lantana camara, Paspalum conjugatum
(Hilo grass), Psidium cattleianum, P.
guajava, Rubus rosifolius, and Schinus
terebinthifolius (Christmas berry),
degrade habitat and outcompete this
and other native species for water,
nutrients, light, and space, or may
produce chemicals that inhibit growth
of other plants (Smith 1985, pp. 180–
250; Vitousek et al. 1987 in Cuddihy
and Stone 1990, p. 74; Wood 1998, in
litt.; HBMP 2010). Kadua fluviatilis is
negatively affected by landslides on
Kauai (HBMP 2010). Fortini et al. (2013,
p. 78) found that, as environmental
conditions are altered by climate
change, K. fluviatilis is unlikely to
tolerate or adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to K.
fluviatilis described above.
The remaining occurrences of Kadua
fluviatilis are at risk. Numbers of
occurrences and individuals are
decreasing on Oahu and Kauai, from 16
occurrences to 3, and from over 1,000
individuals to about 500 individuals
(HBMP 2010; OTFM 2014, in litt.). This
species continues to be negatively
affected by habitat modification and
destruction by feral pigs and goats,
stochastic events such as landslides,
and direct competition from nonnative
plants, combined with herbivory by
nonnative ungulates. Climate change is
likely to further exacerbate these threats.
Because of these threats, we find that
this species is endangered throughout
all of its range, and, therefore, find that
it is unnecessary to analyze whether it
is endangered or threatened in a
significant portion of its range.
Kadua haupuensis (NCN) is a shrub
in the coffee family (Rubiaceae)
(Lorence et al. 2010, p. 137). There is no
historical information for this species as
it was recently discovered and
described from one occurrence just
below and along cliffs in an isolated
area on southern Kauai, from 980 to
1,640 ft (300 to 500 m), in the lowland
mesic ecosystem (TNCH 2007; Lorence
et al. 2010, pp. 137–144). Currently,
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however, there are no known extant
individuals of K. haupuensis; the single
natural occurrence is thought to be
extirpated. Ten individuals were
propagated from seed collected in 1999,
with cuttings from these currently under
cultivation. Seeds are in storage at
NTBG’s seed bank (Lorence 2015, in
litt.).
Feral pigs modify and destroy the
habitat of Kadua haupuensis on Kauai
(Lorence et al. 2010, p. 140). Ungulates
are managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Predation of
fruits and seeds by rats is a threat.
Landslides are an additional threat to
this species at its last known
occurrence. Nonnative plants, such as
Caesalpinia decapetala (wait-a-bit),
Passiflora laurifolia (yellow granadilla),
and various nonnative grasses, modify
and destroy native habitat, outcompete
native plants, and are found at the last
known location of K. haupuensis. The
small number of remaining individuals
limits this species’ ability to adapt to
environmental change. Because of these
threats, we find that K. haupuensis is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Labordia lorenciana (NCN) is a small
tree in the Logania family (Loganiaceae)
(Wood et al. 2007, pp. 195–197). This
species occurs on the island of Kauai at
3,800 ft (1,160 m), in forest in the
montane mesic ecosystem (Wood et al.
2007, pp. 197–198). Currently, there are
four known individuals. Additional
surveys for L. lorenciana have not been
successful; however, experts believe this
species may occur in other areas (Wood
et al. 2007, p. 198).
Labordia lorenciana is at risk from
habitat modification and destruction
and herbivory by nonnative mammals,
displacement of individuals through
competition with nonnative plants,
stochastic events, and problems
associated with small populations. Feral
pigs, goats, and black-tailed deer modify
and destroy the habitat of L. lorenciana
(Wood et al. 2007, p. 198; Kishida 2015,
in litt.). Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction by
these animals (Anderson et al. 2007, in
litt; HAR–DLNR 2010, in litt.). Predation
of seeds by rats is a threat to this species
(Wood et al. 2007, p. 198). Habitat
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destruction and modification by
nonnative plants, and competition with
nonnative plants including Lantana
camara, Passiflora tarminiana, Psidium
cattleianum, and Rubus argutus, are a
threat to Labordia lorenciana, as these
nonnative plants have the ability to
spread rapidly and cover large areas in
the forest understory (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; Wood
et al. 2007, p. 198). Randomly occurring
natural events, such as landslides, flash
floods, fallen tree limbs, and fire, are
threats to L. lorenciana where it occurs
on Kauai (Wood et al. 2007, p. 198).
This species experiences reduced
reproductive vigor as there is no in situ
seedling recruitment and a very small
number of individuals remain (Wood et
al. 2007, p. 198). Infestation by the black
twig borer (Xylosandrus compactus) is a
threat to this species (Kishida 2015, in
litt.). Because of these threats, we find
that L. lorenciana is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Lepidium orbiculare (anaunau) is a
small, many-branched shrub in the
mustard family (Brassicaceae) (St. John
1981, pp. 371–373; Wagner et al. 1999,
p. 409). This species occurs in mesic
forest on the island of Kauai, in the
lowland mesic ecosystem (Wagner et al.
1999, p. 409; TNCH 2007; HBMP 2010;
PEPP 2014, p. 34). Historically, this
species was known from widely
scattered occurrences on Kauai (Wagner
et al. 1999, p. 409). Currently, there is
one occurrence of fewer than 50
individuals (Wagner et al. 2012, p. 19;
PEPP 2014, p. 34; Smithsonian
Institution 2015, in litt.).
Feral pigs and goats have been
documented to modify and destroy
habitat of other rare and endangered
native plant species at the same location
on Kauai (Lorence et al. 2010, p. 140;
Kishida 2015, in litt.); therefore, we
consider that activities of feral pigs and
goats also pose a threat to Lepidium
orbiculare. Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as Melinis
minutiflora (molasses grass) and
Stachytarpheta jamaicensis (Jamaica
vervain), degrade native habitat,
outcompete native plants, and are found
at the last known location of L.
orbiculare (HBMP 2010). Landslides are
an additional threat to this species.
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Because there are fewer than 50
individuals, L. orbiculare experiences
reduced reproductive vigor due to
reduced levels of genetic variability,
leading to diminished capacity to adapt
to environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361;
PEPP 2014, p. 34).
The remaining occurrence of
Lepidium orbiculare is at risk and the
species continues to be negatively
affected by the threats described above.
Because of these threats, we find that
this species is endangered throughout
all of its range, and, therefore, find that
it is unnecessary to analyze whether it
is endangered or threatened in a
significant portion of its range.
Microlepia strigosa var. mauiensis
(NCN) is a terrestrial, medium-sized fern
in the bracken fern family
(Dennstaedtiaceae) (Palmer 2003, p.
186). Typical habitat for M. strigosa var.
mauiensis is mesic to wet forest from
1,400 to 6,000 ft (425 to 1,830 m), in the
lowland mesic (Oahu), montane mesic
(Hawaii Island), and montane wet (Maui
and Hawaii Island) ecosystems (Palmer
2003, p. 186; TNCH 2007: HBMP 2010).
Little is known of the historical
locations of M. strigosa var. mauiensis;
however, it was wide-ranging on the
islands of Hawaii, Maui, and Oahu
(HBMP 2010). Currently, M. strigosa var.
mauiensis is known from nine
occurrences totaling fewer than 100
individuals on the islands of Oahu
(about 40 individuals), Maui (fewer than
20 individuals on east and west Maui),
and Hawaii (35 individuals last
observed in 2004) (Palmer 2003, p. 186;
Lau 2007, pers.comm.; Oppenheimer
2007 and 2008, in litt.; Welton 2008, in
litt.; Ching 2011, in litt.; Ching Harbin
2015, in litt.; Oppenheimer 2015, in
litt.).
Habitat modification and destruction
by feral pigs and goats is a threat to
Microlepia strigosa var. mauiensis
(Oppenheimer 2007, in litt.; Bily 2009,
in litt.; HBMP 2010). Herbivory by feral
pigs is a threat to M. strigosa var.
mauiensis (Oppenheimer 2007, in litt.;
Bily 2009, in litt.; HBMP 2010).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Ageratina adenophora, Juncus
acuminatus (rush), Plantago major
(broad-leaved plantain), and Tibouchina
herbacea, degrade habitat and
outcompete this variety on Maui
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(Oppenheimer 2007, in litt.).
Hybridization with other species and
varieties of Microlepia is a threat to this
plant on Oahu and is compounded by
the low number of individuals (Kawelo
2010, in litt.). Because of these threats,
we find that this variety is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Myrsine fosbergii (kolea) is a branched
shrub or small tree in the myrsine
family (Myrsinaceae) (Wagner et al.
1999, p. 940). Typical habitat for this
species on Oahu is Metrosideros-mixed
native shrubland, from 2,200 to 2,800 ft
(670 to 850 m) (Wagner et al. 1999, p.
940; TNCH 2007; HBMP 2010). Typical
habitat for this species on Kauai is
Metrosideros-Diospyros (ohia-lama)
lowland mesic forest and MetrosiderosCheirodendron (ohia-olapa) montane
wet forest, often on watercourses or
stream banks, from 900 to 4,300 ft (270
to 1,300 m), in the lowland mesic,
lowland wet, and montane wet
ecosystems (TNCH 2007; HBMP 2010;
Wagner et al. 2012, p. 53). Myrsine
fosbergii was historically known from
the summit ridges of the Koolau
Mountains of Oahu (HBMP 2010). This
species was first collected on Kauai in
1987. Currently, on Oahu, there are
fewer than 30 individuals in the Koolau
Mountains (lowland mesic and lowland
wet ecosystems) (HBMP 2010; OTFM
2014, in litt.; Reynolds 2015, in litt.;
Sailer 2015, in litt.). Propagation
attempts of the Oahu plants have been
unsuccessful (Ching Harbin 2015, in
litt.). On Kauai, this species was once
widely scattered in the northwest and
central areas, but is currently known
from only 55 remaining individuals
(Wood 2005e and 2007c, in litt.; HBMP
2010).
Myrsine fosbergii is at risk from
habitat modification and destruction by
nonnative ungulates and plants. On
Oahu, evidence of the activities of feral
pigs has been reported at all summit
occurrences (HBMP 2010). On Kauai,
evidence of the activities of feral pigs
has been reported at the remaining
occurrence (Wood 2005e and 2007c, in
litt.; HBMP 2010), and evidence of the
activities of feral goats has also been
reported (HBMP 2010). Herbivory by
feral pigs and goats is a threat to M.
fosbergii (Wood 2005e and 2007c, in
litt.; HBMP 2010). Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
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HAR–DLNR 2010, in litt.). Nonnative
plants, such as Axonopus fissifolius
(narrow-leaved carpetgrass), Clidemia
hirta, Erigeron karvinskianus, Psidium
cattleianum, P. guajava, and Rubus
rosifolius, compete with M. fosbergii
and modify and destroy its native
habitat on Oahu and Kauai (HBMP
2010). Hybridization is a threat to this
species, as M. fosbergii hybridizes with
other Myrsine species, and the number
of non-hybrid individuals may actually
be lower than estimated (Ching Harbin
2015, in litt.). Fortini et al. (2013, p. 82)
found that, as environmental conditions
are altered by climate change, M.
fosbergii is unlikely to tolerate or adapt
to projected changes in temperature and
moisture, and is unlikely to be able to
move to areas with more suitable
climatic conditions. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to M. fosbergii
described above.
The remaining occurrences of Myrsine
fosbergii are at risk from the threats
described above. The effects of climate
change are likely to exacerbate the
threats described above. Because of
these threats, we find that M. fosbergii
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Nothocestrum latifolium (aiea) is a
small tree in the nightshade family
(Solanaceae) (Symon 1999, p. 1263).
Typical habitat for this species is dry to
mesic forest in the dry cliff (Kauai,
Oahu, Lanai, and Maui), lowland dry
(Oahu, Lanai, and Maui), and lowland
mesic (Oahu, Molokai, Lanai, and Maui)
ecosystems (TNCH 2007; HBMP 2010).
Historically, N. latifolium was known
from the Waianae Mountains of Oahu,
Molokai, Lanai, and Maui (HBMP 2010;
Sailer 2015, in litt.). This species was
collected once on Kauai in 1986, but has
not been observed there before or after
that time (Symon 1999, p. 1263; BISH
504035-Montgomery; Williams 2015, in
litt.). Currently, on the island of Oahu,
there is one individual remaining, with
only one of the other previously extant
individuals represented in an ex situ
collection (Moses 2006, in litt.; Starr
2006, in litt.; HBMP 2010; Kawakami
2010, in litt.; Kawelo 2010, in litt.;
Welton 2010, in litt.; Ching 2011, in litt.;
Ching Harbin 2015, in litt.; Sailer 2015,
in litt.). On Molokai, there a few
individuals on the central south slope
(Oppenheimer 2015, in litt.). There are
18 occurrences totaling approximately
1,600 individuals on east and west Maui
(Ching 2011, in litt.; HBMP 2010;
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Oppenheimer 2015, in litt.). On Lanai,
no individuals were found during
surveys in 2012, and this species may be
extirpated from this island, although
there are plans to continue surveying
suitable habitat (PEPP 2012, p. 129;
Oppenheimer 2015, in litt.). In
summary, the species’ range on each
island has decreased dramatically since
2001 (Kawelo 2005 and 2010, in litt.;
HBMP 2010; Oppenheimer 2011, in
litt.).
Feral pigs (Oahu, Maui, Kauai), goats
(Maui, Kauai), mouflon (Lanai), feral
cattle (Maui), axis deer (Lanai, Maui),
and black-tailed deer (Kauai) modify
and destroy habitat of Nothocestrum
latifolium (HBMP 2010; Oppenheimer
2015, in litt.). Herbivory by these
animals also poses a threat to this
species. Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as Fraxinus
uhdei (tropical ash), Grevillea robusta
(silk oak), Lantana camara, Leucaena
leucocephala (koa haole), Melinis
minutiflora, Passiflora suberosa
(huehue haole), Schinus
terebinthifolius, and Toona ciliata
(Australian red cedar), outcompete N.
latifolium and modify and destroy
habitat at all known occurrences.
Wildfire, and fire caused by military
training activities, is a threat to this
species and its habitat (Sailer 2015, in
litt.). Low numbers of individuals limits
this species’ ability to adapt to
environmental change. Infestation by
the black twig borer is a threat to N.
latifolium (Ching Harbin 2015, in litt.).
This species continues to decline, and,
for unknown reasons, there is an
observed lack of regeneration in N.
latifolium in the wild (HBMP 2010;
Duvall 2015, in litt.). Fortini et al. (2013,
p. 83) found that, as environmental
conditions are altered by climate
change, N. latifolium is unlikely to
tolerate or adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to N.
latifolium described above.
The remaining occurrences of
Nothocestrum latifolium are at risk from
the threats described above. Because of
these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
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67801
endangered or threatened in a
significant portion of its range.
Ochrosia haleakalae (holei) is a tree
in the dogbane family (Apocynaceae)
(Wagner et al. 1999, p. 218). Typical
habitat for this species is dry to mesic
forest, sometimes wet forest, and often
lava, from 2,300 to 4,000 ft (700 to 1,200
m), in the dry cliff (Maui), lowland
mesic (Maui and Hawaii Island), and
montane mesic (Maui) ecosystems
(Medeiros et al. 1986, pp. 27–28;
Wagner et al. 1999, p. 218; TNCH 2007;
HBMP 2010). Historically, this species
was known from east Maui and Hawaii
Island (HBMP 2010). Currently, O.
haleakalae is known from 4 occurrences
totaling about 15 individuals on the
island of Maui (Medeiros 2007, in litt.;
Oppenheimer 2008, in litt.; HBMP 2010;
Oppenheimer 2015, in litt.). On Hawaii
Island, there are two occurrences
totaling at least 150 individuals in
Hawaii Volcanoes National Park, with
150 outplanted in nearby kipuka
(vegetated areas surrounded by lava
flows), and one individual in the
Laupahoehoe section of Hilo Forest
Reserve (Pratt 2005, in litt.; Bio 2008a,
in litt.; HBMP 2010; Pratt 2011, in litt.;
Conry 2012, in litt.; Orlando 2015, in
litt.; Perry 2015, in litt.).
Feral pigs and goats modify and
destroy the habitat of Ochrosia
haleakalae on Maui and Hawaii Island;
in addition, cattle modify and destroy
the habitat of this species on Maui
(Medeiros 1995, in litt.; Pratt 2005, in
litt.; Oppenheimer 2015, in litt.).
Herbivory by these animals also poses a
threat to this species. Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Nonnative
plant species, such as Cestrum diurnum
(day cestrum), Fraxinus uhdei, Psidium
cattleianum, P. guajava, Rubus argutus,
Setaria palmifolia (palmgrass), and
Toona ciliata, modify and destroy
habitat and outcompete native plants,
including O. haleakalae (HBMP 2010).
In dry areas, wildfires affecting the
habitat of this species are exacerbated
by the presence of introduced grass
species such as Pennisetum
clandestinum (kikuyu grass) (HBMP
2010; Oppenheimer 2015, in litt.).
Herbivory and seed predation by slugs
and rats is a threat to this species
(Oppenheimer 2015, in litt.). There is
low to no reproduction observed in the
wild, and this reduced reproductive
vigor is due to reduced levels of genetic
variability resulting from low numbers
of individuals. This decreases the
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species’ capacity to adapt to
environmental changes, and thereby
lessens the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361;
Duvall 2015, in litt.). Fortini et al. (2013,
p. 83) found that, as environmental
conditions are altered by climate
change, O. haleakalae is unlikely to
tolerate or adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to O.
haleakalae described above.
Ochrosia haleakalae is at risk from
habitat degradation and loss by feral
pigs, goats, cattle, and nonnative plants;
the displacement of individuals due to
competition with nonnative plants for
space, nutrients, water, air, and light;
herbivory by feral pigs, goats, and cattle;
seed predation by slugs and rats; and by
the small number of remaining
individuals. The effects of climate
change are likely to further exacerbate
these threats. Because of these threats,
we find that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Phyllostegia brevidens (NCN) is a
scandent (climbing) subshrub in the
mint family (Lamiaceae) (Wagner et al.
1999, pp. 814–815). This species occurs
in wet forest on the islands of Maui and
Hawaii from 2,900 to 3,200 ft (880 to
975 m), in the lowland wet (Maui),
montane wet (Hawaii Island), and wet
cliff (Maui) ecosystems (Wagner et al.
1999, pp. 814–815; TNCH 2007; HBMP
2010). Phyllostegia brevidens is
historically known from Hilo Forest
Reserve, Mauna Kea, and Kulani on
Hawaii Island; and from Kipahulu
Valley on Maui (Haleakala National
Park) (Wagner et al. 1999, p. 815; HBMP
2010; Smithsonian Institution 2015, in
litt.). Currently, there is one individual
on the island of Maui and two
individuals on Hawaii Island (PEPP
2009, p. 90; Wagner et al. 2012, p. 46;
PEPP 2014, p. 136; Gates 2015, in litt.;
Oppenheimer 2015, in litt.; Perry 2015,
in litt.).
Feral pigs modify and destroy habitat
of this species on Maui (PEPP 2014, p.
136). The two remaining individuals on
Hawaii Island are currently fenced
(Perry 2015, in litt.); however, owing to
the potential for accidental damage or
vandalism (irrespective of
maintenance), fences do not guarantee
protection from ungulate ingress.
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Herbivory by feral pigs also poses a
threat to this species on Maui.
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Clidemia hirta and Hedychium
gardnerianum, modify and destroy
habitat and outcompete P. brevidens on
Maui (PEPP 2009, p. 90). Herbivory by
slugs is a threat to the remaining
individual on Maui (PEPP 2014, p. 136).
In addition, natural events such as
landslides and erosion are threats to the
occurrence on Maui (PEPP 2014, p.
136). The small number of remaining
individuals limits this species’ ability to
adapt to environmental change. Fortini
et al. (2013, p. 84) found that, as
environmental conditions are altered by
climate change, P. brevidens is unlikely
to tolerate or adapt to projected changes
in temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to P.
brevidens described above.
The remaining occurrences of
Phyllostegia brevidens are at risk. The
species continues to be negatively
affected by habitat modification and
destruction by ungulates and nonnative
plants, and by direct competition from
nonnative plants, combined with
herbivory by ungulates and slugs. The
effects of climate change are likely to
further exacerbate these threats. We find
that P. brevidens is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Phyllostegia helleri (NCN) is a weakly
erect to climbing shrub in the mint
family (Lamiaceae) (Wagner et al. 1999,
pp. 816–817). This species occurs on
ridges or spurs from 2,800 to 4,000 ft
(860 to 1,200 m) in diverse forest on
Kauai in the lowland wet, montane wet,
and wet cliff ecosystems (Wagner et al.
1999, p. 817; TNCH 2007; HBMP 2010).
Historically, P. helleri was wide-ranging
on the island of Kauai, from the north
and east sides throughout the central
plateau (Wagner et al. 1999, p. 817;
HBMP 2010). Currently, this species is
limited to one occurrence of four
individuals (PEPP 2014, p. 35; Kishida
2015, in litt.).
Feral pigs and goats modify and
destroy the habitat of Phyllostegia
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helleri on Kauai (HBMP 2010).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Herbivory on fruits and seeds by
rats negatively affects the remaining
individuals (HBMP 2010). The only
known occurrence of this species is
located at the base of cliffs, and
landslides are an additional threat
(HBMP 2010). Nonnative plants, such as
Erigeron karvinskianus, Kalanchoe
pinnata (air plant), Psidium guajava,
Rubus rosifolius, and various grasses,
modify and destroy native habitat,
outcompete native plants, and are found
at the last known occurrence of
Phyllostegia helleri (HBMP 2010). This
species experiences reduced
reproductive vigor due to reduced levels
of genetic variability, leading to
diminished capacity to adapt to
environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 84) found that, as
environmental conditions are altered by
climate change, P. helleri is unlikely to
tolerate or adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to P.
helleri described above.
The remaining occurrence of
Phyllostegia helleri is at risk. The
numbers of individuals are decreasing
on Kauai, as this species was wideranging on the island, extending from
the north and east sides throughout the
central plateau, and is now known from
only one occurrence of four individuals.
These four individuals continue to be
negatively affected by habitat
modification and destruction by
ungulates and nonnative plants, direct
competition by nonnative plants, and by
seed predation by rats. Natural events
such as landslides may damage or
destroy the remaining four individuals.
The small number of remaining
individuals limits this species’ ability to
adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that P. helleri
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
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endangered or threatened in a
significant portion of its range.
Phyllostegia stachyoides (NCN) is a
weakly erect to climbing subshrub in
the mint family (Lamiaceae) (Wagner et
al. 1999, p. 823). This species occurs in
mesic to wet forest from 3,600 to 4,600
ft (1,000 to 1,400 m), in the montane wet
(Hawaii Island, Maui, and Molokai) and
montane mesic (Hawaii Island and
Maui) ecosystems (Wagner et al. 1999,
p. 823; TNCH 2007; HBMP 2010).
Phyllostegia stachyoides is historically
known from the eastern and central
Molokai, west Maui, and wide-ranging
occurrences on Hawaii Island (Wagner
et al. 1999, p. 823; HBMP 2010;
VanDeMark 2016, in litt.). Currently,
occurrences on west Maui total about 15
individuals (Oppenheimer 2015, in
litt.). Those on Molokai occur at 5
locations and total fewer than 30
individuals (Orlando 2015, in litt.; PEPP
2012, p. 156). Plants on Hawaii Island
are now considered to be P. ambigua
(VanDeMark 2016, in litt.).
Feral pigs, goats, and axis deer modify
and destroy the habitat of Phyllostegia
stachyoides on Maui, with evidence of
the activities of these animals reported
in areas where this species occurs
(HBMP 2010; PEPP 2014, p. 141).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Ageratina adenophora, Erigeron
karvinskianus, and Tibouchina
herbacea, compete with P. stachyoides,
modify and destroy its native habitat,
and displace other native plant species
(Smith 1985, pp. 180–250; Vitousek et
al. 1987 in Cuddihy and Stone 1990, p.
74; PEPP 2014, pp. 141–142). Herbivory
by slugs and rats on leaves and nutlets
of P. stachyoides poses a threat to this
species at known locations on Maui and
Molokai (PEPP 2014, pp. 140–142). On
Maui, stochastic events such as floods
and drought (with ensuing erosion) pose
a threat to small, isolated occurrences of
P. stachyoides; rockfalls and landslides
are a threat to occurrences on Molokai
(PEPP 2014, pp. 140–142). This species
experiences reduced reproductive vigor
due to reduced levels of genetic
variability, leading to diminished
capacity to adapt to environmental
changes, and thereby lessening the
probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman
and Pilson 1997, p. 361). Fortini et al.
(2013, p. 84) found that, as
environmental conditions are altered by
climate change, P. stachyoides is
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unlikely to tolerate or adapt to projected
changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to P. stachyoides described
above.
The remaining occurrences of
Phyllostegia stachyoides are at risk. The
known individuals are restricted to
small areas on west Maui and Molokai,
and continue to be negatively affected
by habitat modification and destruction
by ungulates and by direct competition
with nonnative plants, combined with
herbivory by slugs and rats. The small
number of remaining individuals limits
this species’ ability to adapt to
environmental changes. Flooding,
drought, and the effects of climate
change are likely to further exacerbate
these threats. Because of these threats,
we find that this species is endangered
throughout all of its range, and,
therefore, find that it is unnecessary to
analyze whether it is endangered or
threatened in a significant portion of its
range.
Portulaca villosa (ihi) is a perennial
herb in the purslane family
(Portulacaceae) (Wagner et al. 1999, p.
1074). Portulaca villosa occurs on dry,
rocky, clay, lava, or coralline reef sites,
from sea level to 1,600 ft (490 m), in the
coastal (Lehua, Kaula, Oahu,
Kahoolawe, Maui, and Hawaii Island)
and lowland dry (Oahu, Molokai, Lanai,
Kahoolawe, Maui, and Hawaii Island)
ecosystems, and one reported
occurrence in the montane dry (Hawaii
Island) ecosystem (Wagner et al. 1999,
p. 1074; TNCH 2007; HBMP 2010).
Portulaca villosa is historically known
from all the main Hawaiian Islands
except Niihau and Kauai (Wagner et al.
1999, p. 1074). Portulaca villosa has
been observed on the small islets of
Kaula and Lehua (west of Kauai and
Niihau), and on Nihoa (NWHI);
however, the current status of these
occurrences is unknown. This species
has not been observed on Oahu since
the 1960s, when it was locally abundant
at Kaohikaipu Island (HBMP 2010).
Historically, on the island of Hawaii,
this species occurred in the coastal area
of Hawaii Volcanoes National Park west
of Kamoamoa, but was extirpated in
1993 by lava flows (Orlando 2015, in
litt.). On the island of Lanai, two
individuals were last observed in 1996
(HBMP 2010). Currently, P. villosa is
known from a few individuals on
Molokai, 2 individuals on east Maui and
24 individuals on west Maui, fewer than
15 individuals on Kahoolawe, and five
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occurrences totaling 10 individuals on
Hawaii Island (MNTF 2010, in litt.;
Evans 2015a, in litt.).
Axis deer (Maui and Lanai), goats
(Maui), mouflon (Lanai), and cattle
(Hawaii Island) modify and destroy the
habitat of Portulaca villosa (HBMP
2010; Oppenheimer 2015, in litt.). These
ungulates also forage directly on this
species. Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as Lantana
camara, Nicotiana glauca (tree tobacco),
Pennisetum ciliare (buffelgrass), and
Prosopis pallida (kiawe, mesquite),
compete with Portulaca villosa, modify
and destroy its native habitat, displace
other native plant species, and pose a
threat to the known occurrences on
Hawaii Island, Maui, Kahoolawe, and
Molokai (Smith 1985, pp. 180–250;
Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74). P. villosa occurs in
drier coastal and lowland habitats, all of
which are affected by wildfires. Some
coastal habitat includes exposed cliffs,
which erode and cause landslides and
rockfalls in areas where P. villosa occurs
(Kahoolawe), posing a threat to this
species (HBMP 2010). This species
experiences reduced reproductive vigor
due to low levels of genetic variability,
leading to diminished capacity to adapt
to environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 86) found that, as
environmental conditions are altered by
climate change, P. villosa is unlikely to
tolerate or adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to P.
villosa described above.
The remaining occurrences of
Portulaca villosa are at risk; the
numbers of individuals are decreasing
on Maui, Molokai, and Hawaii Island,
and the species continues to be
negatively affected by continued habitat
modification and destruction by feral
ungulates and nonnative plants, and by
competition with nonnative plants.
Because of its small and isolated
remaining occurrences, natural events
such as rockfalls, landslides, and
wildfires may pose a threat to this
species. The small number of remaining
individuals limits this species’ ability to
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adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Pritchardia bakeri (Baker’s loulu) is a
small to medium-sized tree in the palm
family (Arecaceae) (Hodel 2009, pp.
173–179; Hodel 2012, pp. 70–73). This
species occurs in the lowland mesic
ecosystem in the Koolau Mountains on
Oahu, from 1,500 to 2,100 ft (457 to 640
m), in disturbed, windswept, and
mostly exposed shrubby or grassy areas,
and sometimes on steep slopes in these
areas (Bacon et al. 2012, pp. 1–17;
Hodel 2012, pp. 71–73). Currently,
occurrences total fewer than 100
individuals (Ching Harbin 2015, in litt.).
Habitat modification and destruction
by feral pigs impact the range and
abundance of Pritchardia bakeri.
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants compete with
and degrade and destroy native habitat
of P. bakeri, and displace native plant
species by competing for water,
nutrients, light, and space, or they may
produce chemicals that inhibit growth
of other plants (Smith 1985, pp. 180–
250; Vitousek et al. 1987 in Cuddihy
and Stone 1990, p. 74). Stochastic
events such as hurricanes modify and
destroy the habitat of P. bakeri, and can
directly damage or kill plants. Rats eat
the fruit before they mature, leading to
minimal or no recruitment (Hodel 2012,
pp. 42, 73). This species experiences
reduced reproductive vigor due to low
levels of genetic variability caused by
seed predation by rats and widely
separated occurrences, leading to
diminished capacity to adapt to
environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361;
Hodel 2012, p. 73).
The remaining occurrences of
Pritchardia bakeri are at risk; the known
individuals are restricted to small areas
on Oahu, and continue to be negatively
affected by habitat degradation and loss
by feral pigs and nonnative plants, fruit
predation by rats, and the small number
and reduced range of remaining
individuals. Although we cannot
predict the timing, extent, or magnitude
of specific impacts, we do expect the
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effects of climate change to exacerbate
the threats to P. bakeri described above.
Based on these threats, we find that this
species is endangered throughout all of
its range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Pseudognaphalium sandwicensium
var. molokaiense (enaena) is a perennial
herb in the sunflower family
(Asteraceae) (Wagner et al. 1999, p.
321). Typical habitat for this variety is
strand vegetation in dry consolidated
dunes, in the coastal ecosystem (Wagner
et al. 1999, p. 321; TNCH 2007; HBMP
2010). Historically, this variety was
known from Molokai, Oahu, Maui, and
Lanai (HBMP 2010; MNTF 2010, in
litt.). Currently, P. sandwicensium var.
molokaiense is known only from two
locations on Molokai (as many as 20,000
individuals, depending on rainfall), and
from fewer than 25 individuals on the
northwest coast of Maui (Moses 2006, in
litt.; Starr 2006, in litt.; Kallstrom 2008,
in litt.; Oppenheimer 2015, in litt.). This
variety was last observed on Lanai in
1960, and on Oahu (5 individuals) in
the 1980s (HBMP 2010).
Goats and axis deer modify and
destroy the habitat of
Pseudognaphalium sandwicensium var.
molokaiense, with evidence of the
activities of these animals reported in
the areas where this plant occurs (Moses
2006, in litt.; Starr 2006, in litt.;
Kallstrom 2008, in litt; HBMP 2010).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Additionally, nonnative plants,
such as Atriplex semibaccata
(Australian saltbush), Chenopodium
murale (aheahea, goosefoot),
Pennisetum ciliare, Prosopis pallida,
and Setaria parviflora (foxtail), compete
with and displace native plant species
by competing for water, nutrients, light,
and space, or they may produce
chemicals that inhibit growth of other
plants (Smith 1985, pp. 180–250;
Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74; Moses 2006, in litt.).
This variety experiences reduced
reproductive vigor due to low levels of
genetic variability, leading to
diminished capacity to adapt to
environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Rockfalls and landslides are a threat to
the occurrence of this variety on a sea
cliff on west Maui (HBMP 2010). Fortini
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et al. (2013, p. 86) found that, as
environmental conditions are altered by
climate change, P. sandwicensium var.
molokaiense is unlikely to tolerate or
adapt to projected changes in
temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to P.
sandwicensium var. molokaiense
described above.
The remaining occurrences of
Pseudognaphalium sandwicensium var.
molokaiense on Molokai and Maui are
at risk; individuals no longer occur on
Oahu and Lanai. Occurrences on Maui
and Molokai continue to be negatively
affected by habitat modification and
destruction by ungulates, and by direct
competition with nonnative plants. The
small number of remaining occurrences
limits this plant’s ability to adapt to
environmental changes. The effects of
climate change are likely to further
exacerbate these threats. Because of
these threats, we find that this variety is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Ranunculus hawaiensis (makou) is an
erect or ascending perennial herb in the
buttercup family (Ranunculaceae)
(Duncan 1999, p. 1088). Typical habitat
is mesic forest on grassy slopes and
scree, and in open pastures, from 6,000
to 6,700 ft (1,800 to 2,000 m), in the
montane mesic (Hawaii Island),
montane dry (Hawaii Island), and
subalpine (Hawaii Island and Maui)
ecosystems (Bio 2008a, in litt; Pratt
2007, in litt.; Duncan 1999, p. 1088;
TNCH 2007; HBMP 2010). Historically,
R. hawaiensis was wide-ranging on the
island of Hawaii. On Maui, this species
was known from Haleakala National
Park (HBMP 2010). In the 1980s and
1990s, this species numbered several
hundred individuals on both islands.
Currently, there are six occurrences
totaling 14 individuals on Hawaii Island
(Bio 2008a, in litt.; PEPP 2008, p. 108;
Pratt 2008, in litt.; HBMP 2010;
Agorastos 2011, in litt.; Imoto 2013, in
litt.; Orlando 2015, in litt.). On Maui, a
few individuals were observed on a cliff
in 1994; however, this occurrence was
not relocated in further surveys (PEPP
2013, p. 177). Additionally, no
individuals were re-observed in
Haleakala National Park (DLNR 2006, p.
61).
Feral pigs, mouflon, and cattle modify
and destroy the habitat of Ranunculus
hawaiensis on Hawaii Island, with
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evidence of the activities of these
animals reported in the areas where this
species occurs (HBMP 2010). These
ungulates also forage on R. hawaiensis.
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as Ehrharta
stipoides (meadow ricegrass), Holcus
lanatus (common velvetgrass), and
various grasses, modify and destroy
native habitat, outcompete native
plants, and have been reported in areas
where R. hawaiensis occurs (HBMP
2010). Drought and erosion pose a threat
in the areas of the last known
occurrences of R. hawaiensis on Maui
(PEPP 2013, p. 177). This species
experiences reduced reproductive vigor
due to low levels of genetic variability,
leading to diminished capacity to adapt
to environmental changes, and thereby
lessening the probability of long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 86) found that, as
environmental conditions are altered by
climate change, R. hawaiensis is
unlikely to tolerate or adapt to projected
changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to R. hawaiensis described
above.
The remaining occurrences of
Ranunculus hawaiensis are at risk; the
known individuals are restricted to
small areas on Hawaii Island and
continue to be negatively affected by
habitat modification and destruction by
feral ungulates, and by direct
competition with nonnative plants,
combined with predation by ungulates.
Drought and erosion pose a threat in the
areas of the last known occurrences on
Maui. The small number of remaining
individuals limits this species’ ability to
adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Ranunculus mauiensis (makou) is an
erect to weakly ascending perennial
herb in the buttercup family
(Ranunculaceae) (Duncan 1999, p.
1089). Typical habitat for R. mauiensis
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is open sites in mesic to wet forest and
along streams, from 3,500 to 5,600 ft
(1,060 to 1,700 m), in the montane wet
(Kauai, Oahu, Molokai, and Maui),
montane mesic (Kauai, Molokai, Maui,
and Hawaii Island), and wet cliff
(Molokai and Maui) ecosystems
(Duncan 1999, p. 1089; TNCH 2007;
HBMP 2010). Historically, R. mauiensis
was known from Kauai, Oahu, Molokai,
Maui, and Hawaii (HBMP 2010). Oahu
occurrences have not been observed
since the 1800s, and Hawaii Island
occurrences have not been observed
since 1980 (HBMP 2010). Currently, R.
mauiensis is known from Kauai (53
individuals) and east Maui (112
individuals). Two individuals formerly
known from Molokai have not been
observed on recent surveys (Bily 2007,
in litt.; Perlman 2007a, in litt.; Wood
2007b, in litt.; HBMP 2010; PEPP 2010,
p. 105; Bakutis 2011, in litt.; PEPP 2011,
p. 161; PEPP 2013, p. 177; Oppenheimer
2015, in litt.).
Feral pigs, goats, axis deer, blacktailed deer, and cattle modify and
destroy the habitat of Ranunculus
mauiensis on Kauai, Molokai, and Maui,
with evidence of the activities of these
animals reported in the areas where this
species occurs (HBMP 2010; PEPP 2014,
pp. 155–156). Ungulates are managed in
Hawaii as game animals (except for
cattle), but public hunting does not
adequately control the numbers of
ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Buddleja asiatica (dog tail), Clidemia
hirta, Erigeron karvinskianus,
Hedychium gardnerianum, Lantana
camara, Passiflora edulis (passion fruit),
P. tarminiana, Psidium cattleianum,
Rubus argutus, R. rosifolius, and
Tibouchina herbacea, modify and
destroy the native habitat of Ranunculus
mauiensis and displace native plant
species by competing for water,
nutrients, light, and space; they may
also produce chemicals that inhibit the
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010; PEPP 2014, p. 155). Herbivory by
slugs (Maui) and seed predation by rats
(Maui, Kauai) are both reported as
threats to R. mauiensis (HBMP 2010;
PEPP 2014, pp. 154–155). Stochastic
events such as drought (Maui),
landslides (Kauai), and fire (Maui) are
also reported as threats to R. mauiensis
(HBMP 2010). Erosion is a threat to
occurrences on Maui and Kauai (PEPP
2014, pp. 155–156). This species
experiences reduced reproductive vigor
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67805
due to low levels of genetic variability,
leading to diminished capacity to adapt
to environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 86) found that, as
environmental conditions are altered by
climate change, R. mauiensis is unlikely
to tolerate or adapt to projected changes
in temperature and moisture, and is
unlikely to be able to move to areas with
more suitable climatic conditions.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to R.
mauiensis described above.
The remaining occurrences of
Ranunculus mauiensis are at risk, the
known individuals are restricted to
small areas on Kauai and Maui, and
continue to be negatively affected by
habitat modification and destruction by
ungulates, direct competition with
nonnative plants, and herbivory and
predation by slugs and rats. Because of
its small, isolated occurrences,
landslides, drought, and erosion also
negatively affect this species. The small
number of remaining individuals limits
this species’ ability to adapt to
environmental changes. The effects of
climate change are likely to further
exacerbate these threats. Because of
these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Sanicula sandwicensis (NCN) is a
stout, erect, perennial herb in the
parsley family (Apiaceae) (Constance
and Affolter 1999, p. 210). This species
occurs from 6,500 to 8,500 ft (2,000 to
2,600 m) in shrubland and woodland on
the islands of Maui and Hawaii Island,
in the montane mesic (Hawaii Island
and Maui), montane dry (Hawaii
Island), and subalpine (Hawaii Island
and Maui) ecosystems (Constance and
Affolter 1999, p. 210; TNCH 2007;
NTBG Database 2014, in litt.). Sanicula
sandwicensis is historically known from
Haleakala on Maui and from Mauna
Kea, Mauna Loa, and Hualalai on
Hawaii Island (Constance and Affolter
1999, p. 210). Currently, there are more
than 50 individuals of S. sandwicensis
on east and west Maui (MNTF 2010, in
litt.; PEPP 2011, pp. 162–164;
Oppenheimer 2015, in litt.). In 2008, an
occurrence of fewer than 20 individuals
was found in Hawaii Volcanoes
National Park (Benitez et al. 2008, p.
59). Following ungulate removal, this
occurrence increased to as many as 45
individuals, with many juvenile plants
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(Orlando 2015, in litt.). A single
individual was found farther east at
about 7,400 ft (Orlando 2015, in litt.).
Feral pigs and goats modify and
destroy the habitat of Sanicula
sandwicensis on Maui, with evidence of
the activities of these animals reported
in the areas where this species occurs
(PEPP 2011, pp. 162–164; Oppenheimer
2015, in litt.). Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants modify and
destroy the habitat of S. sandwicensis
and displace native plant species by
competing for water, nutrients, light,
and space; they may also produce
chemicals that inhibit the growth of
other plants (Smith 1985, pp. 180–250;
Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74; PEPP 2011, pp. 162–
164). Those nonnative plants observed
to directly affect S. sandwicensis and its
habitat are Ageratina adenophora,
Anthoxanthum odoratum (sweet
vernalgrass), Epilobium ciliatum
(willow herb), Holcus lanatus (common
velvetgrass), Pinus spp., Prunella
vulgaris, and Rubus argutus (PEPP 2011,
pp. 162–164). Stochastic events such as
drought, flooding, and fires are all
reported to pose threats to this species
(PEPP 2011, pp. 162–164). Erosion is a
threat to occurrences on Maui (PEPP
2011, pp. 162–163). Herbivory by rats
also is a threat because they eat the
taproot, killing the plant (Oppenheimer
2015, in litt.). This species experiences
reduced reproductive vigor due to low
levels of genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 88) found that, as
environmental conditions are altered by
climate change, S. sandwicensis is
unlikely to tolerate or adapt to projected
changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to S. sandwicensis described
above.
The remaining occurrences of
Sanicula sandwicensis are at risk; the
known individuals are restricted to
small areas on Maui and Hawaii Island
and continue to be negatively affected
by habitat modification and destruction
by feral pigs and goats and by direct
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competition with nonnative plants.
Stochastic events such as drought,
flooding, erosion, and fires are threats to
this species. The small number of
remaining individuals limits this
species’ ability to adapt to
environmental changes. The effects of
climate change are likely to further
exacerbate these threats. Because of
these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Santalum involutum (iliahi) is a shrub
or small tree in the sandalwood family
(Santalaceae) (Harbaugh et al. 2010, pp.
827–838). Habitat for S. involutum is
mesic and wet forest on Kauai, from 400
to 2,500 ft (120 to 750 m), in the
lowland mesic and lowland wet
ecosystems (TNCH 2007; Harbaugh et
al. 2010, pp. 827–838). Historically, this
species was known from northern Kauai
at Kee, Hanakapiai, and Wainiha, and
from southern Kauai at Wahiawa, but
has not been observed in these areas for
30 years (Harbaugh et al. 2010, p. 835).
Currently, approximately 50 to 100
individuals occur in isolated forest
pockets on Kauai (Harbaugh et al. 2010,
p. 835; Wood 2015, in litt.).
Feral pigs and goats modify and
destroy the habitat of Santalum
involutum on Kauai, with evidence of
the activities of these animals reported
in the areas where this species occurs
(Harbaugh et al. 2010, pp. 835–836;
Wood 2015, in litt.). Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat modification and
destruction or herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Nonnative
plants modify and destroy the native
habitat of S. involutum and displace
native plant species by competing for
water, nutrients, light, and space; they
may also produce chemicals that inhibit
the growth of other plants (Smith 1985,
pp. 180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010). Nonnative plants reported to
modify and destroy habitat of S.
involutum include Clidemia hirta,
Hedychium gardnerianum, Lantana
camara, Melinis minutiflora, Psidium
cattleianum, P. guajava, and Rubus
argutus (Harbaugh et al. 2010, p. 836).
Herbivory and seed predation by rats is
a threat to this species (Harbaugh et al.
2010, p. 836; Wood 2015, in litt.).
Wildfire is a threat to this species in
mesic areas (Harbaugh et al. 2010, p.
836). This species experiences reduced
reproductive vigor due to low levels of
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genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
The remaining occurrences of
Santalum involutum are at risk; the
known individuals are restricted to a
small area on Kauai and continue to be
negatively affected by habitat
modification and destruction by
ungulates, direct competition with
nonnative plants, and by herbivory and
fruit predation by rats. The small
number of remaining individuals limits
this species’ ability to adapt to
environmental changes. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to S. involutum
described above. Because of these
threats, we find that this species is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Schiedea diffusa ssp. diffusa (NCN) is
a reclining or weakly climbing vine in
the pink family (Caryophyllaceae)
(Wagner et al. 1999, pp. 511–512;
Wagner et al. 2005, pp. 103–106).
Schiedea diffusa ssp. diffusa occurs in
wet forest from 3,000 to 5,300 ft (915 to
1,600 m) on Molokai, and to 6,700 ft
(2,050 m) on Maui, in the lowland wet
(Maui) and montane wet (Maui and
Molokai) ecosystems (Wagner et al.
1999, p. 512; TNCH 2007; HBMP 2010).
Historically, on Molokai, this subspecies
was known from Kawela to Waikolu
valleys, and on Maui it was wideranging on both the east and west
mountains (Wagner et al. 2005, p. 106).
Currently, S. diffusa ssp. diffusa is
known only from east Maui in scattered
occurrences (fewer than 50 individuals
total), in a much smaller range, with
some remaining in Haleakala National
Park (HBMP 2010; Gates 2015, in litt.).
Two occurrences were observed within
Hanawi NAR in 2005; however, their
current status is unknown (Vetter 2015,
in litt.). On Molokai, there were two
occurrences totaling fewer than 10
individuals; however, these have not
been seen since the 1990s (HBMP 2010;
Oppenheimer 2015, in litt.).
Feral pigs modify and destroy the
habitat of Schiedea diffusa ssp. diffusa
on Maui and Molokai, with evidence of
the activities of these animals reported
in the areas where this subspecies
occurs (HBMP 2010; PEPP 2014, p. 159).
Ungulates are managed in Hawaii as
game animals, but public hunting does
not adequately control the numbers of
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ungulates to eliminate habitat
modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Adiantum raddianum (NCN), Ageratina
adenophora, Hypochaeris radicata
(hairy cat’s ear), Juncus planifolius
(rush), Passiflora tarminiana, Prunella
vulgaris, Rubus argutus, and R.
rosifolius, modify and destroy the native
habitat of S. diffusa ssp. diffusa and
displace native plant species by
competing for water, nutrients, light,
and space; they may also produce
chemicals that inhibit the growth of
other plants (Smith 1985, pp. 180–250;
Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74; HBMP 2010; PEPP
2014, p. 159). Herbivory by slugs and
seed predation by rats are both reported
as threats to this subspecies (HBMP
2010; PEPP 2014, p. 159; Duvall 2015,
in litt.). This subspecies experiences
reduced reproductive vigor due to low
levels of genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
The remaining occurrences of
Schiedea diffusa ssp. diffusa are at risk.
The known individuals are restricted to
small areas on Maui and continue to be
negatively affected by habitat
modification and destruction by
ungulates, direct competition with
nonnative plants, and herbivory and
predation by slugs and rats. The small
number of remaining individuals limits
this subspecies’ ability to adapt to
environmental changes. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to S. diffusa ssp.
diffusa described above. Because of
these threats, we find that this
subspecies is endangered throughout all
of its range, and, therefore, find that it
is unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Schiedea pubescens (maolioli) is a
reclining or weakly climbing vine in the
pink family (Caryophyllaceae) (Wagner
et al. 1999, p. 519; Wagner et al. 2005,
pp. 99–102). This species occurs in
diverse mesic to wet Metrosideros forest
from 2,000 to 4,000 ft (640 to 1,220 m)
in the lowland wet, montane wet,
montane mesic, and wet cliff
ecosystems (Wagner et al. 1999, 519;
Wagner et al. 2005, p 100; TNCH 2007;
HBMP 2010). Historically, on Molokai,
this species was known from Kalae to
Pukoo ridge; on Lanai, it was known
from the Lanaihale summit area but has
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not been observed since 1922; on Maui,
it was known from the western
mountains at Olowalu, Kaanapali, and
Waihee, with a possible occurrence the
eastern mountains at Makawao (HBMP
2010). Currently, this species is known
from one occurrence on Molokai
totaling fewer than 30 individuals. The
occurrence on east Maui has not been
re-observed, but this species is found at
seven locations on west Maui (Wood
2001, in litt.; Oppenheimer 2006, in litt.;
Bakutis 2010, in litt.; HBMP 2010;
MNTF 2010, in litt.; Oppenheimer 2010,
in litt.; PEPP 2014, pp. 162–163;
Oppenheimer 2015, in litt.). It was
determined that the report of 4 to 6
individuals of S. pubescens at the PTA
on Hawaii Island was a
misidentification of individuals from
the species Schiedea hawaiiensis
(Wagner et al. 2005, pp. 93, 95).
Feral pigs, goats, axis deer, and cattle
modify and destroy the habitat of
Schiedea pubescens on Maui and
Molokai, with evidence of the activities
of these animals reported in the areas
where this species occurs (HBMP 2010;
PEPP 2014, p. 162). Ungulates are
managed in Hawaii as game animals
(except for cattle), but public hunting
does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Buddleja asiatica, Cestrum nocturnum
(night cestrum), Clidemia hirta, Erigeron
karvinskianus, Psidium cattleianum,
Rubus rosifolius, and Tibouchina
herbacea, modify and destroy the native
habitat of S. pubescens and displace
native plant species by competing for
water, nutrients, light, and space; they
may also produce chemicals that inhibit
the growth of other plants (Smith 1985,
pp. 180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010; PEPP 2014, pp. 162–163).
Herbivory by slugs and seed predation
by rats are both reported to be threats to
S. pubescens on Maui (HBMP 2010;
PEPP 2014, p. 162; Duvall 2015, in litt.).
Stochastic events such as drought,
erosion, fire, and flooding are also
reported as threats to S. pubescens
(HBMP 2010; PEPP 2014, p. 162;
Oppenheimer 2015, in litt.). This
species is outcrossing; however, very
low population sizes may have reduced
its genetic variation (Weller 2015, in
litt.). Fortini et al. (2013, p. 88) found
that, as environmental conditions are
altered by climate change, S. pubescens
is unlikely to tolerate or adapt to
projected changes in temperature and
moisture, and is unlikely to be able to
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move to areas with more suitable
climatic conditions. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to S. pubescens
described above.
The remaining occurrences of
Schiedea pubescens are at risk. The
known individuals are restricted to
small areas on Molokai and Maui, and
continue to be negatively affected by
habitat modification and destruction by
ungulates, direct competition with
nonnative plants, and herbivory and
predation by slugs and rats. Landslides,
flooding, fire, and drought impact this
species. The small number of remaining
individuals limits this species’ ability to
adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Sicyos lanceoloideus (anunu) is a
perennial vine in the gourd family
(Cucurbitaceae) (Telford 1999, p. 581;
Wagner and Shannon 1999, p. 444).
Sicyos lanceoloideus occurs on ridges or
spurs in mesic forest from 1,800 to 2,700
ft (550 to 800 m), in the dry cliff (Oahu),
lowland mesic (Oahu, Kauai), and
montane mesic (Kauai) ecosystems
(Telford 1999, p. 581; TNCH 2007;
HBMP 2010). Sicyos lanceoloideus was
historically found at Kalalau Valley and
Waimea Canyon on Kauai and in the
Waianae Mountains on Oahu (Telford
1999, p. 581). Currently, on Kauai, there
are four individuals in three locations
(Kishida 2015, in litt.). On Oahu, this
species occurs in 5 locations in the
Waianae Mountains totaling fewer than
35 individuals (HBMP 2010; U.S. Army
2014 database). Because this species is
a vine, determining exact numbers is
difficult (PEPP 2013, p. 189). In
addition, occurrences and numbers vary
widely as individuals have been
observed to persist for fewer than 7
years (Sailer 2015, in litt.).
Feral pigs, goats, and black-tailed deer
modify and destroy the habitat of Sicyos
lanceoloideus on Kauai and Oahu, with
evidence of the activities of these
animals reported in the areas where this
species occurs (HBMP 2010; PEPP 2013,
p. 189; PEPP 2014, p. 166; Williams
2015, in litt.). Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
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litt.). Nonnative plants, such as
Clidemia hirta, Lantana camara, Melia
azedarach (chinaberry), Paspalum
urvillei (vasey grass), Passiflora edulis,
Pluchea carolinensis (sourbush),
Psidium cattleianum, P. guajava,
Ricinus communis (castor bean), Rubus
argutus, Schinus terebinthifolius, and
Stachytarpheta jamaicensis, modify and
destroy the native habitat of Sicyos
lanceoloideus, and displace native plant
species by competing for water,
nutrients, light, and space; they may
also produce chemicals that inhibit the
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010; Sailer 2015, in litt.). Drought is
also reported as a threat to S.
lanceoloideus (PEPP 2014, p. 166;
HBMP 2010; Sailer 2015, in litt.). Fires
are a threat to the occurrence in the
Waianae Mountains of Oahu (Sailer
2015, in litt.). Because of the small
remaining number of individuals, this
species experiences reduced
reproductive vigor due to low levels of
genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 89) found that, as
environmental conditions are altered by
climate change, S. lanceoloideus is
unlikely to tolerate or adapt to projected
changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to S. lanceoloideus described
above.
The remaining occurrences of Sicyos
lanceoloideus are at risk. The known
individuals are restricted to small areas
on Kauai and Oahu and continue to be
negatively affected by habitat
modification and destruction by
ungulates, direct competition with
nonnative plants, and stochastic events
such as drought and fire. The small
number of remaining individuals limits
this species’ ability to adapt to
environmental change. The effects of
climate change are likely to further
exacerbate these threats. Because of
these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Sicyos macrophyllus (anunu) is a
perennial vine in the gourd family
(Cucurbitaceae) (Telford 1999, p. 578;
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Wagner and Shannon 1999, p. 444).
Typical habitat is wet Metrosideros
polymorpha forest and Sophora
chrysophylla-Myoporum sandwicense
(mamane-naio) forest, from 4,000 to
6,600 ft (1,200 to 2,000 m) in the
montane mesic (Hawaii Island),
montane wet (Maui), and montane dry
(Hawaii Island) ecosystems (Telford
1999, p. 578; TNCH 2007; HBMP 2010).
Historically, S. macrophyllus was
known from Puuwaawaa, Laupahoehoe,
Puna, and South Kona on Hawaii Island,
and from Kipahulu Valley on the island
of Maui (HBMP 2010). Currently, S.
macrophyllus is known from 10
occurrences totaling between 24 and 26
individuals on Hawaii Island (Bio 2008,
pers. comm.; Pratt 2008, in litt.; HBMP
2010; Evans 2015b, in litt.; Orlando
2015, in litt.). This species has been
outplanted at several sites in Hawaii
Volcanoes National Park and is
persisting (Orlando 2015, in litt.). The
individual on Maui has not been
observed since 1987 (HBMP 2010).
Feral pigs, mouflon, and cattle modify
and destroy the habitat of Sicyos
macrophyllus on the island of Hawaii,
with evidence of the activities of these
animals reported in the areas where this
species occurs (HBMP 2010). Ungulates
are managed in Hawaii as game animals
(except for cattle), but public hunting
does not adequately control the
numbers of ungulates to eliminate
habitat modification and destruction or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as
Cenchrus setaceus, Delairea odorata
(German ivy), Ehrharta stipoides, and
Pennisetum clandestinum, modify and
destroy the native habitat of S.
macrophyllus and displace native plant
species by competing for water,
nutrients, light, and space; they may
also produce chemicals that inhibit the
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010). Seed predation by rats is reported
to pose a threat to this species (HBMP
2010). Stochastic events such as fire are
also reported as a threat to S.
macrophyllus (HBMP 2010). This
species experiences reduced
reproductive vigor due to low levels of
genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 89) found that, as
environmental conditions are altered by
climate change, S. macrophyllus is
unlikely to tolerate or adapt to projected
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changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to S. macrophyllus described
above.
The remaining occurrences of Sicyos
macrophyllus are at risk. The only
known individuals are restricted to
small areas on Hawaii Island and
continue to be negatively affected
habitat modification and destruction by
ungulates, direct competition with
nonnative plants, and seed predation by
rats. Fire is also a threat to this species.
The small number of remaining
individuals limits this species’ ability to
adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Solanum nelsonii (popolo) is a
sprawling or trailing shrub up to 3 ft (1
m) tall, in the nightshade family
(Solanaceae) (Symon 1999, pp. 1273–
1274). Typical habitat for this species is
coral rubble or sand in coastal sites up
to 490 ft (150 m), in the coastal
ecosystem (Symon 1999, pp. 1273–1274;
TNCH 2007; HBMP 2010). Historically,
S. nelsonii was known from Kaalualu,
Kamilo, and Kaulana Bay, and South
Point (5 individuals total) on Hawaii
Island; from Kealea Bay, Kawaewaae,
and Leahi on Niihau; and from the
Northwest Hawaiian Islands of Nihoa,
Laysan, Pearl and Hermes, and Kure
Atoll (Green Island) (Lamoureux 1963,
p. 6; Clapp et al. 1977, p. 36; HBMP
2010). This species was last collected on
Niihau in 1949 (HBMP 2010). The only
known individual on Maui was reported
to have disappeared in the mid-1990s
after cattle had been allowed to graze in
its last known habitat (HBMP 2010;
Duvall 2015, in litt.). Currently, S.
nelsonii occurs in the coastal ecosystem
on the islands of Hawaii and Molokai
(approximately 50 individuals), and on
the Northwest Hawaiian Islands of Kure
(an unknown number of individuals),
Midway (approximately 260 individuals
on Sand, Eastern, and Spit islands),
Laysan (approximately 490 individuals),
Pearl and Hermes (30 to 100
individuals), and Nihoa (8,000 to 15,000
individuals) (Aruch 2006, in litt.;
Rehkemper 2006, in litt.; Tangalin 2006,
in litt.; Bio 2008 a and 2008b, in litt.;
Vanderlip 2011, in litt.; Conry 2012, in
litt.; PEPP 2013, pp. 190–191).
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Axis deer and feral cattle modify and
destroy the habitat of Solanum nelsonii
on the main Hawaiian islands of Maui,
Molokai, and Hawaii, with evidence of
the activities of these animals reported
in the areas where this species occurs
(HBMP 2010). Ungulates are managed in
Hawaii as game animals (except for
cattle), but public hunting does not
adequately control the numbers of
ungulates to eliminate habitat
modification and destruction, and
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants, such as Lantana
camara, Leucaena leucocephala,
Pennisetum ciliare, Prosopis pallida,
and Setaria verticillata (bristly foxtail),
modify and destroy the native habitat of
S. nelsonii both on the main Hawaiian
Islands and on some of the Northwest
Hawaiian Islands (HBMP 2010).
Nonnative plants displace native plant
species by competing for water,
nutrients, light, and space, or they may
produce chemicals that inhibit the
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; PEPP
2008, p. 110; HBMP 2010). Seed
predation by rats has been reported as
a threat to S. nelsonii on Molokai (PEPP
2014, p. 167). Stochastic events such as
drought, erosion, fire, and flooding are
also reported as threats to S. nelsonii
(HBMP 2010; PEPP 2014, p. 167). In
2011, a tsunami swept over Midway
Atoll’s Eastern Island and Kure Atoll’s
Green Island, inundating S. nelsonii
plants, spreading plastic debris, and
destroying seabird nesting areas,
reaching about 500 ft (150 m) inland
(DOFAW 2011, in litt.; Starr 2011, in
litt.; USFWS 2011, in litt.). Occurrences
of this species on the main Hawaiian
Islands and on some of the Northwest
Hawaiian Islands experience reduced
reproductive vigor due to low levels of
genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
The effects of climate change resulting
in sea-level rise will alter environmental
conditions and ecosystem that support
this species. Fortini et al. (2013, p. 89)
found that, as environmental conditions
are altered by climate change, S.
nelsonii is unlikely to tolerate or adapt
to projected changes in temperature and
moisture, and is unlikely to be able to
move to areas with more suitable
climatic conditions. Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
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exacerbate the threats to S. nelsonii
described above.
The remaining occurrences of
Solanum nelsonii on the main Hawaiian
Islands are restricted to small areas of
Molokai and Hawaii Island, and
continue to be negatively affected by
habitat modification and destruction by
ungulates, direct competition with
nonnative plants, and herbivory and
predation by rats. Even though most
individuals of S. nelsonii in the
Northwestern Hawaii Islands are found
on lands managed by the Service as part
of the Hawaiian Islands National
Wildlife Refuge, the relatively isolated
occurrences of S. nelsonii there are
negatively affected by nonnative plants.
The small number of remaining
individuals limits this species’ ability to
adapt to environmental changes. A
tsunami occurred and impacted habitat
for this species, and sea level rise
associated with global warming will
modify and destroy habitat for S.
nelsonii in the low-lying Northwestern
Hawaiian Islands. Because of these
threats, we find that this species is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Stenogyne kaalae ssp. sherffii (NCN)
is a climbing vine in the mint family
(Lamiaceae) (Wagner and Weller 1999,
pp. 448–449; Weller and Sakai 1999, p.
838). This species occurs in the Koolau
Mountains of Oahu, in diverse forest
from 1,500 to 1,600 ft (450 to 490 m) in
the lowland wet ecosystem (Wagner and
Weller 1999, pp. 448–449; TNCH 2007;
HBMP 2010; U.S. Army 2014 database).
Stenogyne kaalae ssp. sherffii is
historically known from diverse mesic
forest in the Waianae Mountains of
Oahu, and from the lowland wet
ecosystem of the Koolau Mountains
(although, as described in the proposed
rule, it was thought to be a different
species, S. sherffii, until the mid-1990s).
This subspecies occurred within a very
small range in the northern Koolau
Mountains, but now all wild individuals
are extirpated. There are propagules
from collections from those plants that
have been outplanted in the same areas
(PEPP 2014, p. 169; Ching Harbin 2015,
in litt.).
Feral pigs modify and destroy the
habitat of Stenogyne kaalae ssp. sherffii
on Oahu, with evidence of the activities
of these animals reported in the areas
where this subspecies occurred (HBMP
2010; PEPP 2014, p. 169). Ungulates are
managed in Hawaii as game animals,
but public hunting does not adequately
control the numbers of ungulates to
eliminate habitat destruction and
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modification, and herbivory by these
animals (Anderson et al. 2007, in litt.;
HAR–DLNR 2010, in litt.). Nonnative
plants, such as Blechnum
appendiculatum (NCN), Clidemia hirta,
Cyclosorus parasiticus (NCN), Psidium
cattleianum, and Rubus rosifolius,
destroy and modify the native habitat of
S. kaalae ssp. sherffii and displace
native plant species by competing for
water, nutrients, light, and space; they
may also produce chemicals that inhibit
the growth of other plants (Smith 1985,
pp. 180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010). This subspecies experiences
reduced reproductive vigor due to low
levels of genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361).
Fortini et al. (2013, p. 90) found that, as
environmental conditions are altered by
climate change, S. kaalae ssp. sherffii is
unlikely to tolerate or adapt to projected
changes in temperature and moisture,
and is unlikely to be able to move to
areas with more suitable climatic
conditions. Although we cannot predict
the timing, extent, or magnitude of
specific impacts, we do expect the
effects of climate change to exacerbate
the threats to S. kaalae ssp. sherffii
described above.
Any remaining occurrences of
Stenogyne kaalae ssp. sherffii are at risk;
the last known wild individuals were
restricted to a very small area on Oahu,
and the habitat continues to be
negatively affected by habitat
modification and destruction by
ungulates and direct competition with
nonnative plants. The small number of
remaining individuals (outplanted only)
limits this subspecies’ ability to adapt to
environmental changes. The effects of
climate change are likely to further
exacerbate these threats. Because of
these threats, we find that this
subspecies is endangered throughout all
of its range, and, therefore, find that it
is unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Wikstroemia skottsbergiana (akia) is a
shrub or small tree in the akia family
(Thymelaceae) (Peterson 1999, p. 1290).
Wikstroemia skottsbergiana occurs in
wet forest on the island of Kauai, in the
lowland wet ecosystem (Peterson 1999,
p. 1290; TNCH 2007). Wikstroemia
skottsbergiana is historically known
from the Wahiawa Mountains, Hanalei
Valley, and Kauhao Valley, on the
island of Kauai (Peterson 1999, p. 1290).
Currently, this species is limited to 30
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individuals at one location (PEPP 2012,
p. 26; Wood 2015, in litt.).
Feral pigs and goats destroy and
modify the habitat of Wikstroemia
skottsbergiana on Kauai, with evidence
of the activities of these animals
reported in the areas where this species
occurs (DLNR 2005, in litt.; Wood 2015,
in litt.). Ungulates are managed in
Hawaii as game animals, but public
hunting does not adequately control the
numbers of ungulates to eliminate
habitat destruction and modification or
herbivory by these animals (Anderson et
al. 2007, in litt.; HAR–DLNR 2010, in
litt.). Nonnative plants destroy and
modify the native habitat of W.
skottsbergiana and displace native plant
species by competing for water,
nutrients, light, and space; they may
also produce chemicals that inhibit the
growth of other plants (Smith 1985, pp.
180–250; Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74; HBMP
2010). Predation of seeds by rats is a
threat to this species (DLNR 2005, in
litt.). Landslides are a threat to the only
known occurrence of this species (Wood
2015, in litt.). This species experiences
reduced reproductive vigor due to low
levels of genetic variability, leading to
diminished capacity to adapt to
environmental changes, thereby
lessening the probability of its long-term
persistence (Barrett and Kohn 1991, p.
4; Newman and Pilson 1997, p. 361;
DLNR 2005, in litt.).
The remaining occurrences of
Wikstroemia skottsbergiana are at risk.
The known individuals are restricted to
a very small area on Kauai and continue
to be negatively affected by habitat
modification and destruction by
ungulates, direct competition with
nonnative plants, and seed predation by
rats. The small number of remaining
individuals limits this species’ ability to
adapt to environmental changes.
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the threats to W.
skottsbergiana described above. Because
of these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Animals
Band-Rumped Storm-Petrel
(Oceanodroma castro)—Hawaii
Population
The band-rumped storm-petrel, a
small seabird, is a member of the family
Hydrobatidae (order Procellariiformes)
and a member of the Northern
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Hemisphere subfamily Hyrdrobatinae
(Slotterback 2002, p. 2). This seabird is
found in several areas of the subtropical
Pacific and Atlantic Oceans (del Hoyo et
al. 1992 in Bird Life International 2015,
in litt.). The Atlantic breeding
populations are restricted to islands in
the eastern portions: Cape Verde,
Ascension, Madeira, and the Azores
Islands (Allan 1962, p. 274; Harrison
1983, p. 274). Wintering birds may
occur as far west as the mid-Atlantic;
however, Atlantic breeding populations
are not within the borders of the United
States or in areas under U.S.
jurisdiction. Three widely separated
breeding areas occur in the Pacific: in
Japan, Hawaii, and Galapagos
(Richardson 1957, p. 19; Harris 1969, p.
96; Harrison 1983, p. 274). The Japanese
population, which breeds on islets off
the east coast of Japan (Hidejima and
Sanganjima in Allan 1962, p. 274; Harris
1969, p. 96), ranges within 860 mi
(1,400 km) east and south of the
breeding colonies. Populations in Japan
and Galapagos total as many as 23,000
pairs (Boersma and Groom 1993, p.
114); however, a recent survey on
Hidejima Island revealed only 117
burrows, some of which were occupied
by Leach’s storm petrels (Oceanodroma
leucorhoa) (Biodiversity Center of Japan
2014, p. 1). Surveyors noted that the
nesting area had been affected by
extensive erosion caused by the 2011
earthquake and tsunami (Biodiversity
Center of Japan 2014, p. 1).
When Polynesians arrived about 1,500
years ago, the band-rumped storm-petrel
probably was common on all of the
main Hawaiian Islands (Harrison et al.
1990, pp. 47–48). As evidenced by
bones found in middens on Hawaii
Island (Harrison et al. 1990, pp. 47–48)
and in excavation sites on Oahu and
Molokai (Olson and James 1982, pp. 30,
33), band-rumped storm-petrels were
once numerous enough to be harvested
for food and possibly for their feathers
(Harrison et al. 1990, p. 48).
In Hawaii, band-rumped storm-petrels
are known to nest in remote cliff
locations on Kauai and Lehua Island, in
steep open to vegetated cliffs, and in
little vegetated, high-elevation lava
fields on Hawaii Island (Wood et al.
2002, p. 17–18; VanderWerf et al. 2007,
pp. 1, 5; Joyce and Holmes 2010, p. 3;
Banko 2015 in litt.; Raine 2015, in litt.).
Vocalizations were heard in Haleakala
Crater on Maui in 1992 (Johnston 1992,
in Wood et al. 2002, p. 2), on Lanai
(Penniman 2015, in litt.), and in Hawaii
Volcanoes National Park (Orlando 2015,
in litt.). Based on the scarcity of known
breeding colonies in Hawaii and their
remote, inaccessible locations today
compared to prehistoric population
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levels and distribution, the bandrumped storm-petrel appears to be
significantly reduced in numbers and
range following human occupation of
the Hawaiian Islands, likely as a result
of predation by nonnative mammals and
habitat loss.
Taxonomists have typically combined
the Pacific populations of band-rumped
storm-petrel into a single taxon, and
currently the American Ornithologist’s
Union (AOU) regards the species as
monotypic (2015, in litt.). However,
molecular studies are ongoing and
indicate genetic differences between
populations in different oceans and
archipelagos (Friesen et al. 2007b, pp.
18590–18952; Smith et al. 2007, p. 770;
Taylor et al., in prep in Raine 2015, in
litt.) and between sympatric populations
that breed in different seasons (e.g., in
the Galapagos Islands; Smith and
Friesen 2007, pp. 1599–1560; Smith et
al. 2007, p. 756).
Band-rumped storm-petrels are
regularly observed in coastal waters
around Kauai, Niihau, and Hawaii
Island (Harrison et al. 1990, p. 49;
Holmes and Joyce 2009, 4 pp.), and in
‘‘rafts’’ (regular concentrations) of a few
birds to as many as 100, possibly
awaiting nightfall before coming ashore
to breeding colonies. Kauai likely has
the largest population, with an
estimated 221 nesting pairs in cliffs
along the north shore of the island in
2002, and additional observations on
the north and south side of the island
in 2010 (Harrison et al. 1990, p. 49;
Wood et al. 2002, pp. 2–3; Holmes and
Joyce 2009, 4 pp.; Joyce and Holmes
2010, pp. 1–3). Audio detections for
Kauai indicate this species may be
predominantly breeding on the Na Pali
coast and Waimea Canyon, with a very
small number in Wainiha Valley (Raine
2015, in litt.). The band-rumped stormpetrel is also known from Lehua Island
(as detected there by auditory surveys)
(VanderWerf et al. 2007, p.1; Raine
2015, in litt.), Maui (Mitchell et al. 2005,
in litt.), Kahoolawe (Olson 1992, pp. 38,
112), Lanai (Penniman 2015, in litt.) and
Hawaii Island (Mitchell et al. 2005, in
litt.; Orlando 2015, in litt.). Additional
surveys have been conducted on several
islands in recent years, including
surveys confirming the presence of
band-rumped storm-petrels at the PTA
on Hawaii Island, but further data are
not yet available (Swift 2015, in litt.).
The species likely once nested in coastal
Maui, where the remains of a chick were
found in 1999, and islands such as
Niihau and Kaula, where surveys have
not been conducted, likely have suitable
nesting habitat and may harbor the
species (Penniman 2015, in litt.). We do
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not have a current estimate of total
numbers in Hawaii at this time.
Nesting sites are in burrows and in
crevices, holes, and on protected ledges
along cliff faces, where a single egg is
laid (Allan 1962, p. 274–275; Harris
1969, pp. 104–105; Slotterback 2002, p.
11). Predation by nonnative animals on
nests and adults during the breeding
season is the greatest threat to the
Hawaiian population of the bandrumped storm-petrel. These predators
include feral cats (Felis catus), barn
owls (Tyto alba), small Indian mongoose
(Herpestes auropunctatus), black rats
(Rattus rattus), Norway rats (R.
norvegicus), and Polynesian rats (R.
exulans) (Scott et al. 1986, pp. 1, 363–
364; Tomich 1986, pp. 37–45; Harrison
et al. 1990, pp. 47–48; Slotterback 2002,
p. 19; Raine 2015, in litt.). Attraction of
fledglings to artificial lights and
collisions with structures, such as
communication towers and utility lines,
is also a threat (Reed et al. 1985, p. 377;
Telfer et al. 1987, pp. 412–413; Harrison
et al. 1990, p. 49; Banko et al. 1991, p.
651; Cooper and Day 1998, p. 18;
Podolsky et al. 1998, pp. 21, 27–30;
Holmes and Joyce 2009, p. 2).
Monitoring of power lines on Kauai has
recorded over 1,000 strikes by seabirds
annually (mostly Newell’s shearwaters
(Puffinus newelli); Travers et al. 2014,
pp. 19, 42) that may result in injury or
death. Recent studies of attraction of
seabirds to artificial lights indicate that
40 percent of those downed by
exhaustion (from circling the lights) are
killed by collisions with cars or other
objects (Anderson 2015, p. 4–13). The
small numbers of these birds and their
nesting areas on remote cliffs make
population-level impacts difficult to
document. However, the band-rumped
storm-petrel has similar behavior, lifehistory traits, and habitat needs to the
Newell’s shearwater, a threatened
species that has sustained major losses
as a result of light attraction and
collisions with lines or other objects
(Banko et al. 1991, p. 651; Banko 2015,
in litt.; Raine 2015, in litt.). Therefore,
we conclude that these are threats to the
band-rumped storm-petrel as well.
Erosion and landslides at nest sites
caused by the actions of nonnative
ungulates is a threat in some locations
on the island of Kauai (Raine 2015, in
litt.). Nonnative plants outcompete
native plants and can also affect nesting
sites of the band-rumped storm-petrel
by accelerating erosion, leading to
landslides and rockfalls (Wood et al.
2002, pp. 7–19). Regulatory mechanisms
(e.g., the Migratory Bird Treaty Act
(MBTA; 16 U.S.C. 703 et seq.))
contribute minimally to the active
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recovery and management of this
species (USFWS 2013, in litt.). The
small population size and limited
distribution of the band-rumped stormpetrel in Hawaii is a threat to this
´
population (Soule 1987, p. 8; Lande
1988, pp. 1455, 1458–1459; Harrison et
al. 1990, p. 50; Furness 2003, p. 33).
During the breeding season, a single
hurricane or landslide caused by
erosion could cause reproductive failure
and kill a significant number of adult
birds. Commercial fisheries and ocean
pollution have negative impacts to
seabirds, and also are likely to have
negative impacts to the band-rumped
storm petrel, although the information
about the impacts of fisheries and
plastics on storm-petrel species is
limited. In this rule, our listing
determination applies only to the
Hawaiian population of the bandrumped storm-petrel (see Distinct
Population Segment (DPS) Analysis,
below). Because of the deleterious and
cumulative effects to the band-rumped
storm-petrel caused by the threats
described above, we find that the
Hawaii population is endangered
throughout its range, and, therefore, we
find that it is unnecessary to analyze
whether it is endangered or threatened
in a significant portion of its range.
Yellow-Faced Bees (Hylaeus spp.)
Bees in the genus Hylaeus (family
Colletidae), which includes the seven
species in this final rule, are commonly
known as yellow-faced bees or masked
bees for their yellow-to-white facial
markings. All Hylaeus bees roughly
resemble small wasps in appearance;
however, Hylaeus bees have plumose
(branched) hairs on the body that are
longest on the sides of the thorax, which
readily distinguish them from wasps
(Michener 2000, in litt.). Bees in the
family Colletidae are also referred to as
plasterer bees because they line their
nests with a self-secreted, cellophanelike material. Eggs hatch and develop
into larvae (immature stage) and as
larvae grow, they molt through three
successive stages (instars), then change
into pupae (a resting form) in which
they metamorphose and emerge as
adults (Michener 2000, in litt.). The diet
of the larval stages is unknown,
although it is presumed the larvae feed
on stores of pollen and nectar collected
and deposited in the nest by the adult
female.
Yellow-Faced Bee (Hylaeus
anthracinus)
Hylaeus anthracinus was historically
known from numerous coastal and
lowland dry forest habitats up to 2,000
ft (610 m) in elevation on the islands of
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67811
Hawaii, Maui, Lanai, Molokai, and
Oahu, and in some areas was ‘‘locally
abundant.’’ Between 1997 and 1998,
surveys for Hawaiian Hylaeus were
conducted at 43 sites that were either
historical collecting localities or
potential suitable habitat. Hylaeus
anthracinus was observed at 13 of the
43 survey sites, but was not found at
any of the 9 historically occupied sites
(Daly and Magnacca 2003, p. 217).
Several of the historical collection sites
have been urbanized or are dominated
by nonnative vegetation (Liebherr and
Polhemus 1997, pp. 346–347; Daly and
Magnacca 2003, p. 55; Magnacca 2007,
pp. 186–188). There has been a dramatic
decline in abundance or presence of H.
anthracinus since surveys conducted in
1999 through 2002, noted on surveys
conducted between 2011 and 2013
(Magnacca 2015, in litt.). Currently, H.
anthracinus is known from 15 small
patches of coastal and lowland dry
forest habitat (Magnacca 2005a, in litt.,
p. 2); 5 locations on the island of Hawaii
in the coastal ecosystem; 2 locations on
Maui in the coastal and lowland dry
ecosystems; 1 location on Kahoolawe in
the lowland dry ecosystem; 3 locations
on Molokai in the coastal ecosystem,
and 4 locations on Oahu in the coastal
ecosystem (Daly and Magnacca 2003, p.
217; Magnacca 2005a, in litt., p. 2;
Magnacca and King 2013, pp. 13–14;
Graham 2015, in litt.). These 15
locations supported small populations
of H. anthracinus, but the number of
individual bees is unknown. In 2004, a
single individual was collected in
montane dry forest on the island of
Hawaii (possibly a vagrant); however,
the presence of additional individuals
has not been confirmed at this site
(Magnacca 2005a, in litt., p. 2).
Although this species was previously
unknown from the island of Kahoolawe,
it was observed at one location on the
island in 2002 (Daly and Magnacca
2003, p. 55). Additionally, during
surveys between 1997 and 2008, H.
anthracinus was absent from 17 other
sites with potentially suitable habitat
from which other species of Hylaeus
were collected (Daly and Magnacca
2003, pp. 4, 55) on Hawaii Island, Maui,
Lanai, Molokai, and Oahu.
Habitat destruction and modification
by urbanization and land use
conversion lead to the direct
fragmentation of foraging and nesting
areas used by Hylaeus anthracinus.
Habitat destruction and modification by
nonnative plants adversely impacts
native plant species by modifying the
availability of light, altering soil-water
regimes, modifying nutrient cycling,
altering the fire characteristics
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(increasing the fire cycle), and
ultimately converting native dominated
plant communities to nonnative plant
communities, and results in removal of
food sources and nesting sites for H.
anthracinus (Graham 2015, in litt.).
Habitat modification and destruction by
nonnative animals such as feral pigs,
goats, axis deer, and cattle, is
considered one of the primary factors
underlying degradation of native
vegetation in the Hawaiian Islands, and
these habitat changes also remove food
sources and nesting sites for H.
anthracinus (Stone 1985, pp. 262–263;
Cuddihy and Stone 1990, pp. 60–66,
73). Fire is a threat to H. anthracinus,
as it destroys native coastal and lowland
plant communities on which the species
depends, and opens habitat for
increased invasion by nonnative plants.
Because of the greater frequency,
intensity, and duration of fires that have
resulted from the human alteration of
landscapes and the introduction of
nonnative plants, especially grasses,
fires are now more destructive to the
coastal and lowland dry ecosystems
(Brown and Smith 2000, p. 172). A
single grass-fueled fire often kills most
native trees and shrubs in the area
(D’Antonio and Vitousek 1992, p. 74)
and could destroy food and nesting
resources for H. anthracinus. The
number and size of wildfires are
increasing in the main Hawaiian
Islands; however, their occurrences and
locations are unpredictable, and could
affect habitat for yellow-faced bees at
any time (Gima 1998, in litt.; County of
Maui 2009, Ch. 3, p. 3; Hamilton 2009,
in litt.; Honolulu Advertiser 2010, in
litt.; Pacific Disaster Center 2011, in
litt.). Random, naturally occurring
events such as hurricanes, tsunami, and
drought can also modify and destroy
habitat of H. anthracinus by creating
disturbed areas conducive to invasion
by nonnative plants and by eliminating
food and nesting resources (Kitayama
and Mueller-Dombois 1995, p. 671;
Businger 1998, pp. 1–2; Magnacca 2015,
in litt.). Predation by nonnative ants
including the big-headed ant (Pheidole
megacephala), the yellow crazy ant
(Anoplolepis gracilipes), Solenopsis
papuana (NCN), and S. geminata (NCN)
on Hylaeus egg, larvae, and pupal stages
is a threat to H. anthracinus, and ants
also compete with H. anthracinus for
their nectar food and nesting resources
(Howarth 1985, p. 155; Hopper et al.
1996, p. 9; Holway et al. 2002, pp. 188,
209; Daly and Magnacca 2003, p. 9;
Lach 2008, p. 155; Graham 2015, in
litt.). Predation by nonnative western
yellow jacket wasps (Vespula
pensylvanica) is a threat to H.
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anthracinus because the wasp is an
aggressive, generalist predator, and
occurs in great numbers in many habitat
types, from sea level to over 8,000 ft
(2,450 m), including areas where H.
anthracinus and other yellow-faced bees
occur (Gambino et al. 1987, p. 169;
Graham 2015, in litt.). Existing
regulatory mechanisms and agency
policies do not address the primary
threats to the yellow-faced bees and
their habitat from nonnative ungulates.
Competition with nonnative bees
(honeybees, carpenter bees, sweat bees
(Lasioglossum spp.), and alien Hylaeus
bees) for nectar and pollen, and by
exclusion from foraging, is a potential
threat to H. anthracinus (Magnacca
2007, p. 188; Graham 2015, in litt.;
Magnacca 2015, in litt.). The small
number of populations and individuals
of H. anthracinus makes this species
more vulnerable to extinction because of
the higher risks from genetic
bottlenecks, random demographic
fluctuations, and localized catastrophes
such as hurricanes, tsunami, and
drought (Daly and Magnacca 2003, p. 3;
Magnacca 2007, p. 173; Magnacca 2015,
in litt.). Although we cannot predict the
timing, extent, or magnitude of specific
impacts, we do expect the effects of
climate change to exacerbate the threats
to H. anthracinus described above. In
addition, disease has been suggested as
a threat, as pathogens carried by
nonnative bees, wasps, and ants could
be transmitted to H. anthracinus
through shared food sources (Graham
2015, in litt.); however, we have no
reports of this type of disease
transmission at this time.
The remaining populations of Hylaeus
anthracinus and its habitat are at risk.
The known individuals are restricted to
15 locations on Hawaii, Maui,
Kahoolawe, Molokai, and Oahu and
continue to be negatively affected by
habitat destruction and modification by
urbanization and land-use conversion,
and by habitat destruction and removal
of food and nesting sites by nonnative
ungulates and nonnative plants. Habitat
destruction by fire is a threat. Randomly
occurring events such as hurricanes and
drought modify habitat and remove food
and nesting sources for H. anthracinus.
Predation by nonnative ants and wasps
is a threat. Existing regulatory
mechanisms and agency policies do not
address the primary threats to the
yellow-faced bees and their habitat from
nonnative ungulates. Competition with
nonnative bees for food and nesting
sites is a threat. The small number of
remaining populations limits this
species’ ability to adapt to
environmental changes. The effects of
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climate change are likely to further
exacerbate these threats. Because of
these threats, we find that H.
anthracinus is endangered throughout
all of its range, and, therefore, find that
it is unnecessary to analyze whether it
is endangered or threatened in a
significant portion of its range.
Yellow-Faced Bee (Hylaeus assimulans)
Historically, Hylaeus assimulans was
known from numerous coastal and
lowland dry forest habitats up to 2,000
ft (610 m) in elevation on the islands of
Maui (coastal and lowland dry
ecosystems), Lanai (lowland dry
ecosystem), and Oahu (coastal and
lowland dry ecosystem). There are no
collections from Molokai although it is
likely H. assimulans occurred there
because all other species of Hylaeus
known from Maui, Lanai, and Oahu also
occurred on Molokai (Daly and
Magnacca 2003, pp. 217–229). Between
1997 and 1998, surveys for Hawaiian
Hylaeus were conducted at 25 sites on
Maui, Kahoolawe, Lanai, Molokai, and
Oahu. Hylaeus assimulans was absent
from 6 of its historical localities on
Maui, Lanai, and Oahu, and was not
observed at the remaining 19 sites with
potentially suitable habitat (Daly and
Magnacca 2003, pp. 56, 217; Magnacca
2005b, in litt., p. 2; Magnacca 2007, pp.
177, 181, 183; Xerces Society 2009, p.
4). Currently, H. assimulans is known
from a few small patches of coastal and
lowland dry forest habitat (Magnacca
2005b, in litt., p. 2) in two locations on
Maui in the lowland dry ecosystem; one
location on Kahoolawe in the coastal
ecosystem; and two locations on Lanai
in the lowland dry ecosystem (Daly and
Magnacca 2003, p. 58; Magnacca 2005b,
in litt., p. 2). This species has likely
been extirpated from Oahu because it
has not been observed since Perkin’s
1899 surveys, and was not found during
recent surveys of potentially suitable
habitat on Oahu at Kaena Point,
Makapuu, and Kalaeloa (Daly and
Magnacca 2003, p. 217; Magnacca
2005b, in litt., p. 2).
Habitat destruction and modification
by urbanization and land use
conversion lead to fragmentation of, and
eventual loss, of foraging and nesting
areas used by Hylaeus assimulans.
Habitat destruction and modification by
nonnative plants (Asystasia gangetica
(Chinese violet), Atriplex semibaccata,
Cenchrus ciliaris (buffelgrass), Chloris
barbata (swollen fingergrass), Digitaria
insularis (sourgrass), Leucaena
leucocephala, Melinis minutiflora,
Pluchea indica (Indian fleabane), P.
carolinensis, Prosopis pallida, Schinus
terebinthifolius, and Verbesina
encelioides (golden crown-beard)
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adversely impact native plant species by
modifying the availability of light,
altering soil-water regimes, modifying
nutrient cycling, altering the fire
characteristics, and ultimately
converting native dominated plant
communities to nonnative plant
communities, and results in removal of
food sources and nesting sites for H.
assimulans (Xerces Society 2009, p. 21;
76 FR 55170, September 6, 2011, p.
55184). Habitat modification and
destruction by nonnative animals such
as feral pigs, goats, axis deer, and cattle
is considered one of the primary factors
underlying destruction of native
vegetation in the Hawaiian Islands, and
these habitat changes also remove food
sources and nesting sites of H.
assimulans (Stone 1985, pp. 262–263;
Cuddihy and Stone 1990, pp. 60–66,
73). Fire is a threat to H. assimulans, as
it destroys native plant communities on
which the species depends, and opens
habitat for increased invasion by
nonnative plants. Because of the greater
frequency, intensity, and duration of
fires that have resulted from the human
alteration of landscapes and the
introduction of nonnative plants,
especially grasses, fires are now more
destructive to the coastal and lowland
dry ecosystems (Brown and Smith 2000,
p. 172), and a single grass-fueled fire
often kills most native trees and shrubs
in the area (D’Antonio and Vitousek
1992, p. 74) and could destroy food and
nesting resources for H. assimulans. The
numbers of wildfires and the acreages
involved are increasing in the main
Hawaiian Islands; however, their
occurrences and locations are
unpredictable, and could affect habitat
for yellow-faced bees at any time (Gima
1998, in litt.; County of Maui 2009, ch.
3, p. 3; Hamilton 2009, in litt.; Honolulu
Advertiser 2010, in litt.; Pacific Disaster
Center 2011, in litt.). Random, naturally
occurring events such as hurricanes,
tsunami, and drought modify and
destroy habitat of H. assimulans by
creating disturbed areas conducive to
invasion by nonnative plants,
eliminating food and nesting sources
(Kitayama and Mueller-Dombois 1995,
p. 671; Businger 1998, pp. 1–2;
Magnacca 2015, in litt.). Predation by
nonnative ants (the big-headed ant, the
yellow crazy ant, Solenopsis papuana,
and S. geminata) on Hylaeus egg, larvae,
and pupal stages is a threat to H.
assimulans; additionally, ants compete
with H. assimulans for their nectar food
source (Howarth 1985, p. 155; Hopper et
al. 1996, p. 9; Holway et al. 2002, pp.
188, 209; Daly and Magnacca 2003, p. 9;
Lach 2008, p. 155). Predation by
nonnative western yellow jacket wasps
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is a threat to H. assimulans because the
wasp is an aggressive, generalist
predator, and occurs in great numbers in
many habitat types, from sea level to
over 8,000 ft (2,450 m), including areas
where H. assimulans and other yellowfaced bees occur (Gambino et al. 1987,
p. 169). Existing regulatory mechanisms
and agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees (honeybees, carpenter bees, sweat
bees, and alien Hylaeus bees) for nectar
and pollen is a threat to H. assimulans
(Magnacca 2007, p. 188; Graham 2015,
in litt; Magnacca 2015, in litt.). The
small number of populations and
individuals of H. assimulans makes this
species more vulnerable to extinction
because of the higher risks from genetic
bottlenecks, random demographic
fluctuations, and localized catastrophes
such as hurricanes and drought (Daly
and Magnacca 2003, p. 3; Magnacca
2007, p. 173). Although we cannot
predict the timing, extent, or magnitude
of specific impacts, we do expect the
effects of climate change to exacerbate
the threats to H. assimulans described
above.
The remaining populations of Hylaeus
assimulans and its habitat are at risk.
The known individuals are restricted to
5 locations: 2 on Maui, 1 on Kahoolawe,
and 2 on Lanai, and is likely extirpated
from Oahu. This species continues to be
negatively affected by habitat
destruction and modification by
urbanization and land-use conversion,
and by habitat destruction and removal
of food and nesting sites by nonnative
ungulates and nonnative plants. Habitat
destruction by fire is a threat. Randomly
occurring events such as hurricanes and
drought modify habitat and remove food
and nesting sources for H. assimulans.
Predation by nonnative ants and wasps
is a threat. Existing regulatory
mechanisms and agency policies do not
address the primary threats to the
yellow-faced bees and their habitat from
nonnative ungulates. Competition with
nonnative bees for food and nesting
sites is a threat. The small number of
remaining populations limits this
species’ ability to adapt to
environmental changes. The effects of
climate change are likely to further
exacerbate these threats. Because of
these threats, we find that H. assimulans
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
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Yellow Faced Bee (Hylaeus facilis)
Historically, Hylaeus facilis was
known from Maui, Lanai, Molokai, and
Oahu, in dry shrubland to wet forest
from sea level to 3,000 ft (1,000 m)
(Gagne and Cuddihy 1999, p. 93; Daly
and Magnacca 2003, pp. 81, 83). Perkins
(1899, p. 77) remarked H. facilis was
among the most common and
widespread Hylaeus species on Oahu,
Maui, Lanai, and Molokai (Magnacca
2007, p. 183). Although the species was
collected in a wide range of habitat
types, it likely prefers dry to mesic
forest and shrubland (Magnacca 2005c,
in litt., p. 2), which are increasingly rare
and patchily distributed habitats (Smith
1985, pp. 227–233; Juvik and Juvik
1998, p. 124; Gagne and Cuddihy 1999,
pp. 66–67, 75; Magnacca 2005c, in litt.,
p. 2). Researchers believe the wet forest
site on Oahu where H. facilis was
observed likely had a more open
understory (more mesic conditions), and
represented an outlier or residual
population (Perkins 1899, p.76; Liebherr
and Polhemus 1997; p. 347). Hylaeus
facilis has almost entirely disappeared
from most of its historical range (Maui,
coastal and lowland mesic; Lanai,
lowland dry and lowland mesic; and
Oahu, coastal and lowland dry) (Daly
and Magnacca 2003, p. 7; Magnacca
2007, p. 183). Between 1998 and 2006,
39 sites on Maui, Lanai, Molokai, and
Oahu were surveyed, including 13
historical sites. Hylaeus facilis was
absent from all 13 locations (Magnacca
2007, p. 183) and was not observed at
26 additional sites with potentially
suitable habitat (Daly and Magnacca
2003, pp. 7, 81–82; Magnacca 2007, p.
183). Likely extirpated from Lanai, H.
facilis is currently known from only two
locations, one on Molokai in the coastal
ecosystem, and one on Oahu in the
coastal ecosystem (Daly and Magnacca
2003, pp. 81–82; Magnacca 2005c, in
litt., p. 2). In addition, in 1990, a single
individual was collected on Maui near
Makawao at 1,500 ft (460 m); however,
this site is urbanized and devoid of
native plants, and it is likely this
collection was a vagrant individual.
Habitat destruction and modification
by urbanization and land use
conversion lead to fragmentation of, and
eventual loss of, foraging and nesting
areas used by Hylaeus facilis. Habitat
destruction and modification by
nonnative plants adversely impacts
native plant species by modifying the
availability of light, altering soil-water
regimes, modifying nutrient cycling,
altering the fire characteristics, and
ultimately converting native dominated
plant communities to nonnative plant
communities, and results in removal of
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food sources and nesting sites for the H.
facilis. In addition to the nonnative
plant species noted above that modify
and destroy habitat of H. assimulans,
Urochloa mutica, Prosopis pallida,
Psidium cattleianum, and Rubus spp.
are noted to negatively affect the habitat
of H. facilis (Cuddihy and Stone 1990,
p. 105; Hawaii Division of Forestry and
Wildlife (DOFAW) 2007, pp. 20–22).
Habitat modification and destruction by
nonnative animals such as feral pigs,
goats, axis deer, and cattle is considered
one of the primary factors underlying
destruction of native vegetation in the
Hawaiian Islands, and these habitat
changes also remove food sources and
nesting sites for H. facilis (Stone 1985,
pp. 262–263; Cuddihy and Stone 1990,
pp. 60–66, 73). Fire is a threat to H.
facilis, as it destroys native plant
communities on which the species
depends, and opens habitat for
increased invasion by nonnative plants.
Because of the greater frequency,
intensity, and duration of fires that have
resulted from the human alteration of
landscapes and the introduction of
nonnative plants, especially grasses,
fires are now more destructive to the
coastal and lowland dry ecosystems
(Brown and Smith 2000, p. 172), and a
single grass-fueled fire often kills most
native trees and shrubs in the area
(D’Antonio and Vitousek 1992, p. 74)
and could destroy food and nesting
resources for H. facilis. The numbers of
wildfires and the acreages involved are
increasing in the main Hawaiian
Islands; however, their occurrences and
locations are unpredictable, and could
affect habitat for yellow-faced bees at
any time (Gima 1998, in litt.; County of
Maui 2009, ch. 3, p. 3; Hamilton 2009,
in litt.; Honolulu Advertiser 2010, in
litt.; Pacific Disaster Center 2011, in
litt.). Random, naturally occurring
events such as hurricanes, tsunami, and
drought modify and destroy habitat of
H. facilis by creating disturbed areas
conducive to invasion by nonnative
plants, eliminating food and nesting
resources (Kitayama and MuellerDombois 1995, p. 671; Businger 1998,
pp. 1–2; Magnacca 2015, in litt.).
Predation by nonnative ants (the bigheaded ant, the yellow crazy ant,
Solenopsis papuana, and S. geminata)
on Hylaeus egg, larvae, and pupal stages
is a threat to H. facilis; additionally, ants
compete with H. facilis for their nectar
food source (Howarth 1985, p. 155;
Hopper et al. 1996, p. 9; Holway et al.
2002, pp. 188, 209; Daly and Magnacca
2003, p. 9; Lach 2008, p. 155). Predation
by nonnative western yellow jacket
wasps is a threat to H. facilis because
the wasp is an aggressive, generalist
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predator, and occurs in great numbers in
many habitat types, from sea level to
over 8,000 ft (2,450 m), including areas
where H. facilis and other yellow-faced
bees occur (Gambino et al. 1987, p. 169).
Existing regulatory mechanisms and
agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees (honeybees, carpenter bees, sweat
bees, and alien Hylaeus bees) for nectar
and pollen is a threat to H. facilis
(Magnacca 2007, p. 188; Magnacca 2015,
in litt.). The small number of
populations and individuals of H. facilis
makes this species more vulnerable to
extinction because of the higher risks
from genetic bottlenecks, random
demographic fluctuations, and localized
catastrophes such as hurricanes and
drought (Daly and Magnacca 2003, p. 3;
Magnacca 2007, p. 173). Although we
cannot predict the timing, extent, or
magnitude of specific impacts, we do
expect the effects of climate change to
exacerbate the threats to H. facilis
described above.
The remaining populations of Hylaeus
facilis and its habitat are at risk. The
known individuals are restricted to one
location on Molokai and one location on
Oahu, and continue to be negatively
affected by habitat destruction and
modification by urbanization and landuse conversion, and by habitat
destruction and removal of food and
nesting sites by nonnative ungulates and
nonnative plants. Habitat destruction by
fire is a threat. Randomly occurring
events such as hurricanes and drought
modify habitat and remove food and
nesting sources for H. facilis. Predation
by nonnative ants and wasps is a threat.
Existing regulatory mechanisms and
agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees for food and nesting sites is a
threat. The small number of remaining
populations limits this species’ ability
to adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that H. facilis
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Yellow-Faced Bee (Hylaeus hilaris)
Historically, Hylaeus hilaris was
known from coastal habitat on Maui,
Lanai, and Molokai; and lowland dry
habitat on Maui. It is believed to have
occurred along much of the coast of
these islands because its primary hosts,
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H. anthracinus, H. assimulans, and H.
longiceps likely occurred throughout
this habitat. First collected on Maui in
1879, H. hilaris has only been collected
twice in the last 100 years. Hylaeus
hilaris was absent from three of its
historical population sites revisited by
researchers between 1998 and 2006
(Magnacca 2007, p. 181). It was also not
observed in 2003 at 10 additional sites
with potentially suitable habitat (Daly
and Magnacca 2003, pp. 103, 106).
Currently, the only known population of
H. hilaris is located on Molokai, in the
coastal ecosystem (Daly and Magnacca
2003, pp. 103, 106; Magnacca 2005d, in
litt., p. 2; Magnacca 2007, p. 181).
Because Hylaeus hilaris is an obligate
parasite on H. anthracinus, H.
assimulans, and H. longiceps, its
occurrences are determined by the
remaining populations of these three
species. Habitat destruction and
modification by urbanization and land
use conversion leads to fragmentation
of, and eventual loss of, foraging and
nesting areas of H. hilaris, and of those
Hylaeus species that H. hilaris is
dependent upon. Habitat destruction
and modification by nonnative plants
adversely impacts native plant species
by modifying the availability of light,
altering soil-water regimes, modifying
nutrient cycling, altering the fire
characteristics, and ultimately
converting native dominated plant
communities to nonnative plant
communities, and results in removal of
food sources and nesting sites for the
Hylaeus species that H. hilaris is
dependent upon. Nonnative plant
species that modify and destroy habitat
of H. hilaris are noted in the description
for H. assimulans, above. Habitat
modification and destruction by
nonnative animals such as feral pigs,
goats, axis deer, and cattle is considered
one of the primary factors underlying
destruction of native vegetation in the
Hawaiian Islands, and these habitat
changes also remove food sources and
nesting sites for the host species of H.
hilaris (Stone 1985, pp. 262–263;
Cuddihy and Stone 1990, pp. 60–66,
73). Fire is a threat to H. hilaris, as it
destroys native plant communities, and
opens habitat for increased invasion by
nonnative plants. Because of the greater
frequency, intensity, and duration of
fires that have resulted from the human
alteration of landscapes and the
introduction of nonnative plants,
especially grasses, fires are now more
destructive to the coastal and lowland
dry ecosystems (Brown and Smith 2000,
p. 172), and a single grass-fueled fire
often kills most native trees and shrubs
in the area (D’Antonio and Vitousek
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1992, p. 74) and could destroy food and
nesting resources for H. hilaris. The
numbers of wildfires and the acreages
involved are increasing in the main
Hawaiian Islands; however, their
occurrences and locations are
unpredictable, and could affect habitat
for yellow-faced bees at any time (Gima
1998, in litt.; County of Maui 2009, ch.
3, p. 3; Hamilton 2009, in litt.; Honolulu
Advertiser 2010, in litt.; Pacific Disaster
Center 2011, in litt.). Random, naturally
occurring events such as hurricanes,
tsunami, and drought can modify and
destroy habitat of H. hilaris by creating
disturbed areas conducive to invasion
by nonnative plants, eliminating food
and nesting sources of its host species
(Kitayama and Mueller-Dombois 1995,
p. 671; Businger 1998, pp. 1–2;
Magnacca 2015, in litt.). Predation by
nonnative ants (the big-headed ant, the
long-legged ant, Solenopsis papuana,
and S. geminata) on Hylaeus egg, larvae,
and pupal stages is a threat to H. hilaris;
additionally, ants compete with the
yellow-faced bees that H. hilaris is
dependent on for their food resources
(Howarth 1985, p. 155; Hopper et al.
1996, p. 9; Holway et al. 2002, pp. 188,
209; Daly and Magnacca 2003, p. 9;
Lach 2008, p. 155). Predation by
nonnative western yellow jacket wasps
is a threat to H. hilaris because the wasp
is an aggressive, generalist predator, and
occurs in great numbers in many habitat
types, from sea level to over 8,000 ft
(2,450 m), including areas where H.
hilaris and other yellow-faced bees
occur (Gambino et al. 1987, p. 169).
Existing regulatory mechanisms and
agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees (honeybees, carpenter bees, sweat
bees, and alien Hylaeus bees) for nectar
and pollen is a threat to the host yellowfaced bees of H. hilaris (Magnacca 2007,
p. 188; Graham 2015, in litt.; Magnacca
2015, in litt.). The small number of
populations and individuals of H.
hilaris makes this species more
vulnerable to extinction because of the
higher risks from genetic bottlenecks,
random demographic fluctuations, and
localized catastrophes such as
hurricanes and drought (Daly and
Magnacca 2003, p. 3; Magnacca 2007, p.
173). Although we cannot predict the
timing, extent, or magnitude of specific
impacts, we do expect the effects of
climate change to exacerbate the threats
to H. hilaris described above.
The remaining populations of Hylaeus
hilaris and its habitat are at risk. There
is one known occurrence on Molokai.
Hylaeus hilaris and its host species
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continue to be negatively affected by
habitat destruction and modification by
urbanization and land-use conversion,
and by habitat destruction and removal
of food and nesting sites (for host
species) by nonnative ungulates and
nonnative plants. Habitat destruction by
fire is a threat. Randomly occurring
events such as hurricanes and drought
modify habitat and remove food and
nesting sources for H. hilaris and its
host species. Predation by nonnative
ants and wasps is a threat. Existing
regulatory mechanisms and agency
policies do not address the primary
threats to the yellow-faced bees and
their habitat from nonnative ungulates.
Competition with nonnative bees for
food and nesting sites is a threat. The
small number of remaining populations
limits this species’ ability to adapt to
environmental changes, especially
because it is an obligate parasite of other
rare Hylaeus bees. Because of these
threats, we find that H. hilaris is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Yellow-Faced Bee (Hylaeus kuakea)
Because the first collection of Hylaeus
kuakea was not made until 1997, its
historical range is unknown (Magnacca
2005e, in litt., p. 2; Magnacca 2007, p.
184). Phylogenetically, H. kuakea
belongs in a species-group primarily
including species inhabiting mesic
forests (Magnacca and Danforth 2006, p.
405). Only four individuals (all males)
have been collected from two different
sites in the Waianae Mountains of Oahu
in the lowland mesic ecosystem
(Magnacca 2007, p. 184). The species
has never been collected in any other
habitat type or area, including some
sites that have been more thoroughly
surveyed (Magnacca 2011, in litt.). Not
all potentially suitable habitat has been
surveyed due to the remote and rugged
locations, small size, rareness, and
distant spacing among large areas of
nonnative forest (Smith 1985, pp. 227–
233; Juvik and Juvik 1998, p. 124;
Wagner et al. 1999, pp. 66–67, 75).
Habitat destruction and modification
by feral pigs leads to fragmentation of,
and eventual loss of, foraging and
nesting areas of Hylaeus kuakea. Habitat
destruction and modification by
nonnative plants adversely impacts
native plant species by modifying the
availability of light, altering soil-water
regimes, modifying nutrient cycling,
altering the fire characteristics, and
ultimately converting native dominated
plant communities to nonnative plant
communities, and results in removal of
PO 00000
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67815
food sources and nesting sites for H.
kuakea. Nonnative plant species that
modify and destroy habitat of H. kuakea
are noted in the descriptions for H.
assimulans and H. facilis, above. Fire is
a threat to H. kuakea because it destroys
native plant communities and opens
habitat for increased invasion by
nonnative plants. Because of the greater
frequency, intensity, and duration of
fires that have resulted from the human
alteration of landscapes and the
introduction of nonnative plants,
especially grasses, fires are now more
destructive, including in lowland mesic
areas (Brown and Smith 2000, p. 172),
and a single grass-fueled fire often kills
most native trees and shrubs in the area
(D’Antonio and Vitousek 1992, p. 74)
and could destroy food and nesting
resources for H. kuakea. The numbers of
wildfires and the acreages involved are
increasing in the main Hawaiian
Islands; however, their occurrences and
locations are unpredictable, and could
affect habitat for yellow-faced bees at
any time (Gima 1998, in litt.; County of
Maui 2009, ch. 3, p. 3; Hamilton 2009,
in litt.; Honolulu Advertiser 2010, in
litt.; Pacific Disaster Center 2011, in
litt.). Random, naturally occurring
events such as hurricanes and drought
can modify and destroy habitat of H.
kuakea by creating disturbed areas
conducive to invasion by nonnative
plants, eliminating food and nesting
resources (Kitayama and MuellerDombois 1995, p. 671; Businger 1998,
pp. 1–2). Predation by nonnative ants
(the big-headed ant, the long-legged ant,
Solenopsis papuana, and S. geminata)
on Hylaeus egg, larvae, and pupal stages
is a threat to H. kuakea; additionally,
ants compete with H. kuakea for their
nectar food source (Howarth 1985, p.
155; Hopper et al. 1996, p. 9; Holway et
al. 2002, pp. 188, 209; Daly and
Magnacca 2003, p. 9; Lach 2008, p. 155).
Predation by nonnative western yellow
jacket wasps is a threat to H. kuakea
because the wasp is an aggressive,
generalist predator, and occurs in great
numbers in many habitat types, from sea
level to over 8,000 ft (2,450 m),
including areas where H. kuakea and
other yellow-faced bees occur (Gambino
et al. 1987, p. 169). Existing regulatory
mechanisms and agency policies do not
address the primary threats to the
yellow-faced bees and their habitat from
nonnative ungulates. Competition with
nonnative bees (honeybees, carpenter
bees, sweat bees, and alien Hylaeus
bees) for nectar and pollen is a threat to
H. kuakea (Magnacca 2007, p. 188;
Graham 2015, in litt.; Magnacca 2015, in
litt.). The small number of populations
and individuals of H. kuakea makes this
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species more vulnerable to extinction
because of the higher risks from genetic
bottlenecks, random demographic
fluctuations, and localized catastrophes
such as hurricanes and drought (Daly
and Magnacca 2003, p. 3; Magnacca
2007, p. 173). Although we cannot
predict the timing, extent, or magnitude
of specific impacts, we do expect the
effects of climate change to exacerbate
the threats to H. kuakea described
above.
The remaining populations of Hylaeus
kuakea and its habitat are at risk. The
known individuals are restricted to
mesic forest in one area of one island
(Oahu), and continue to be negatively
affected by habitat destruction and
removal of food and nesting sites by
nonnative ungulates and nonnative
plants. Habitat destruction by fire is a
threat. Randomly occurring events such
as hurricanes and drought modify
habitat and remove food and nesting
sources for H. kuakea. Predation by
nonnative ants and wasps is a threat.
Existing regulatory mechanisms and
agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees for food and nesting sites is a
threat. The small number of remaining
populations limits this species’ ability
to adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that H. kuakea
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Yellow-faced bee (Hylaeus longiceps)
Hylaeus longiceps is historically
known from coastal and lowland dry
shrubland habitat up to 2,000 ft (610 m)
in numerous locations on the islands of
Maui, Lanai, Molokai, and Oahu.
Perkins (1899, p. 98) noted H. longiceps
was locally abundant, and probably
occurred throughout much of the
leeward and lowland areas on these
islands. Hylaeus longiceps is now
restricted to small populations in
patches of coastal and lowland dry
habitat on the Maui, Lanai, Molokai,
and Oahu (Magnacca 2005f, in litt., p. 2;
Magnacca and King 2013, pp. 13, 16).
Twenty-five sites that were either
historical collecting localities or
contained potentially suitable habitat
for this species were surveyed between
1997 and 2008 (Magnacca and King
2013, p. 16). Hylaeus longiceps was
observed at only seven of the surveyed
sites: Three sites on Lanai (in the coastal
and lowland dry ecosystems), two sites
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on Oahu (in the coastal ecosystem), and
one site on each of the islands of Maui
(in the coastal ecosystem) and Molokai
(in the coastal ecosystem) (Daly and
Magnacca 2003, p. 135; Magnacca and
King 2013, pp. 11–12).
Most of the coastal and lowland
habitat of Hylaeus longiceps has been
developed or degraded, and is no longer
suitable (Liebherr and Polhemus 1997,
pp. 346–347; Magnacca 2007, pp. 186–
188). Habitat destruction and
modification by axis deer (Lanai) and
urbanization (Maui and Molokai) leads
to fragmentation of, and eventual loss
of, foraging and nesting areas of H.
longiceps (Daly and Magnacca 2003, pp.
217–229). Habitat modification and
destruction by human impacts in areas
accessible by four-wheel drive vehicles
on Lanai is a threat because these
vehicles can destroy plants used as food
sources and destroy ground nesting sites
for H. longiceps (Daly and Magnacca
2003, p. 135). Habitat destruction and
modification by nonnative plants
adversely affects native plant species
used by H. longiceps as a food source by
modifying the availability of light,
altering soil-water regimes, modifying
nutrient cycling, altering the fire
characteristics, and ultimately
converting native-dominated plant
communities to nonnative plant
communities. Nonnative plant species
that modify and destroy habitat of H.
longiceps are noted in the descriptions
for H. assimulans and H. facilis, above.
Fire is a threat to H. longiceps because
it destroys native plant communities,
and opens habitat for increased invasion
by nonnative plants. Because of the
greater frequency, intensity, and
duration of fires that have resulted from
the human alteration of landscapes and
the introduction of nonnative plants,
especially grasses, fires are now more
destructive to the coastal and lowland
dry ecosystems (Brown and Smith 2000,
p. 172), and a single grass-fueled fire
often kills most native trees and shrubs
in the area (D’Antonio and Vitousek
1992, p. 74) and could destroy food and
nesting resources for H. longiceps. The
numbers of wildfires and the acreages
involved are increasing in the main
Hawaiian Islands; however, their
occurrences and locations are
unpredictable, and could affect habitat
for yellow-faced bees at any time (Gima
1998, in litt.; County of Maui 2009, ch.
3, p. 3; Hamilton 2009, in litt.; Honolulu
Advertiser 2010, in litt.; Pacific Disaster
Center 2011, in litt.). Random, naturally
occurring events such as hurricanes,
tsunami, and drought modify and
destroy habitat of H. longiceps by
creating disturbed areas conducive to
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invasion by nonnative plants,
eliminating food and nesting resources
(Kitayama and Mueller-Dombois 1995,
p. 671; Businger 1998, pp. 1–2;
Magnacca 2015, in litt.). Predation by,
and competition for food sources, by
nonnative ants and the nonnative
western yellow jacket wasp is a threat
to H. longiceps (see H. kuakea, above)
(Gambino et al. 1987, p. 169; Howarth
1985, p. 155; Hopper et al. 1996, p. 9;
Holway et al. 2002, pp. 188, 209; Daly
and Magnacca 2003, p. 9; Lach 2008, p.
155). Existing regulatory mechanisms
and agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees (honeybees, carpenter bees, sweat
bees, and alien Hylaeus bees) for nectar
and pollen is a threat to H. longiceps
(Magnacca 2007, p. 188; Graham 2015,
in litt.; Magnacca 2015, in litt.). The
small number of populations and
individuals of H. longiceps makes this
species more vulnerable to extinction
because of the higher risks from genetic
bottlenecks, random demographic
fluctuations, and localized catastrophes
such as hurricanes and drought (Daly
and Magnacca 2003, p. 3; Magnacca
2007, p. 173). Although we cannot
predict the timing, extent, or magnitude
of specific impacts, we do expect the
effects of climate change to exacerbate
the threats to H. longiceps described
above.
The remaining population of Hylaeus
longiceps and its habitat are at risk. The
known individuals are restricted to
seven locations, three on Lanai, two on
Oahu, and one each on Maui and
Molokai, and continue to be negatively
affected by habitat destruction and
modification by urbanization and landuse conversion, by habitat destruction
and removal of food and nesting sites by
nonnative ungulates and nonnative
plants, and by recreational use vehicles
on Lanai. Habitat destruction by fire is
a threat. Randomly occurring events
such as hurricanes and drought may
modify habitat and remove food and
nesting sources for H. longiceps.
Predation by nonnative ants and wasps
is a threat. Existing regulatory
mechanisms and agency policies do not
address the primary threats to the
yellow-faced bees and their habitat from
nonnative ungulates. Competition with
nonnative bees for food and nesting
sites is a threat. The small number of
remaining populations limits this
species’ ability to adapt to
environmental changes. Because of
these threats, we find that H. longiceps
is endangered throughout all of its
range, and, therefore, find that it is
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threat to H. mana, as it destroys native
plant communities on which the species
depends, and opens habitat for
increased invasion by nonnative plants.
Yellow-Faced Bee (Hylaeus mana)
Because of the greater frequency,
Hylaeus mana is known only from
intensity, and duration of fires that have
lowland mesic forest dominated by
resulted from the human alteration of
native Acacia koa in the Koolau
landscapes and the introduction of
Mountains of Oahu, at 1,400 ft (430 m).
nonnative plants, especially grasses,
Few other Hylaeus species have been
fires are now more destructive,
found in this type of forest on Oahu
including in lowland mesic ecosystems
(Daly and Magnacca 2003, p. 138). This
(Brown and Smith 2000, p. 172). A
type of native forest is increasingly rare
single grass-fueled fire often kills most
and patchily distributed because of
native trees and shrubs in the area
competition and encroachment into
(D’Antonio and Vitousek 1992, p. 74)
habitat by nonnative plants (Smith
and could destroy food and nesting
1985, pp. 227–233; Juvik and Juvik
resources for H. mana. The numbers of
1998, p. 124; Wagner et al. 1999, pp. 66– wildfires and the acreages involved are
67, 75). Decline of this forest type could increasing in the main Hawaiian
lead to decline in populations and
Islands; however, their occurrences and
numbers of H. mana. Three additional
locations are unpredictable, and could
population sites were discovered on
affect habitat for yellow-faced bees at
Oahu in 2012, including a new
any time (Gima 1998, in litt.; County of
observation of the species at the original Maui 2009, ch. 3, p. 3; Hamilton 2009,
site (Magnacca and King 2013, pp. 17–
in litt.; Honolulu Advertiser 2010, in
18). The three new sites are within a
litt.; Pacific Disaster Center 2011, in
narrow range of lowland mesic forest at
litt.). Random, naturally occurring
1,400 ft (430 m), bordered by nonnative
events such as hurricanes and drought
plant habitat at lower elevations and
can modify and destroy habitat of H.
wetter native forest habitat above
mana by creating disturbed areas
(Magnacca and King 2013, pp. 17–18).
conducive to invasion by nonnative
Hylaeus mana was most often observed
plants (Kitayama and Mueller-Dombois
on Santalum freycinetianum var.
1995, p. 671; Businger 1998, pp. 1–2).
freycinetianum, which suggests that H.
Predation and competition for food
mana may be closely associated with
sources by nonnative ants and the
this plant species (Magnacca and King
nonnative western yellow jacket wasp
2013, p. 18). Additional surveys may
are threats to H. mana (see H. kuakea,
reveal more populations; however, the
above) (Howarth 1985, p. 155; Gambino
extreme rarity of this species, its
absence from many survey sites, the fact et al. 1987, p. 169; Hopper et al. 1996,
p. 9; Holway et al. 2002, pp. 188, 209;
that it was not discovered until very
Daly and Magnacca 2003, p. 9; Lach
recently, and the limited range of its
2008, p. 155). Existing regulatory
possible host plant, all suggest that few
populations remain (Magnacca 2005g, in mechanisms and agency policies do not
address the primary threats to the
litt., p. 2; Magnacca and King 2013, pp.
yellow-faced bees and their habitat from
17–18).
nonnative ungulates. Competition with
Habitat destruction and modification
nonnative bees (honeybees, carpenter
by feral pigs leads to fragmentation of,
bees, sweat bees, and alien Hylaeus
and eventual loss of, foraging and
bees) for nectar and pollen is a threat to
nesting areas of Hylaeus mana (Daly
H. mana (Magnacca 2007, p. 188;
and Magnacca 2003, pp. 217–229).
Habitat destruction and modification by Graham 2015, in litt.; Magnacca 2015, in
litt.). The small number of populations
nonnative plants adversely impacts
native plant species used by H. mana as and individuals of H. mana makes this
species more vulnerable to extinction
a food source by modifying the
because of the higher risks from genetic
availability of light, altering soil-water
bottlenecks, random demographic
regimes, modifying nutrient cycling,
fluctuations, and localized catastrophes
altering the fire characteristics, and
ultimately converting native dominated such as hurricanes and drought (Daly
and Magnacca 2003, p. 3; Magnacca
plant communities to nonnative plant
2007, p. 173). Although we cannot
communities. Nonnative plant species
predict the timing, extent, or magnitude
that modify and destroy habitat of H.
of specific impacts, we do expect the
mana are noted in the descriptions for
H. assimulans and H. facilis, above, and effects of climate change to exacerbate
the threats to H. mana described above.
can outcompete native canopy species
The remaining populations of Hylaeus
such as Acacia koa, the known
preferred native canopy type of H. mana mana and its habitat are at risk. The
known individuals are restricted to
(GISD 2011, in litt.; State of Hawaii
three locations of native koa forest on
2013, in litt. (S.C.R. No. 74)). Fire is a
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unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
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Oahu, and continue to be negatively
affected by habitat destruction and
removal of food and nesting sites by
nonnative ungulates and nonnative
plants. Habitat destruction by fire is a
threat. Randomly occurring events such
as hurricanes and drought may modify
habitat and remove food and nesting
sources for H. mana. Predation by
nonnative ants and wasps is a threat.
Existing regulatory mechanisms and
agency policies do not address the
primary threats to the yellow-faced bees
and their habitat from nonnative
ungulates. Competition with nonnative
bees for food and nesting sites is a
threat. The small number of remaining
populations limits this species’ ability
to adapt to environmental changes. The
effects of climate change are likely to
further exacerbate these threats. Because
of these threats, we find that H. mana
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Orangeblack Hawaiian Damselfly
(Megalagrion xanthomelas)
The orangeblack Hawaiian damselfly
was once Hawaii’s most abundant
damselfly species likely because of its
ability to use a variety of aquatic
habitats for breeding sites. Historically,
the orangeblack Hawaiian damselfly
probably occurred on all of the main
Hawaiian Islands (except Kahoolawe) in
suitable aquatic habitat within the
anchialine pool, coastal, lowland dry,
and lowland mesic ecosystems (Perkins
1913, p. clxxviii; Zimmerman 1948, p.
379; Polhemus 1996, p. 30). Its
historical range on Kauai is unknown.
On Oahu, it was recorded from
Honolulu, Kaimuki, Koko Head, Pearl
City, Waialua, the Waianae Mountains,
and Waianae (Polhemus 1996, pp. 31,
33). On Molokai, it was known from
Kainalu, Meyer’s Lake (Kalaupapa
Peninsula), Kaunakakai, Mapulehu, and
Palaau (Polhemus 1996, pp. 33–41). On
Lanai, small populations occurred on
Maunalei Gulch, and in ephemeral
coastal ponds at the mouth of Maunalei
Gulch drainage, at Keomuku, and in a
mixohaline (brackish water) habitat at
Lopa (Polhemus 1996, pp. 37–41; HBMP
2010). On Maui, this species was
recorded from an unspecified locality in
the west Maui Mountains (Polhemus
1996, pp. 41–42; Polhemus et al. 1999,
pp. 27–29). On Hawaii Island, it was
known from Hilo, Kona, and Naalehu
(Polhemus 1996, pp. 42–47).
Currently, the orangeblack Hawaiian
damselfly occurs on Oahu, Molokai,
Lanai, Maui, and Hawaii Island. In
1994, on Oahu, a very small population
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was discovered in pools of an
intermittent stream (Englund 2001, p.
256). On Molokai, populations occur at
the mouths of two streams, and in
wetlands on the south coast (Polhemus
1996, p. 47). On Lanai, a large
population occurs in an artificial pond
(Polhemus 1996, p. 47). The species is
present on west Maui at a stream and
near anchialine pools on east Maui
(Polhemus et al. 1999, p. 29). Several
large populations exist in coastal
wetlands on Hawaii Island at 14
locations (Polhemus 1996, pp. 42–47;
Orlando 2015, in litt.). The species is
believed to be extirpated from Kauai
(Asquith and Polhemus 1996, p. 91).
Past and present land use and water
management practices, including
agriculture, urban development, ground
water development, and destruction of
perched aquifer and surface water
resources, and feral ungulates (pigs,
goats, axis deer), modify and destroy
habitat of the orangeblack Hawaiian
damselfly (Harris et al. 1993, pp. 9–13;
Meier et al. 1993, pp. 181–183).
Nonnative plant species such as
Urochloa mutica form dense, monotypic
stands that can completely eliminate
any open water habitat of the
orangeblack Hawaiian damselfly (Smith
1985, p. 186). Stochastic events such as
drought, flooding, and hurricanes can
also modify and destroy habitat, and kill
individuals. Predation of the
orangeblack Hawaiian damselfly by
nonnative fish and nonnative aquatic
invertebrates on the orangeblack
Hawaiian damselfly is a significant
threat; predation by Jackson’s
chameleons (Trioceros jacksonii) may
occur as well (Sailer 2015, in litt.).
Hawaiian damselflies evolved with few,
if any, predatory fish, and the reduced
defensive and evasive behaviors of most
of the fully aquatic species, including
the orangeblack Hawaiian damselfly,
makes them particularly vulnerable to
predation by nonnative fish (Englund
1999, pp. 225–225, 235; Haines 2015, in
litt.). The damselfly is not observed in
any bodies of water that support
nonnative fish (Henrickson 1988, p. 183;
McPeek 1990a, pp. 92–96). Nonnative
backswimmers (aquatic true bugs;
Heteroptera) are voracious predators
and frequently feed on prey much larger
than themselves, such as tadpoles, small
fish, and other aquatic invertebrates and
may be a potential threat to damselfly’s
aquatic larvae (naiads) (Borror et al.
1989, p. 296). In addition, the nonnative
bullfrog (Rana catesbeiana, Lithobates
catesbeianus), found in ponds and along
streams, is a generalist predator, and
eats insects and crustaceans as well as
a wide variety of small vertebrates (Bury
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and Whelan 1985, p. 4). Predation by
the bullfrog is a threat to the
orangeblack Hawaiian damselfly
(Englund et al. 2007, pp. 215, 219;
Haines 2015, in litt.). Also, caddisflies
(Trichoptera spp.) compete with native
aquatic invertebrates for resources and
space (Flint et al. 2003, p. 38; Haines
2015, in litt.) and reduce prey
abundance for orangeblack Hawaiian
damselfly larvae.
Hawaii State law (State Water Code)
does not provide for permanent or
minimal instream flow standards, and
channel modifications or revisions to
flow standards can be undertaken at any
time by the Water Commission, without
regard for changes that degrade or
destroy habitat, food resources, or
aquatic life stages of the orangeblack
Hawaiian damselfly. Therefore, existing
regulatory mechansims do not
adequately address the threat of
modification and destruction of the
aquatic habitat of the orangeblack
Hawaiian damselfly (Hawaii
Administrative Rule (HAR)-State Water
Code, title 13, chapter 169–36; Tango
2010, in litt.).
The remaining populations and
habitat of the orangeblack Hawaiian
damselfly are at risk; numbers are
decreasing on Oahu, Molokai, Lanai,
Maui, and Hawaii Island, and both the
species and its habitat continue to be
negatively affected by modification and
destruction by development and water
management practices, drought, feral
ungulates, and by nonnative plants,
combined with predation by nonnative
fish and other nonnative vertebrates.
Competition with caddisflies is a
potential threat to the orangeblack
Hawaiian damselfly. The orangeblack
damselfly was once the most common
Hawaiian damselfly in the State, and
occurred in any suitable aquatic habitat.
Populations no longer occur on Kauai.
The Oahu populations were described
from seven locations, and this species
now only occurs at one location. The
populations on Molokai have declined
from five to three. Populations on Lanai
have declined from four to one in an
artificial pond. On Maui, there are only
two populations, one on east Maui, and
one on west Maui. Of the 21 known
populations on Hawaii Island, only 14
remain. Because of the dramatic decline
in numbers and populations, and
because of the ongoing threats described
above, we find that this species is
endangered throughout all of its range,
and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
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Anchialine Pool Shrimp (Procaris
hawaiana)
The shrimp family Procarididae is
represented by a small number of
species globally, with only two species
within the genus Procaris (Magnacca
2015, in litt.). Procaris hawaiana is an
endemic anchialine pool shrimp species
known only from the islands of Maui
and Hawaii. The second species, P.
ascensionis, is restricted to similar
habitat on Ascension Island in the
South Atlantic Ocean. Of the anchialine
pools on Hawaii Island, only 25 are
known to contain P. hawaiana. During
nocturnal-diurnal surveys conducted
from 2009 to 2010, 19 pools within
Manuka NAR were found to contain P.
hawaiana. Five additional pools located
on unencumbered State land adjacent to
Manuka NAR also contained P.
hawaiana. An additional separate pool
also contains P. hawaiana, along with
the endangered anchialine pool shrimp
Vetericaris chaceorum (Holthuis 1973,
pp. 12–19; Maciolek 1983, pp. 607–614;
Brock 2004, pp. 30–57). On Maui, P.
hawaiana occurs in two anchialine
pools (Holthuis 1973, pp. 12–19;
Maciolek 1983, pp. 607–614; Brock
2004, pp. 30–57).
Like other anchialine pool shrimp
species, Procaris hawaiana inhabits
extensive networks of water-filled
interstitial spaces (cracks and crevices)
leading to and from the open pools
where they can be detected, a trait
which has precluded accurate estimates
of population size (Holthuis 1973, p. 36;
Maciolek 1983, pp. 613–616). Surveys
for many rare species of anchialine pool
shrimp, including P. hawaiana, often
involve baiting in likely habitat to
determine presence or absence.
Absence, and presumably extirpation, of
shrimp species from suitable habitat is
the best or only measure of species
decline as population sizes are not
easily determined or monitored
(Holthuis 1973, pp. 7–12; Maciolek
1983, pp. 613–616), but owing to the
potential for shrimp to move between
pools through subterranean
connections, the lack of sighting on one
or several visits to a site is not definitive
evidence that the species is extirpated
(Kinzie 2015, in litt.). Extirpation of
anchialine pool shrimp has been
documented definitively in some cases;
for example, Halocaridina rubra
disappeared from an anchialine pool at
Honokohau Harbor (Hawaii Island) as a
result of the use of the pool for dumping
of used oil, grease, and oil filters (Brock
2004, p. 14). To date, however, P.
hawaiana is not known to have been
extirpated from any of the pools where
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it has been documented to occur (Wada
2016, in litt.).
Habitat modification and destruction
by human activities is a significant
threat to Procaris hawaiana. It is
estimated that up to 90 percent of
existing anchialine pools in Hawaii
have been destroyed by filling and
bulldozing (Baily-Brock and Brock 1993,
p. 354; Brock 2004, p. i). Anchialine
pools are used as dumping pits for
bottles, cans, and used oil and grease,
and these activities are a known cause
of the disappearance of other anchialine
pool shrimp species from the pools.
Trampling damage from use of
anchialine pools for swimming and
bathing has been documented (Brock
2004, pp. 13–17). Although a permit
from the State is required to collect
anchialine pool shrimp, unpermitted
collection of shrimp is ongoing (FukuBonsai 2015, in litt.). A single person
with a handnet could do irreparable
damage to a population of P. hawaiana
(Yamamoto 2015, in litt.), but collection
by permitted individuals is not
prohibited at State Parks or City and
County property where some anchialine
pools occur. Predation by nonnative fish
is a direct threat to P. hawaiana.
Nonnative fish (tilapia, Oreochromis
mossambica) also outcompete native
herbivorous species of shrimp that serve
as a prey-base for P. hawaiana,
disrupting the delicate ecological
balance in the anchialine pool system,
and leading to decline of the pools and
the shrimp inhabiting them (Brock 2004,
pp. 13–17). Although anchialine pools
within State of Hawaii NARs are
provided some protection, these areas
are remote and signage does not prevent
human use and damage of the pools (see
Factor B). The persistence of P.
hawaiana is hampered by the small
number of extant populations and the
small geographic range of the known
populations. The populations of P.
hawaiana are at risk of extinction
because of their increased vulnerability
to loss of individuals from disturbance,
habitat destruction, and the effects of
invasive species and because of the
reduction in genetic variability that may
make the species less able to adapt to
changes in the environment (Harmon
and Braude 2010, pp. 125–128). In
addition, large-scale water extraction
from underground water sources
negatively affects the habitat and P.
hawaiana directly (Conry 2012, in litt.).
A threat from development upslope of
anchialine pool habitat is infiltration of
waste water or application of fertilizer
and pesticides that may enter the
ground water system of the anchialine
pools and consequently affect the pool’s
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ecosystem health, food sources of the
pool shrimp, or the pool shrimp directly
(Kinzie 2015, in litt.; Yamamoto et al.
2015, pp. 75–83). Sea-level rise and
coastal inundation resulting from the
effects of climate change is a threat to
P. hawaiana (Sakihara 2015, in litt.).
Sea-level rise would increase surface
connectivity between isolated
anchialine pools, and exacerbate the
spread of nonnative fish into pools not
yet occupied by nonnative fish
(Sakihara 2015, in litt.).
Procaris hawaiana and its habitat are
at risk. There are a total of 700 known
anchialine pools in the State of Hawaii.
Procaris hawaiana is restricted to 25
anchialine pools out of 600 on Hawaii
Island and to 2 anchialine pools on
Maui. These 27 anchialine pools
continue to be negatively affected by
habitat destruction and modification by
human use of the pools for bathing and
for dumping of trash and nonnative fish;
filling and bulldozing; water extraction;
contamination; predation by and
competition with nonnative fish; and
collection for the aquarium trade. The
small number of populations (27) limits
this species’ ability to adapt to
environmental changes. Because of
these threats, we find that this species
is endangered throughout all of its
range, and, therefore, find that it is
unnecessary to analyze whether it is
endangered or threatened in a
significant portion of its range.
Distinct Population Segment (DPS)
Analysis
Band-Rumped Storm-Petrel
(Oceanodroma castro)
Under the Act, we have the authority
to consider for listing any species,
subspecies, or, for vertebrates, any
distinct population segment (DPS) of
these taxa if there is sufficient
information to indicate that such action
may be warranted. To guide the
implementation of the DPS provisions
of the Act, we, and the National Marine
Fisheries Service (National Oceanic and
Atmospheric Administration—
Fisheries), published the Policy
Regarding the Recognition of Distinct
Vertebrate Population Segments Under
the Endangered Species Act (DPS
Policy) in the Federal Register on
February 7, 1996 (61 FR 4722). Under
our DPS Policy, we use two elements to
assess whether a population segment
under consideration for listing may be
recognized as a DPS: (1) The population
segment’s discreteness from the
remainder of the species to which it
belongs, and (2) the significance of the
population segment to the species to
which it belongs. If we determine that
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a population segment being considered
for listing is a DPS, then the population
segment’s conservation status is
evaluated based on the five listing
factors established by the Act to
determine if listing it as either
endangered or threatened is warranted.
In the proposed rule (80 FR 58820;
September 30, 2015), we evaluated the
Hawaii population of the band-rumped
storm-petrel to determine whether it
meets the definition of a DPS under our
DPS Policy.
Discreteness
Under the DPS Policy, a population
segment of a vertebrate taxon may be
considered discrete if it satisfies either
one of the following conditions: (1) It is
markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors
(quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation); or
(2) it is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the Act. The Hawaii
population of the band-rumped stormpetrel meets the first criterion: it is
markedly separated from other
populations of this species by physical
(geographic) and physiological (genetic)
factors, as described below.
The band-rumped storm-petrel is
widely distributed in the tropics and
subtropics, with breeding populations
in numerous island groups in the
Atlantic and in Hawaii, Galapagos, and
Japan in the Pacific (Harrison 1983, p.
274; Carboneras et al. 2014, p. 1 and Fig.
2). The geographic separation of these
breeding populations is widely
recognized, with strong genetic
differentiation between the two ocean
basins and among individual
populations (Friesen et al. 2007a, p.
1768; Smith et al. 2007, p. 768).
Whether individual populations merit
taxonomic separation remains unclear,
and further study is needed (Friesen et
al. 2007b, p. 18591; Smith et al. 2007,
p. 770; reviewed in Howell 2012, pp.
349, 369–370); some populations, such
as those in the Galapagos and Cape
Verde islands, may warrant full species
status (Smith et al. 2007, p. 770). Like
other storm-petrels, the band-rumped
storm-petrel is a highly pelagic (openocean) seabird (Howell 2012, p. 349). In
addition, like other species in the
seabird order Procellariiformes, bandrumped storm-petrels exhibit strong
philopatry, or fidelity to their natal sites
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(Allan 1962, p. 274; Harris 1969, pp. 96,
113, 120; Harrison et al. 1990, p. 49;
Smith et al. 2007, pp. 768–769). Both of
these characteristics contribute to
isolation of breeding populations, in
spite of the absence of physical barriers
such as land masses within ocean basins
(Friesen et al. 2007a, pp. 1777–1778).
Band-rumped storm-petrels from
Hawaii are likely to encounter
individuals from other populations only
very rarely. The approximate distances
from Hawaii to other known breeding
sites are much greater than the birds’
average foraging range of 860 mi (1,200
km): 4,000 mi (6,600 km) to Japan and
4,600 mi (7,400 km) to Galapagos (the
two other Pacific populations), and
7,900 mi (12,700 km) to Madeira, 7,300
mi (11,700 km) to the Azores, and 9,700
mi (15,600 km) to Ascension Island (in
the Atlantic). Data from at-sea surveys of
the eastern tropical Pacific conducted
since 1988 show that the density of
band-rumped storm-petrels attenuates
north and northwest of Galapagos and
that the species rarely occurs in a broad
area southeast of Hawaii (Pitman,
Ballance, and Joyce 2015, unpublished).
This pattern suggests a gap in the at-sea
distribution of this species, and low
likelihood of immigration on an
ecological timescale, between Hawaii
and Galapagos. We are not aware of any
data describing the at-sea distribution of
this species between Hawaii and Japan,
but the absence of breeding records from
western Micronesia (Pyle and Engbring
1985, p. 59) indicates a distributional
gap between these two archipelagoes as
well. Other than occasional encounters
in their foraging habitat, the vast
expanses of ocean between Japan,
Hawaii, and Galapagos provide for no
other sources of potential connectivity
between band-rumped storm-petrel
populations in the Pacific, such as
additional breeding sites.
Even those disparate breeding
populations of pelagic seabirds that do
overlap at sea may remain largely
isolated otherwise and exhibit genetic
differentiation (e.g., Walsh and Edwards
2005, pp. 290, 293). Despite the birds’
capacity to move across large areas of
ocean, genetic differentiation among
breeding populations of band-rumped
storm-petrels is high (Friesen et al.
2007b, p. 18590; Smith et al. 2007, p.
768), even between populations nesting
in different seasons on the same island
(in Galapagos; Smith and Friesen 2007,
p. 1599). Genetic analysis found low
relatedness (1) between Atlantic and
Pacific populations; (2) among Japan,
Hawaii, and Galapagos populations; or
(3) among Cape Verde, Ascension, and
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northeast Atlantic breeding populations
(Smith et al. 2007, p. 768). Hawaiian
birds have not been well-sampled for
genetic analysis, but the few individuals
from Hawaii included in a rangewide
analysis showed that Hawaiian birds
differed from all other populations, and
were most closely related to birds from
Japan (Friesen et al. 2007b, p. 18590).
We have determined that the Hawaii
population of the band-rumped stormpetrel is discrete from the rest of the
taxon because its breeding and foraging
range are markedly separated from those
of other populations. The Hawaii
population is geographically isolated
from populations in Japan and
Galapagos, as well as from populations
in very distant island groups in the
central and western Atlantic Ocean.
Molecular evidence indicates that the
genetic structure of the species reflects
the spatial or temporal separation of
individual populations; the scant
molecular data from Hawaii suggest that
this holds for the Hawaii population as
well.
Significance
Under our DPS Policy, once we have
determined that a population segment is
discrete, we consider its biological and
ecological significance to the larger
taxon to which it belongs. This
consideration may include, but is not
limited to: (1) Evidence of the
persistence of the discrete population
segment in an ecological setting that is
unusual or unique for the taxon, (2)
evidence that loss of the population
segment would result in a significant
gap in the range of the taxon, (3)
evidence that the population segment
represents the only surviving natural
occurrence of a taxon that may be more
abundant elsewhere as an introduced
population outside its historical range,
or (4) evidence that the discrete
population segment differs markedly
from other populations of the species in
its genetic characteristics. We have
found substantial evidence that the
Hawaii population of the band-rumped
storm-petrel meets two of the
significance criteria listed above: the
loss of this population would result in
a significant gap in the range of the
taxon, and this population persists in a
unique ecological setting. As described
above, the physical isolation that
defines the discreteness of Hawaii
population is likely reflected in genetic
differentiation from other populations,
but at this time we lack sufficient data
to consider genetic characteristics as an
independent factor in our determination
of the Hawaii population’s significance
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to the rest of the taxon. Genetic patterns
on an ocean-basin or species-wide scale,
however, have implications for
connectivity and potential gaps in the
band-rumped storm-petrel’s range
(described below).
Dispersal between populations of
seabird species with ranges fragmented
by large expanses of ocean may play a
vital role in the persistence of
individual populations (Bicknell et al.
2012, p. 2872). No evidence currently
exists of such dispersal among Pacific
populations of band-rumped stormpetrels at frequencies or in numbers that
would change the population status
between years, for example, by
providing immigrants that compensate
for breeding failure or adult mortality
resulting from predation, as has been
hypothesized for Leach’s storm-petrel in
the Atlantic (Bicknell et al. 2012, p.
2872). Given the remnant population of
band-rumped storm-petrels in Hawaii
and recently documented decline in
Japan (Biodiversity Center of Japan
2014, p. 1), we would not expect to see
exchange on such short timescales.
However, genetic evidence is suggestive
of exchange between these two
populations on an evolutionary
timescale (Friesen et al. 2007b, p.
18590).
The loss of this population would
result in a significant gap in the range
of the band-rumped storm-petrel. As
noted above, seabirds in the order
Procellariiformes, including the bandrumped storm-petrel, exhibit very high
natal site fidelity, and so are slow to
recolonize extirpated areas or rangegaps (Jones 2010, p. 1214), and may lack
local adaptations; they thus face a
potentially increased risk of extinction
with the loss of individual populations
(Smith et al. 2007, p. 770). The Hawaii
population of the band-rumped storm
petrel constitutes the entire Central
Pacific distribution of the species,
located roughly half-way between the
populations in Galapagos and Japan (see
Figure 1, below), and its loss would
create a gap of approximately 8,500 mi
(13,680 km) between them and
significantly reducing the likelihood of
connectivity and genetic exchange.
Such exchange would be reliant on
chance occurrences, such as severe
storms that could result in birds being
displaced to the opposite side of the
Pacific Ocean basin, and such chance
dispersal events would not necessarily
result in breeding.
BILLING CODE 4333–15–P
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The Hawaii population of the bandrumped storm-petrel is significant also
because it persists in a unique
ecological setting. This is the only
population of the species known to nest
at high-elevation sites (above 6,000 ft
(1,800 m)) (Banko et al. 1991, pp. 651–
653; Athens et al. 1991, p. 95). In
prehistory, the species likely nested in
lowland habitats and more accessible
habitats in Hawaii as well as in the highelevation and otherwise remote areas
where the species is found today;
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archaeological evidence suggests that
band-rumped storm-petrels were once
sufficiently common at both high (5,260
and 6,550 ft (1,600 and 2,000 m)) and
low elevations on Hawaii Island to be
used as a food source by humans
(Ziegler pers. comm. in Harrison et al.
1990, pp. 47–48; Athens et al. 1991, pp.
65, 78–80; Banko et al. 1991, p. 650). In
lowland areas, the species was common
enough for the Hawaiians to name it and
to identify it by its call (Harrison et al.
1990, p. 47; Banko et al. 1991, p. 650).
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67821
In addition to the impacts of harvest by
humans in prehistory, seabirds in
Hawaii, including the band-rumped
storm-petrel, were negatively affected by
the proliferation of nonnative predators
such as rats and pigs, and, later, cats
and mongoose, and by loss of habitat
(reviewed in Duffy 2010, pp. 194–196).
Predation and habitat loss combined
likely led to the extirpation of the bandrumped storm-petrel from coastal and
lowland habitats and other accessible
nesting areas, as occurred in the
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endangered Hawaiian petrel
(Pterodroma sandwichensis) and
threatened Newell’s shearwater
(Puffinus newelli), which have similar
nesting habits and life histories (Olson
and James 1982, p. 43; Slotterback 2002,
p. 6; Troy et al. 2014, pp. 315, 325–326).
The band-rumped storm-petrel’s
persistence in sites such as the
Southwest Rift Zone (6,900 ft (2,100 m))
on Mauna Loa (Hawaii Island) has
required them to surmount
physiological challenges posed by
nesting in high-elevation conditions
(cold temperatures, low humidity, and
less oxygen). They may possess special
adaptations for this, such as reduction
in porosity and other eggshell
modifications to reduce the loss of water
and carbon dioxide during incubation at
high elevation (Rahn et al. 1977, p.
3097; Carey et al. 1982, p. 716; Carey et
al. 1983, p. 349). In sum, the remnant
distribution of band-rumped stormpetrel breeding sites in only the most
remote and rugged terrain in Hawaii
reflects the conditions necessary for the
species’ persistence in Hawaii
(relatively undisturbed habitat in areas
least accessible to predators) and also
reflects unique adaptations that
facilitate the species’ persistence in
high-elevation areas.
We have determined that the Hawaii
population of band-rumped storm-petrel
is significant to the rest of the taxon. Its
loss would result in a gap in the range
of the species of more than 8,500 mi
(13,680 km), reducing and potentially
precluding connectivity between the
two remaining populations in the
Pacific Basin. In addition, the Hawaii
population nests at high elevation on
some islands, constituting a unique
ecological setting represented nowhere
else in the species’ breeding range.
DPS Conclusion
We have evaluated the Hawaii
population of band-rumped storm-petrel
to determine if it meets the definition of
a DPS, considering its discreteness and
significance as required by our policy.
We have found that this population is
markedly separated from other
populations by geographic distance, and
this separation is likely reflected in the
population’s genetic distinctiveness.
The Hawaii population is significant to
the rest of the species because its loss
would result in a significant gap in the
species’ range; Hawaii is located
roughly half-way between the other two
populations in the Pacific Ocean, and
little or no evidence exists of current
overlap at sea between the Hawaii
population and either the Japan or
Galapagos populations. The Hawaii
population of band-rumped storm-petrel
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also nests at high elevation in Hawaii—
conditions at high elevation constitute
an ecological setting unique to the
species. We conclude that the Hawaii
population of band-rumped storm-petrel
is a distinct vertebrate population
segment under our February 7, 1996,
DPS Policy (61 FR 4722), and that it
warrants review for listing under the
Act. Therefore, we have incorporated
the Hawaii DPS of the band-rumped
storm-petrel in our evaluation of threats
affecting the other 48 species addressed
in this rule (summarized above; see also
Summary of Factors Affecting the 49
Species From the Hawaiian Islands,
below).
Summary of Factors Affecting the 49
Species From the Hawaiian Islands
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations (50
CFR part 424), set forth the procedures
for adding species to the Federal Lists
of Endangered and Threatened Wildlife
and Plants. A species may be
determined to be an endangered or
threatened species due to one or more
of the five factors described in section
4(a)(1) of the Act: (A) The present or
threatened destruction, modification, or
curtailment of its habitat or range; (B)
overutilization for commercial,
recreational, scientific, or educational
purposes; (C) disease or predation; (D)
the inadequacy of existing regulatory
mechanisms; and (E) other natural or
manmade factors affecting its continued
existence. Listing actions may be
warranted based on any of the above
threat factors, singly or in combination.
Each of these factors is discussed below.
In considering factors that might
constitute threats to a species, we must
look beyond the exposure of the species
to a factor to evaluate whether the
species responds to the factor in a way
that causes impacts to the species or is
likely to cause impacts in the future. If
a species responds negatively to such
exposure, the factor may be a threat and,
during the status review, our aim is to
determine whether impacts are or will
be of an intensity or magnitude to place
the species at risk. The factor is a threat
if it drives, or contributes to, the risk of
extinction of the species such that the
species warrants listing as an
endangered or threatened species as
those terms are defined by the Act. This
does not necessarily require empirical
proof of a threat. The combination of
exposure and some corroborating
evidence of how the species is likely
affected could suffice. In sum, the mere
identification of factors that could affect
a species negatively is not sufficient to
compel a finding that listing is
appropriate; we require evidence that
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Fmt 4701
Sfmt 4700
these factors act on the species to the
point that the species meets the
definition of an endangered or
threatened species.
If we determine that the threats posed
to a species by one or more of the five
listing factors are, or are likely to
become, of such magnitude and/or
intensity that the species meets the
definition of either endangered or
threatened under section 3 of the Act,
that species may then be listed as
endangered or threatened. The Act
defines an endangered species as ‘‘in
danger of extinction throughout all or a
significant portion of its range,’’ and a
threatened species as ‘‘likely to become
an endangered species within the
foreseeable future throughout all or a
significant portion of its range.’’ The
threats to each of the individual 49
species are summarized in Table 2, and
discussed in detail below.
We acknowledge that the specific
threats to the individual species in this
final rule are not all completely
understood. Scientific study of each of
the 49 species is limited because of their
rarity and the challenging logistics
associated with conducting field work
in Hawaii (areas are typically remote,
difficult to access, challenging work
environments, and expensive to survey
in a comprehensive manner). However,
information is available on many of the
threats that act on Hawaiian ecosystems,
and, for some ecosystems, these threats
are well studied and understood. Each
of the native species that occurs in
Hawaiian ecosystems suffers from
exposure to those threats to differing
degrees. For the purposes of our listing
determination, the best available
scientific information leads us to
conclude that the threats that act at the
ecosystem level also act on each of the
species that occurs in those ecosystems.
In some cases we have additionally
identified species-specific threats, such
as loss of host plants.
The following threats affect the 49
species in one or more of the ecosystems
addressed in this rule:
(1) Modification and destruction of
habitat, including streams, ponds, and
anchialine pools, by urban development
and water extraction. Human activities
also contribute to increased
sedimentation in anchialine pools.
(2) Habitat destruction and
modification by feral ungulates
including pigs, goats, axis deer, blacktailed deer, mouflon, sheep, and cattle.
The disturbance of soils by these
animals causes erosion and creates
fertile seedbeds for nonnative plants,
leading to further habitat degradation.
Ungulates also trample seedlings.
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(3) Habitat destruction and
modification by nonnative plants.
Nonnative plants modify availability of
light, alter soil-water regimes, modify
nutrient cycling, alter fire regimes, and
ultimately convert native dominated
plant communities to nonnative plant
communities. They also cause or
contribute to loss of host plants used for
food and nesting by the yellow-faced
bees.
(4) Habitat destruction by wildfires
caused naturally or by humans. Fires
also destroy the native plant seedbank,
and contribute to habitat conversion of
native forest to nonnative grasslands
(grass/fire cycle).
(5) Habitat destruction and
modification, or direct damage and
death, by stochastic events including
drought, erosion, flooding, tree falls,
rock falls, landslides, hurricanes, and
tsunamis.
(6) Illegal collection of anchialine
pool shrimp for personal use or
commercial trade.
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(7) Herbivory or defoliation of native
plants by ungulates, rats, slugs, and
black twig borers, which have been
observed to contribute to the decline or
death of 35 the 39 plant species (except
for Cyperus neokunthianus, Cyrtandra
hematos, Lepidium orbiculare, and
Stenogyne kaalae ssp. sherffii).
Herbivory also destroys seeds and fruit
and contributes to lack of reproduction
in the wild and low genetic diversity
compounding the decline of native
plants.
(8) Predation of the band-rumped
storm-petrel by rats, barn owls, cats, and
mongoose.
(9) Predation of the orangeblack
Hawaiian damselfly by bullfrogs,
backswimmers, Jackson’s chameleons,
and nonnative fish.
(10) Predation of the anchialine pool
shrimp by nonnative fish.
(11) Predation of Hylaeus bees by ants
and wasps.
(12) Competition for food and nesting
sites of the Hylaeus yellow-faced bees
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67823
by nonnative ants, wasps, and bees, and
competition for food and habitat of the
orangeblack Hawaiian damselfly by
caddisflies. Competition for space and
food resources of the anchialine pool
shrimp by nonnative fish.
(13) Injury and mortality of the bandrumped storm-petrel caused by artificial
lighting, communication towers, and
power lines.
(14) Injury and mortality of the bandrumped storm-petrel by the activities of
fisheries and encounters with marine
debris.
(15) Low numbers and/or no
reproduction of all 49 species
exacerbated by one or more of the above
threats, combined with inability of the
species to adapt to sea-level rise or other
factors associated with climate change.
Existing regulatory mechanisms do
not ameliorate these threats for any of
the 49 species such that listing is not
warranted. Each of the threats listed
above is discussed in more detail below,
and summarized in Table 2.
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LM ......................
CO ......................
MM, MD, SA .......
MW, MM, WC .....
MM, MD, SA .......
LM, LW ...............
LW, MW .............
LW, MW, MM,
WC.
LM, MM, DC .......
MW, MM, MD .....
CO ......................
LW ......................
LW ......................
CO, SA, DC, WC
AP, CO, LD, LM
AP .......................
CO, LD ...............
CO, LD ...............
CO, LD, LM ........
Pritchardia bakeri ....................................................................
Pseudognaphalium sandwicensium var. molokaiense ...........
Ranunculus hawaiensis ..........................................................
Ranunculus mauiensis ............................................................
Sanicula sandwicensis ............................................................
Santalum involutum ................................................................
Schiedea diffusa ssp. diffusa ..................................................
Schiedea pubescens ...............................................................
Sfmt 4700
Sicyos lanceoloideus ..............................................................
Sicyos macrophyllus ...............................................................
Solanum nelsonii .....................................................................
Stenogyne kaalae ssp. sherffii ................................................
Wikstroemia skottsbergiana ....................................................
ANIMALS:
Band-rumped storm-petrel (Oceanodroma castro) .................
Orangeblack Hawaiian damselfly (Megalagrion xanthomelas)
Anchialine pool shrimp (Procaris hawaiana) ..........................
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Yellow-faced bee (Hylaeus anthracinus) ................................
Yellow-faced bee (Hylaeus assimulans) .................................
Yellow-faced bee (Hylaeus facilis) ..........................................
X ..................
X ..................
X ..................
X, WE ..........
X, WE ..........
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CO, LD, MD .......
Portulaca villosa ......................................................................
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LM, MM, DC .......
LW, MW, WC .....
LW, MW, WC .....
MW, MM .............
LM, LW ...............
MW .....................
LM, MM, MD ......
MD ......................
LM, LW ...............
MW .....................
MW .....................
LM, LW, MW,
MM.
LM, LW ...............
LM ......................
MM .....................
LM ......................
LM, MW, MM ......
LM, LW, MW ......
LD, LM, DC ........
.....................
.....................
.....................
X, WE ..........
.....................
.....................
Agriculture
and urban
development
Ochrosia haleakalae ...............................................................
Phyllostegia brevidens ............................................................
Phyllostegia helleri ..................................................................
Phyllostegia stachyoides .........................................................
Kadua fluviatilis .......................................................................
Kadua haupuensis ..................................................................
Labordia lorenciana ................................................................
Lepidium orbiculare .................................................................
Microlepia strigosa var. mauiensis .........................................
Myrsine fosbergii .....................................................................
Nothocestrum latifolium ..........................................................
Deparia kaalaana ....................................................................
Dryopteris glabra var. pusilla ..................................................
Exocarpos menziesii ...............................................................
Festuca hawaiiensis ................................................................
Gardenia remyi .......................................................................
Huperzia stemmermanniae .....................................................
Hypolepis hawaiiensis var. mauiensis ....................................
Joinvillea ascendens ssp. ascendens ....................................
MM .....................
MW .....................
LW ......................
LW, MW .............
LW ......................
MW .....................
Ecosystem
P, G, D, C
P, G, D,
M, C.
P, G, D,
SH, C.
P, G, D ...
.................
G .............
P, G, BTD
P, M, C ...
D, C ........
P .............
P, G ........
G, D, M,
C.
P .............
G, D ........
P, M, C ...
P, G, D,
BTD, C.
P, G ........
P, G ........
P .............
P, G, D, C
P .............
P, BTD ....
G, M, SH
G, SH .....
P, G, D ...
P, G, D, C
.................
P, G, D,
BTD.
P, G ........
P .............
P, G, BTD
P, G ........
P, G ........
P, G ........
P, G, D,
BTD, M,
C.
P, G, C ...
P .............
P, G ........
P, G, D ...
P, G, BTD
P .............
.................
P .............
P .............
P, G ........
Ungulates
............
............
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X ............
X ............
X ............
X ............
...............
X ............
X
X
X
X
X
X
X
X
X
X
X
X
X
X ............
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Nonnative
plants
Factor A
X .............
X .............
X .............
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X .............
X .............
X .............
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X .............
X .............
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X .............
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X .............
X .............
X .............
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X .............
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X .............
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X .............
X .............
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X .............
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Fire
DR, HUR,
TS.
DR, HUR,
TS.
DR, HUR,
TS.
DR, F, HUR
.....................
E, L, HUR ....
DR ...............
.....................
DR, E, F, TS
.....................
L ..................
DR, E, F ......
.....................
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DR, E, F ......
HUR ............
L, RF ...........
DR, E ..........
DR, E, L ......
.....................
E, L ..............
L ..................
DR, E, F, L,
RF.
L, RF ...........
L ..................
L ..................
F, L, TF .......
L ..................
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DR, F ...........
L ..................
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L ..................
DR ...............
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L ..................
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E, F, L .........
DR, F, L ......
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Stochastic
events
.................
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X .............
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Overutilization
Factor B
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X .................
X .................
X .................
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X .................
X .................
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X .................
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X .................
X .................
X .................
X .................
X .................
X .................
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X .................
X .................
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X .................
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X .................
X .................
X .................
X .................
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X .................
X .................
X .................
X .................
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X .................
X .................
X .................
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X .................
....................
....................
Predation/
herbivory by
ungulates
Factor D
................
................
................
................
....................
....................
....................
R, O, CA,
MO.
FS, BF, JC
FS ..............
....................
R ................
R ................
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R ................
R
R
R
R
R ................
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R ................
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R ................
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R ................
R ................
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R ................
R ................
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R ................
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R ................
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R ................
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R ................
R ................
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A, W .............
A, W .............
A, W, ............
BS ................
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S ..................
S ..................
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S ..................
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S ..................
S ..................
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S ..................
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BTB ..............
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BTB ..............
S ..................
S ..................
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S ..................
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S ..................
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.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
.................
X .................
X .................
X .................
X .................
X .................
X .................
X
X
X
X
X
X
X
X
X
X
X
X
X
X .................
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Inadequate
existing
regulatory
mechanisms
Predation/
herbivory by
NN
invertebrates
Factor C
Predation/
herbivory by
other NN
vertebrates
TABLE 2—SUMMARY OF PRIMARY THREATS IDENTIFIED FOR EACH OF THE 49 HAWAIIAN ISLANDS SPECIES
PLANTS:
Asplenium diellaciniatum ........................................................
Calamagrostis expansa ..........................................................
Cyanea kauaulaensis ..............................................................
Cyclosorus boydiae .................................................................
Cyperus neokunthianus ..........................................................
Cyrtandra hematos .................................................................
Species
asabaliauskas on DSK3SPTVN1PROD with RULES
NR
........
........
........
........
........
SL ..
........
........
........
........
........
........
........
........
........
........
LI, ST,
H, LN.
LN, CD
FS, H,
LN,
RU,
SD,
SL.
LN, W,
B,
LHP.
LN, W,
B,
LHP.
LN, W,
B,
LHP.
LN
LN
LN,
LN
LN
LN
LN
LN
LN
LN
LN
LN
LN
LN ........
LN,
LN
LN
LN
LN ........
LN ........
LN, NR
LN ........
HY, LN
HY, LN
LN, NR
LN ........
LN ........
LN, NR
LN ........
LN ........
HY, LN,
NR.
LN ........
LN ........
LN ........
LN ........
LN, NR
LN ........
LN ........
LN, NR
Ft.
Ft.
Ft.
Ft.
Ft.
Ft.
Ft.
Ft.
Ft.
Ft.
Ft.
X.
X.
X.
X.
X.
X.
X,
X,
X,
X,
X.
Ft.
Ft.
Ft.
Ft.
X, Ft.
X.
X.
X, Ft.
X.
X, Ft.
X, Ft.
X, Ft.
X, Ft.
X,
X,
X,
X,
X, Ft.
X.
X.
X.
X.
X, Ft.
X, Ft.
X.
X,
X,
X,
X,
X,
X,
X,
X.
X, Ft.
X.
X, Ft.
X.
X, Ft.
Climate
change
Factor E
Other
speciesspecific
threats
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CO, LD ...............
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Yellow-faced bee (Hylaeus longiceps) ...................................
.....................
X ..................
.....................
X ..................
P .............
D .............
P .............
P, G, D, C
X ............
X ............
X ............
X ............
X .............
X .............
X .............
X .............
DR, HUR .....
DR, HUR,
TS.
DR, HUR .....
DR, HUR,
TS.
.................
.................
.................
.................
....................
....................
....................
....................
....................
....................
....................
....................
A, W .............
A, W .............
A, W .............
A, W .............
X .................
X .................
X .................
X .................
LN, W,
B,
LHP.
LN, W,
B,
LHP.
LN, W,
B,
LHP,
RU.
LN, W,
B,
LHP.
X.
X.
X.
X.
Factor A = Habitat Modification; Factor B = Overutilization; Factor C = Disease or Predation; Factor D = Inadequacy of Regulatory Mechanisms: the Xs in this column indicate that existing regulatory mechanisms do not ameliorate the threats to the
species such that listing is not warranted (we do not identify Factor D, in and of itself, as a threat to the species); Factor E = Other Species-Specific Threats.
AP = Anchialine Pools; CO = Coastal; LD = Lowland Dry; LM = Lowland Mesic; LW = Lowland Wet; MW = Montane Wet; MM = Montane Mesic; MD = Montane Dry; SA = Subalpine; DC = Dry Cliff; WC = Wet Cliff.
A = Ants, B = Bees (competition); BF = Bullfrog; BS = Backswimmer; BTB = Black Twig Borer; BTD = Black-tailed Deer; C = Cattle; CA = Cats; CD = Caddisflies; D = Axis Deer; ; FS = Fish; G = Goats; JC = Jackson’s Chameleons; M = Mouflon;
MO = Mongoose; O = Barn Owls; P = Pigs; R = Rats; S = Slugs; SH = Sheep; W = Wasps (competition, predation).
DR = Drought; E = Erosion; F = Flooding; Ft = assessed in Fortini et al. (2013) climate change vulnerability analysis; H = Human (fisheries, marine debris, contamination); HUR = Hurricanes; HY = Hybridization; L = Landslides; LHP = Loss of Host
Plants; LI = Lights; LN = Low Numbers; NR = No Regeneration; RF = Rockfalls; RU = Recreational Use (swimming, fishing, dumping trash and nonnative fish); SD = Sedimentation; SL = Sea Level Rise; ST = Structures; TF = Tree Fall; TS = Tsunami;
WE = Water Extraction.
LM ......................
CO, LD ...............
Yellow-faced bee (Hylaeus kuakea) .......................................
Yellow-faced bee (Hylaeus mana) ..........................................
LM ......................
Yellow-faced bee (Hylaeus hilaris) .........................................
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Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of Their Habitat or Range
The Hawaiian Islands are located over
2,000 mi (3,200 km) from the nearest
continent. This isolation has allowed
the few plants and animals transported
to the islands by wind, water, or birds
to evolve into many varied and endemic
species. The only native terrestrial
mammals on the Hawaiian Islands
include two bat taxa, the Hawaiian
hoary bat (Lasiurus cinereus semotus),
and an extinct, unnamed insectivorous
bat (Ziegler 2002, p. 245). The native
plants of the Hawaiian Islands therefore
evolved in the absence of mammalian
predators, browsers, or grazers, and
subsequently, many native species lost
unneeded defenses against threats
typical of continental environments
such as herbivory and competition with
aggressive, weedy plant species (Loope
1992, p. 11; Gagne and Cuddihy 1999,
p. 45; Wagner et al. 1999, pp. 3–6). For
example, Carlquist (in Carlquist and
Cole 1974, p. 29) notes, ‘‘Hawaiian
plants are notably nonpoisonous, free
from armament, and free from many
characteristics thought to be deterrents
to herbivores (oils, resins, stinging hairs,
coarse texture).’’ In addition, species
restricted to highly specialized habitats
(e.g., Hawaiian damselflies) or food and
nesting sources (e.g., Hawaiian yellowfaced bees) are particularly vulnerable
to changes in their habitat (Carlquist
and Cole 1974, pp. 28–29; Loope 1992,
pp. 3–6).
asabaliauskas on DSK3SPTVN1PROD with RULES
Habitat Destruction and Modification by
Agriculture and Urban Development
Past land use practices such as
agriculture or urban development have
resulted in little or no native vegetation
below 2,000 ft (600 m) throughout the
Hawaiian Islands (TNC 2006). These
land use practices negatively affect the
anchialine pool, coastal, lowland dry,
and lowland mesic ecosystems,
including streams and wetlands that
occur within these areas. Hawaii’s
agricultural industries (e.g., sugar cane,
pineapple) have been declining in
importance, and large tracts of former
agricultural lands are being converted
into residential areas or left fallow (TNC
2007). In addition, Hawaii’s population
has increased almost 10 percent in the
past 10 years, further increasing
demands on limited land and water
resources in the islands (Hawaii
Department of Business, Economic
Development and Tourism 2013, in
litt.).
Development and urbanization of
anchialine pool, coastal, lowland dry,
and lowland mesic ecosystems on Oahu,
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Molokai, Maui, Lanai, and Hawaii
Island are a threat to some species:
• On Oahu, the plant Cyclosorus
boydiae, the orangeblack Hawaiian
damselfly, and the yellow-faced bees
Hylaeus anthracinus, H. assimulans, H.
facilis, and H. longiceps.
• On Molokai, the orangeblack
Hawaiian damselfly and the yellowfaced bees Hylaeus anthracinus, H.
facilis, H. hilaris, and H. longiceps.
• On Maui, the plant Cyclosorus
boydiae, the orangeblack Hawaiian
damselfly, the anchialine pool shrimp
Procaris hawaiana, and the yellowfaced bees Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H.
longiceps.
• On Lanai, the orangeblack Hawaiian
damselfly, and the yellow-faced bees
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, and H. longiceps.
• On Hawaii Island, the orangeblack
Hawaiian damselfly, the anchialine pool
shrimp Procaris hawaiana, and the
yellow-faced bee Hylaeus anthracinus.
(Daly and Magnacca 2003, pp. 55, 173;
Palmer 2003, p. 88; Magnacca 2007, p.
188; Magnacca and King 2013, pp. 22–
25).
Although we are unaware of any
comprehensive, site-by-site assessment
of wetland development in Hawaii
(Erikson and Puttock 2006, p. 40), Dahl
(1990, p. 7) estimated that at least 12
percent of lowland to upper-elevation
wetlands in Hawaii had been converted
to non-wetland habitat by the 1980s. If
only coastal plain (below 1,000 ft (300
m)) marshlands and wetlands are
considered, it is estimated that 30
percent were developed or converted to
agricultural use (Kosaka 1990, in litt.).
Records show the modification and
reduction in area of these marshlands
and wetlands that provided habitat for
many damselfly species, including the
orangeblack Hawaiian damselfly
(Englund 2001, p. 256; Rees and Reed
2013, Fig 2S). Once modified, these
areas then lack the aquatic habitat
features that the orangeblack Hawaiian
damselfly requires for essential lifehistory needs, such as pools of
intermittent streams, ponds, and coastal
springs (Polhemus 1996 pp. 30–31, 36).
Although the filling of wetlands is
regulated by section 404 of the Clean
Water Act (33 U.S.C. 1251 et seq.), the
loss of riparian or wetland habitats
utilized by the orangeblack Hawaiian
damselfly may still occur due to
Hawaii’s population growth and
development, with concurrent demands
on limited developable land and water
resources. The State’s Commission of
Water Resource Management (CWRM)
recognizes the need to update the 2008
water resource protection plan, and an
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update is currently under development
(CWRM 2014, in litt.). In addition,
marshes have been slowly filled and
converted to meadow habitat as a result
of sedimentation from increased storm
water runoff from upslope development,
the accumulation of uncontrolled
growth of invasive vegetation, and
blockage of downslope drainage (Wilson
Okamoto & Associates, Inc. 1993, pp. 3–
4–3–5). Agriculture and urban
development have thus contributed to
habitat destruction and modification,
and continue to be a threat to the habitat
of the orangeblack Hawaiian damselfly
and the fern, Cyclosorus boydiae.
On Hawaii Island, it is estimated that
up to 90 percent of the anchialine pools
have been destroyed or altered by
human activities, including bulldozing
and filling of pools (Brock 2004, p. i;
Bailey-Brock and Brock 1993, p. 354).
Dumping of trash and nonnative fish
has affected anchialine pools on this
island (Brock 2004, pp. 13–17) (see
Factor E. Other Natural or Manmade
Factors Affecting Their Continued
Existence, below). Brock also noted that
garbage like bottles and cans appear to
have no net negative impact, while the
dumping of used oil, oil filters, and
grease has resulted in the disappearance
of the anchialine pool shrimp
Halaocaridina rubra from a pool
adjacent to Honokohau Harbor on
Hawaii Island. Lua O Palahemo (where
Procaris hawaiana occurs) on Hawaii
Island is accessible to the public, and
dumping has occurred there (Brock
2004, pp. 13–17). We are not aware of
any dumping activities within the two
Maui anchialine pools known to be
occupied by P. hawaiana; however, this
threat remains a possibility (Brock 2004,
pp. 13–17).
Destruction and modification of
Hylaeus habitat by urbanization and
land use conversion, including
agriculture, has led to the fragmentation
of foraging and nesting habitat of these
species. In particular, because native
host plant species are known to be
essential to the yellow-faced bees for
foraging of nectar and pollen, any
further loss of this habitat may reduce
their long-term chances for recovery.
Additionally, further destruction and
modification of Hylaeus habitat is also
likely to facilitate the introduction and
spread of nonnative plants within these
areas (see ‘‘Habitat Destruction and
Modification by Nonnative Plants,’’
below).
Habitat Destruction and Modification by
Nonnative Ungulates
Nonnative ungulates have greatly
affected the native vegetation, as well as
the native fauna, of the Hawaiian
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asabaliauskas on DSK3SPTVN1PROD with RULES
Islands. Impacts to the native species
and ecosystems accelerated following
the arrival of Captain James Cook in
1778. The Cook expedition and
subsequent explorers intentionally
introduced a European race of pigs (i.e.,
boars) and other livestock such as goats
to serve as food sources for seagoing
explorers (Tomich 1986, pp. 120–121;
Loope 1998, p. 752). The mild climate
of the islands, combined with lack of
competitors or predators, led to the
successful establishment of large
populations of these feral mammals, to
the detriment of native Hawaiian
species and ecosystems (Cox 1992, pp.
116–117). The presence of introduced
mammals is considered one of the
primary factors underlying the
modification and destruction of native
vegetation and habitats of the Hawaiian
Islands (Cox 1992, pp. 118–119). All of
the 11 ecosystems on the main islands
(except Kahoolawe) are currently
affected by habitat destruction resulting
from the activities of various
combinations of nonnative ungulates,
including pigs (Sus scrofa), goats (Capra
hircus), axis deer (Axis axis), blacktailed deer (Odocoileus hemionus
columbianus), sheep (Ovis aries),
mouflon (Ovis gmelini musimon) and
mouflon-sheep hybrids, and cattle (Bos
taurus). Habitat destruction or
modification by ungulates is a threat to
37 of the 39 plant species, the bandrumped storm-petrel, the orangeblack
Hawaiian damselfly, and the seven
yellow-faced bees (see Table 2).
Pigs (Sus scrofa)
The destruction or modification of
habitat by pigs is currently a threat to
four of the ecosystems (lowland mesic,
lowland wet, montane wet, and
montane mesic) in which these species
occur. Feral pigs are known to cause
deleterious impacts to ecosystem
processes and functions throughout
their worldwide distribution (Campbell
and Long 2009, p. 2319). Pigs have been
described as having the most pervasive
and disruptive nonnative influences on
the unique ecosystems of the Hawaiian
Islands and are widely recognized as
one of the greatest current threats (Aplet
et al. 1991. p. 56; Anderson and Stone
1993, p. 195; Anderson et al. 2007, in
litt.). Introduced European pigs
hybridized with smaller, domesticated
Polynesian pigs, became feral, and
invaded forested areas, especially mesic
and wet forests, from low to high
elevations; they are present on all the
main Hawaiian Islands except Lanai and
Kahoolawe, where they have been
eradicated (Tomich 1986, pp. 120–121;
Munro (1911–1930) 2007, p. 85). By the
early 1900s, feral pigs were already
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recognized as a serious threat to these
areas, and an eradication project was
conducted by the Hawaii Territorial
Board of Agriculture and Forestry,
which removed 170,000 pigs from
forests Statewide (Diong 1982, p. 63).
Feral pigs are extremely destructive
and have both direct and indirect
impacts on native plant communities.
While rooting in the earth in search of
invertebrates and plant material, pigs
directly affect native plants by
disturbing and destroying vegetative
cover and by trampling plants and
seedlings. It has been estimated that at
a conservative rooting rate of 2 square
yards (sq yd) (1.7 square meters (sq m))
per minute and only 4 hours of foraging
per day, a single pig could disturb over
1,600 sq yd (1,340 sq m) (or
approximately 0.3 acres (ac) (0.1
hectares (ha)) of groundcover per week
(Anderson et al. 2007, in litt.). Feral pigs
are a major vector for the establishment
and spread of invasive nonnative plant
species, such as Passiflora tarminiana
and Psidium cattleianum, by dispersing
seeds carried on their hooves and coats
and in their feces (which also serve to
fertilize disturbed soil) (Diong 1982, pp.
169–170; Matson 1990, p. 245; Siemann
et al. 2009, p. 547). Pigs also feed
directly on native plants such as
Hawaiian tree ferns. Pigs preferentially
eat the core of tree-fern trunks, and
these cored trunks then fill with
rainwater and serve as breeding sites for
introduced mosquitoes that spread
avian malaria, with devastating
consequences for Hawaii’s native forest
birds (Baker 1975, p. 79). Additionally,
rooting pigs contribute to erosion,
especially on slopes, by clearing
vegetation and creating large areas of
disturbed soil (Smith 1985, pp. 190,
192, 196, 200, 204, 230–231; Stone
1985, pp. 254–255, 262–264; Medeiros
et al. 1986, pp. 27–28; Scott et al. 1986,
pp. 360–361; Tomich 1986, pp. 120–
126; Cuddihy and Stone 1990, pp. 64–
65; Aplet et al. 1991, p. 56; Loope et al.
1991, pp. 1–21; Gagne and Cuddihy
1999, p. 52; Nogueira-Filho et al. 2009,
pp. 3677–3682; Dunkell et al. 2011, pp.
175–177). The resulting erosion alters
native plant communities by damaging
individual plants, contributing to
watershed degradation, and changing
nutrient availability for plants; erosion
also affects aquatic animals by
increasing sedimentation in streams and
pools (Vitousek et al. 2009, pp. 3074–
3086; Nogueira-Filho et al. 2009, p.
3681; Cuddihy and Stone 1992, p. 667).
The following 15 plants are at risk from
erosion and landslides resulting from
the activities of feral pigs: Cyclosorus
boydiae, Dryopteris glabra var. pusilla,
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Gardenia remyi, Joinvillea ascendens
ssp. ascendens, Kadua fluviatilis, K.
haupuensis, Labordia lorenciana,
Lepidium orbiculare, Phyllostegia
brevidens, P. helleri, P. stachyoides,
Ranunculus hawaiensis, R. mauiensis,
Sanicula sandwicensis, and Schiedea
pubescens. Thirty-two of the 39 plants
(all except for Cyanea kauaulaensis,
Exocarpos menziesii, Festuca
hawaiiensis, Hypolepis hawaiiensis var.
mauiensis, Portulaca villosa,
Pseudognaphalium sandwicensium var.
molokaiense, and Solanum nelsonii) are
at risk of habitat destruction and
modification by feral pigs, and the
orangeblack Hawaiian damselfly and six
of the seven yellow-faced bees (all
except Hylaeus longiceps) are at risk of
habitat destruction and modification by
feral pigs (see Table 2).
Goats (Capra hircus)
Feral goats currently destroy and
modify habitat in 10 of the 11
ecosystems (coastal, lowland dry,
lowland mesic, lowland wet, montane
wet, montane mesic, montane dry,
subalpine, dry cliff, and wet cliff) in
which these species occur. Goats, native
to the Middle East and India, were
successfully introduced to the Hawaiian
Islands in the late 1700s. Actions to
control feral goat populations began in
the 1920s (Tomich 1986, pp. 152–153).
However, goats still occupy a wide
variety of habitats on all the main
islands (except for Kahoolawe; see
below), where they consume native
vegetation, trample roots and seedlings,
strip tree bark, accelerate erosion, and
promote the invasion of nonnative
plants (van Riper and van Riper 1982,
pp. 34–35; Stone 1985, p. 261; Kessler
2010, pers. comm.). Kahoolawe was
negatively affected by ungulates
beginning in 1793, with introduction of
goats and the addition of sheep (up to
15,000) and cattle (about 900) by
ranchers between 1858 and 1941, with
the goat population estimated to be as
high as 50,000 individuals by 1988
(KIRC 2014, in litt.; KIRC 2015, in litt.).
Beginning in 1941, the U.S. military
used the entire island as a bombing
range, and in 1994, control of
Kahoolawe was returned to the State
and the Kahoolawe Island Reserve
Commission. The remaining ungulates
were eradicated in 1993 (McLeod 2014,
in litt.). Because they are able to access
extremely rugged terrain, and have a
high reproductive capacity (Clark and
Cuddihy 1980, pp. C–19–C2–20;
Culliney 1988, p. 336; Cuddihy and
Stone 1990, p. 64), goats are believed to
have completely eliminated some plant
species from certain islands (Atkinson
and Atkinson 2000, p. 21). Goats are
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asabaliauskas on DSK3SPTVN1PROD with RULES
highly destructive to native vegetation
and contribute to erosion by: (1) Eating
young trees and young shoots of plants
before they become established; (2)
creating trails that damage native
vegetative cover; (3) destabilizing
substrate and creating gullies that
convey water; and (4) dislodging stones
from ledges that results in rockfalls and
landslides that damage or destroy native
vegetation below (Cuddihy and Stone
1990, pp. 63–64). Feral goats forage
along some cliffs where band-rumped
storm-petrels nest on Kauai, and may
trample nests and increase erosion
(Scott et al. 1986, pp. 8, 352–357;
Tomich 1986, pp. 152–153). The bandrumped storm-petrel and the following
12 plants are at risk from landslides or
erosion caused by feral goats: Gardenia
remyi, Joinvillea ascendens ssp.
ascendens, Kadua fluviatilis, Labordia
lorenciana, Lepidium orbiculare,
Phyllostegia helleri, P. stachyoides,
Portulaca villosa, Pseudognaphalium
sandwicensium var. molokaiense,
Ranunculus mauiensis, Sanicula
sandwicensis, and Schiedea pubescens.
Twenty-four of the 39 plants (all except
for Calamagrostis expansa, Cyanea
kauaulaensis, Cyclosorus boydiae,
Cyperus neokunthianus, Deparia
kaalaana, Dryopteris glabra var. pusilla,
Hypolepis hawaiiensis var. mauiensis,
Kadua haupuensis, Phyllostegia
brevidens, Pritchardia bakeri,
Ranunculus hawaiensis, Schiedea
diffusa ssp. diffusa, Sicyos
macrophyllus, Solanum nelsonii, and
Stenogyne kaalae ssp. sherffii), the
band-rumped storm-petrel, the
orangeblack Hawaiian damselfly, and
four of the yellow-faced bees (Hylaeus
anthracinus, H. assimulans, H. facilis,
and H. hilaris) are at risk of habitat
destruction and modification by feral
goats.
Axis Deer (Axis axis)
Axis deer destroy and modify 6 of the
11 ecosystems (coastal, lowland dry,
lowland mesic, lowland wet, montane
wet, and montane mesic) in which these
species are found. Axis deer were
introduced to the Hawaiian Islands for
hunting opportunities on Molokai in
1868, on Lanai in 1920, and on Maui in
1959 (Hobdy 1993, p. 207; Erdman
1996, pers. comm. in Waring 1996, in
litt., p. 2; Hess 2008, p. 2). Axis deer are
primarily grazers, but also browse
numerous palatable plant species
including those grown as commercial
crops (Waring 1996, p. 3; Simpson 2001,
in litt.). They prefer the low, openly
vegetated areas for browsing and
grazing, but during episodes of drought
(e.g., from 1998 to 2001 on Maui
(Medeiros 2010, pers. comm.)), axis deer
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move into urban and forested areas in
search of food (Waring 1996, p. 5;
Nishibayashi 2001, in litt.). Like goats,
axis deer are highly destructive to native
vegetation and contribute to erosion by
eating young trees and young shoots of
plants before they can become
established. Other axis deer impacts
include stripping bark from mature
trees, creating trails, promoting erosion
by destabilizing substrate, creating
gullies that convey water, and
dislodging stones from ledges that can
result in rockfalls and landslides that
directly damage vegetation (Cuddihy
and Stone 1990, pp. 63–64).
On Molokai, axis deer likely occur at
all elevations from sea level to almost
5,000 ft (1,500 m) at the summit area
(Kessler 2011, pers. comm.). The most
current population estimate for axis
deer on the island of Molokai is between
4,000 and 5,000 individuals (Anderson
2003, p. 119). Little management for
deer control has been implemented on
Molokai, and this figure from more than
a decade ago is likely an underestimate
of the axis deer population on this
island today (Scott et al. 1986, p. 360;
Anderson 2003, p. 30; Hess 2008, p. 4).
On Lanai, axis deer were reported to
number approximately 6,000 to 8,000
individuals in 2007 (The Aloha Insider
2008, in litt; WCities 2010, in litt.). On
Maui, five adult axis deer were released
east of Kihei in 1959 (Hobdy 1993, p.
207; Hess 2008, p. 2). In 2013, the Maui
Axis Deer Working Group estimated that
there may be 8,000 deer on southeast
Maui alone, based on helicopter surveys
(Star Advertiser 2015, in litt.; Hawaii
News Now 2014, in litt.) According to
Medeiros (2010, pers. comm.) axis deer
can be found in all but high-elevation
ecosystems (subalpine and alpine) and
montane bogs on Maui, and are
increasing in numbers at such high rates
that native forests are changing in
unprecedented ways. Additionally,
Medeiros (2010, pers. comm.) asserted
that native plants will only survive in
habitat that is fenced or otherwise
protected from the browsing and
trampling effects of axis deer. Kessler
(2010, pers. comm.) and Hess (2010,
pers. comm.) reported the presence of
axis deer up to 9,000 ft (2,700 m) on
Maui, and Kessler suggests that no
ecosystem is safe from the negative
impacts of these animals. Montane bogs
are also susceptible to impacts from axis
deer. As the native vegetation is
removed by browsing and trampling, the
soil dries out, and nonnative plants
invade. Eventually, the bog habitat and
its associated native plants and animals
are replaced by grassland or shrubland
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dominated by nonnative plants
(Mitchell et al. 2005, p. 6–32).
While axis deer are allowed as game
animals on these three islands, the State
does not permit their introduction to
other Hawaiian Islands. In 2010–2011,
axis deer were illegally introduced to
Hawaii Island as a game animal (Kessler
2011, pers. comm.; Aila 2012, in litt.),
and deer have now been observed across
the southern portion of the island
including in Kohala, Kau, Kona, and
Mauna Kea (HDLNR 2011, in litt.). The
Hawaii Department of Lands and
Natural Resources (HDLNR) Division of
Forestry and Wildlife (HDOFAW) has
developed a response-and-removal plan,
including a partnership now underway
with the Hawaii Department of
Agriculture (HDOA), the Big Island
Invasive Species Committee (BIISC),
Federal natural resource management
agencies, ranchers, farmers, private
landowners, and concerned citizens (Big
Island.com, June 6, 2011). Also, in
response to the introduction of axis deer
to Hawaii Island, the Hawaii Invasive
Species Council drafted House Bill 2593
to amend House Revised Statutes
(H.R.S.) 91, which allows agencies to
adopt emergency rules in the instances
of imminent peril to public health,
including to livestock and poultry
health (BigIsland.com 2011, in litt.;
Martin 2012, in litt.). This emergency
rule became permanent on June 21,
2012, when House Bill 2593 was
enacted into law as Act 194 (State of
Hawaii 2012, in litt.).
The following 16 species in this rule
are at risk from the activities of axis
deer: Gardenia remyi, Huperzia
stemmermanniae, Joinvillea ascendens
ssp. ascendens, Nothocestrum
latifolium, Phyllostegia stachyoides,
Portulaca villosa, Pseudognaphalium
sandwicensium var. molokaiense,
Ranunculus mauiensis, Schiedea
pubescens, Solanum nelsonii, the
orangeblack Hawaiian damselfly, and
five of the yellow-faced bees (Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, and H. longiceps).
Black-Tailed Deer (Odocoileus
hemionus columbianus)
Black-tailed deer destroy and modify
habitat in 5 of the 11 ecosystems
(lowland mesic, lowland wet, montane
wet, montane mesic, and dry cliff) in
which these species occur. The blacktailed deer is one of nine subspecies of
mule deer (Natural History Museum
2015, in litt.). Black-tailed deer were
first introduced to Kauai in 1961, for the
purpose of sport hunting (Tomich 1986,
pp. 131–134). Currently, these deer are
only known from the western side of the
island, where they feed on a variety of
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native (e.g., Acacia koa and Coprosma
spp.) and nonnative plants (van Riper
and van Riper 1982, pp. 42–46; Tomich
1986, p. 134). In addition to their direct
impacts on native plants (browsing),
black-tailed deer likely affect native
plants indirectly by serving as a primary
vector for the spread of introduced
plants by carrying their seeds or other
propagules on their coats and hooves
and in feces. Black-tailed deer have
been noted as a cause of habitat
alteration in the Kauai ecosystems
(NTBG 2007, in litt.; HBMP 2010).
Seven of the 39 plants (Asplenium
diellaciniatum, Dryopteris glabra var.
pusilla, Joinvillea ascendens ssp.
ascendens, Labordia lorenciana,
Nothocestrum latifolium, Ranunculus
mauiensis, and Sicyos lanceoloideus)
are at risk of habitat destruction and
modification by black-tailed deer.
Sheep (Ovis aries)
Four of the ecosystems on Hawaii
Island (lowland dry, lowland mesic,
montane mesic, and montane dry) in
which these species occur are currently
threatened by habitat destruction and
modification due to the activities of
feral sheep. Sheep were introduced to
Hawaii Island in 1791, when Captain
Vancouver brought five rams and two
ewes from California (Tomich 1986, pp.
156–163). Soon after, stock was brought
from Australia, Germany, and the
Mediterranean for sheep production
(Tomich 1986, pp. 156–163; Cuddihy
and Stone 1990, pp. 65–66), and by the
early 1930s, herds reached close to
40,000 individuals (Scowcroft and
Conrad 1992, p. 627). Capable of
acquiring the majority of their water
needs by consuming vegetation, sheep
can inhabit dry forests in remote regions
of the mountains of Mauna Kea and
Mauna Loa, including the saddle
between the two volcanoes. Feral sheep
browse and trample native vegetation
and have decimated large areas of native
forest and shrubland on Hawaii Island
(Tomich 1986, pp. 156–163; Cuddihy
and Stone 1990, pp. 65–66). Browsing
results in the erosion of top soil that
alters moisture regimes and microenvironments, leading to the loss of
native plants and animals (Tomich
1986, pp. 156–163; Cuddihy and Stone
1990, pp. 65–66). In addition, nonnative
plant seeds are dispersed into native
forest by adhering to sheep’s wool coats
(DOFAW 2002, p. 3). In 1962, game
hunters intentionally crossbred feral
sheep with mouflon sheep and released
them on Mauna Kea, where they have
done extensive damage to the montane
dry ecosystem (Tomich 1986, pp. 156–
163). Over the past 30 years, attempts to
protect the vegetation of Mauna Kea and
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the saddle area between the two
volcanoes have been only sporadically
effective (Hess 2008, pp. 1, 4).
Currently, a large population of sheep
(and mouflon hybrids) extends from
Mauna Kea into the saddle and northern
part of Mauna Loa, including State
forest reserves, where they trample and
browse all vegetation, including
endangered plants (Hess 2008, p. 1).
One study estimated as many as 2,500
mouflon within just the Kau district of
the Kahuku Unit (Volcanoes National
Park) in 2006 (Hess et al. 2006, p. 10).
Two of the 39 plants, Exocarpos
menziesii and Festuca hawaiiensis, and
the yellow-faced bee Hylaeus
anthracinus, are reported to be at risk of
habitat destruction and modification by
feral sheep (see Table 2).
Mouflon (Ovis gmelini musimon)
Mouflon destroy and modify habitat
in 6 of the 11 ecosystems on Maui,
Lanai, and Hawaii Island (lowland dry,
lowland mesic, montane mesic,
montane dry, subalpine, and dry cliff) in
which these species occur. Native to
central Asia, mouflon were introduced
to the islands of Lanai and Hawaii in the
1950s as game species, and are now
widely established on these islands
(Tomich 1986, pp. 163–168; Cuddihy
and Stone 1990, p. 66; Hess 2008, p. 1).
Due to their high reproductive rate, the
original population of 11 mouflon on
the island of Hawaii increased to more
than 2,500 individuals in 36 years, even
though they were hunted for game (Hess
2008, p. 3). Mouflon have decimated
vast areas of native shrubland and forest
through grazing, browsing, and bark
stripping (Stone 1985, p. 271; Cuddihy
and Stone 1990, pp. 63, 66; Hess 2008,
p. 3). Mouflon also create trails and
pathways through vegetation, resulting
in soil compaction and increased runoff
and erosion. In some areas, the
interaction of browsing and soil
compaction has led to a shift from
native forest to grassy scrublands (Hess
2008, p. 3). Mouflon only gather in
herds when breeding, thus complicating
control techniques and hunting
efficiency (Hess 2008, p. 3; Ikagawa
2011, in litt.). Currently, many of the
current and proposed fence exclosures
on Hawaii Island constructed to protect
rare species and habitat are designed to
exclude feral pigs, goats, and sheep and
are only 4 ft (1.3 m) in height; a fence
height of at least 6 ft (2 m) is necessary
to exclude mouflon (Ikagawa 2011, in
litt.). Five of the 39 plant species
(Exocarpos menziesii, Nothocestrum
latifolium, Portulaca villosa,
Ranunculus hawaiensis, and Sicyos
macrophyllus), and the yellow-faced bee
Hylaeus assimulans, are at risk from
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habitat destruction and modification
resulting from the activities of mouflon
(see Table 2).
Cattle (Bos taurus)
Cattle destroy and modify habitat in 7
of the 11 ecosystems on Maui and
Hawaii Island (coastal, lowland dry,
lowland mesic, lowland wet, montane
wet, montane mesic, and montane dry)
in which these species occur. Cattle, the
wild progenitors of which were native
to Europe, northern Africa, and
southwestern Asia, were introduced to
the Hawaiian Islands in 1793, and large
feral herds (as many as 12,000 on the
island of Hawaii) developed as a result
of restrictions on killing cattle decreed
by King Kamehameha I (Cuddihy and
Stone 1990, p. 40). While small cattle
ranches were developed on Kauai,
Oahu, Molokai, west Maui, and
Kahoolawe, very large ranches of tens of
thousands of acres were created on east
Maui and Hawaii Island (Stone 1985,
pp. 256, 260; Broadbent 2010, in litt.).
Feral cattle can be found today on the
islands of Molokai, Maui, and Hawaii.
Feral cattle eat native vegetation,
trample roots and seedlings, cause
erosion, create disturbed areas into
which alien plants invade, and spread
seeds of alien plants carried in their
feces and on their bodies. The forest in
areas grazed by cattle rapidly degrades
into grassland pasture, and plant cover
remains reduced for many years
following removal of cattle from an area.
Increased nitrogen availability through
the feces of cattle contributes to the
ingress of nonnative plant species
(Kohala Mountain Watershed
Partnership (KMWP) 2007, pp. 54–55;
Laws et al. 2010, in litt.). Furthermore,
several alien grasses and legumes
purposely introduced for cattle forage
have become invasive weeds (Tomich
1986, pp. 140–150; Cuddihy and Stone
1990, p. 29). According to Kessler (2011,
pers. comm.) approximately 300
individuals roam east Maui as high as
the subalpine ecosystem (i.e., to 9,800 ft
(3,000 m)), and feral cattle are
occasional observed on west Maui. Feral
cattle (more than 100 individuals) are
reported from remote regions of Hawaii
Island, including the back of Pololu and
Waipio Valleys in the Kohala
Mountains, and the Kona Unit of the
Hakalau Forest National Wildlife Refuge
(NWR) (KMWP 2007, p. 55; USFWS
2010, pp. 3–15, 4–86). Nine of the 39
plant species (Huperzia
stemmermanniae, Nothocestrum
latifolium, Ochrosia haleakalae,
Portulaca villosa, Ranunculus
hawaiensis, R. mauiensis, Schiedea
pubescens, Sicyos macrophyllus, and
Solanum nelsonii) and four of the
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yellow-faced bees (Hylaeus anthracinus,
H. assimulans, H. facilis, and H. hilaris)
are currently at risk of habitat
destruction or modification due to the
activities of feral cattle.
In summary, 37 of the 39 plant
species (all except Cyanea kauaulaensis
and Hypolepis hawaiiensis var.
mauiensis), and 9 of the 10 animals
(except for the anchialine pool shrimp
Procaris hawaiana), are at risk of habitat
destruction and modification by
ungulates including pigs, goats, axis
deer, black-tailed deer, sheep, mouflon,
and cattle (see Table 2). The effects of
these nonnative animals include the
destruction of vegetative cover,
trampling of plants and seedlings, direct
consumption of native vegetation, soil
disturbance and sedimentation (erosion
and landslides), dispersal of nonnative
plant seeds by animals, alteration of soil
nitrogen availability, and creation of
open, disturbed areas conducive to
further invasion by nonnative pest plant
species. All of these impacts also can
lead to the conversion of a native plant
community to one dominated by
nonnative species (see ‘‘Habitat
Destruction and Modification by
Nonnative Plants,’’ below). In addition,
because these animals inhabit terrain
that is often steep and remote, foraging
and trampling contributes to severe
erosion of watersheds and degradation
of streams and wetlands (Cuddihy and
Stone 1990, p. 59; Dunkell et al. 2011,
pp. 175–194).
Habitat Destruction and Modification by
Nonnative Plants
Ten of the 11 ecosystems (excluding
anchialine pool ecosystem) and the
species in this rule that are associated
with them are currently at risk of habitat
destruction and modification by
nonnative plants. Native vegetation on
all of the main Hawaiian Islands has
undergone extreme alteration because of
past and present land management
practices, including ranching, deliberate
introduction of nonnative plants and
animals, and agriculture (Cuddihy and
Stone 1990, pp. 27, 58). The original
native flora of Hawaii (present before
human arrival) consisted of about 1,000
taxa, 89 percent of which are endemic
(Wagner et al. 1999, pp. 3–6). Over 800
plant taxa have been introduced to the
Hawaiian Islands. These were brought
to Hawaii for food or for cultural
reasons, to reforest areas destroyed by
grazing feral and domestic animals, or
for horticultural or agricultural
purposes; some were introduced
unintentionally (Scott et al. 1986, pp.
361–363; Cuddihy and Stone 1990, p.
73). Individual descriptions of 114
nonnative plant species that negatively
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affect the 49 species are provided in our
proposed rule (80 FR 58820, September
30, 2015; see pp. 58869–58881).
Fourteen of these nonnative plants are
included in the Hawaii Noxious Weed
List (Hawaii Department of Agriculture
HAR 1981-title 4, subtitle 6, chapter 68).
Nonnative plants adversely affect
native habitat in Hawaii by (1)
modifying the availability of light, (2)
altering soil-water regimes, (3)
modifying nutrient cycling, and (4)
altering fire regimes of native plant
communities (i.e., the ‘‘grass/fire cycle’’
that converts native-dominated plant
communities to nonnative plant
communities; see below) (Smith 1985,
pp. 180–181; Cuddihy and Stone 1990,
p. 74; D’Antonio and Vitousek 1992, p.
73; Vitousek et al. 1997, p. 6). The
contribution of nonnative plants to the
extinction of native species in the
lowland and upland habitats of Hawaii
is well-documented (Vitousek et al.
1987 in Cuddihy and Stone 1990, p. 74).
The most commonly observed effect of
nonnative plants on native species is
displacement through competition.
Competition occurs for water or
nutrients, or it may involve allelopathy
(chemical inhibition of growth of other
plants), shading, or precluding sites for
seedling establishment (Vitousek et al.
1987 in Cuddihy and Stone 1990, p. 74).
Alteration of fire regimes represents
an ecosystem-level change caused by
the invasion of nonnative plants,
primarily grasses (D’Antonio and
Vitousek 1992, p. 73). Grasses generate
standing dead material that burns
readily, and grass tissues with large
surface-to-volume ratios dry out
quickly, contributing to flammability
(D’Antonio and Vitousek 1992, p. 73).
The finest size classes of grass material
ignite and spread fires under a broader
range of conditions than do woody fuels
or even surface litter (D’Antonio and
Vitousek 1992, p. 73). The grass life
form allows rapid recovery following
fire because there is little above-ground
vegetative structure. Grasslands also
support a microclimate in which surface
temperatures are hotter, contributing to
drier vegetative conditions that favor
fire (D’Antonio and Vitousek 1992, p.
73). In summary, nonnative plants
directly and indirectly affect the 39
plants and 9 of the 10 animals in this
rule (except the anchialine pool shrimp)
by destroying and modifying their
habitat, by removing their native host
plants, or by direct competition.
Habitat Destruction and Modification by
Fire
Seven of the 11 ecosystems (coastal,
lowland dry, lowland mesic, montane
mesic, montane dry, subalpine, and dry
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cliff) and the species in this rule that are
associated with them are at risk of
destruction and modification by fire.
Fire is an increasing, humanexacerbated threat to native species and
ecosystems in Hawaii. The presettlement fire regime in Hawaii was
characterized by infrequent, lowseverity events, as few natural ignition
sources existed (Cuddihy and Stone
1990, p. 91; Smith and Tunison 1992,
pp. 395–397). It is believed that prior to
human colonization fuel was sparse in
wet plant communities and only
seasonally flammable in mesic and dry
plant communities. The only ignition
sources were volcanism and lightning
(Baker et al. 2009, p. 43). Although Vogl
(1969, in Cuddihy and Stone 1990, p.
91) proposed that naturally occurring
fires may have been important in the
development of some of the original
Hawaiian flora, Mueller-Dombois (1981,
in Cuddihy and Stone 1990, p. 91)
asserts that most natural vegetation
types of Hawaii would not carry fire
before the introduction of alien grasses.
Smith and Tunison (in Cuddihy and
Stone 1990, p. 91) state that native plant
fuels typically have low flammability.
Existing fuel loads were often
discontinuous, and rainfall in many
areas on most islands was moderate to
high. Fires inadvertently or
intentionally set by the Polynesian
settlers probably contributed to the
initial decline of native vegetation in the
drier plains and foothills. These early
settlers practiced slash-and-burn
agriculture that created open lowland
areas suitable for the opportunistic
invasion and colonization of nonnative,
fire-adapted grasses (Kirch 1982, pp. 5–
6, 8; Cuddihy and Stone 1990, pp. 30–
31). Beginning in the late 18th century,
Europeans and Americans introduced
plants and animals that further
degraded native Hawaiian ecosystems.
Ranching and the creation of
pasturelands in particular created
highly fire-prone areas of nonnative
grasses and shrubs (D’Antonio and
Vitousek 1992, p. 67). Although fires
were infrequent in mountainous
regions, extensive fires have recently
occurred in lowland dry and lowland
mesic areas, leading to grass/fire cycles
that convert native dry forest and native
wet forest to nonnative grassland
(D’Antonio and Vitousek 1992, p. 77).
Because of the greater frequency,
intensity, and duration of fires that have
resulted from the human alteration of
landscapes and the introduction of
nonnative plants, especially grasses,
fires are now more destructive to native
Hawaiian ecosystems (Brown and Smith
2000, p. 172), and a single grass-fueled
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fire often kills most native trees and
shrubs in the area (D’Antonio and
Vitousek 1992, p. 74). Fire destroys
dormant seeds of native plants, as well
as individual plants and animals
themselves, even in steep, inaccessible
areas or near streams and ponds.
Successive fires remove habitat for
native species by altering microclimate
conditions, creating conditions more
favorable to nonnative plants.
Nonnative grasses (e.g., Cenchrus
setaceus; fountain grass), many of which
may be fire-adapted, produce a high fuel
load that allow fire to burn areas that
would not otherwise burn easily,
regenerate quickly after fire, and
establish rapidly in burned areas
(Fujioka and Fujii 1980 in Cuddihy and
Stone 1990, p. 93; D’Antonio and
Vitousek 1992, pp. 70, 73–74; Tunison
et al. 2002, p. 122). Native woody plants
may recover to some degree, but fire tips
the competitive balance toward
nonnative species (National Park
Service 1989 in Cuddihy and Stone
1990, p. 93). During a post-burn survey
on Hawaii Island, in an area of native
Diospyros forest with undergrowth of
the nonnative grass Pennisetum
setaceum [Cenchrus setaceus], Takeuchi
(1991, p. 2) noted that ‘‘no regeneration
of native canopy is occurring within the
Puuwaawaa burn area.’’ Takeuchi also
stated that ‘‘burn events served to
accelerate a decline process already in
place, compressing into days a sequence
which would ordinarily have taken
decades’’ (Takeuchi 1991, p. 4), and
concluded that, in addition to
increasing the number of fires, the
nonnative Pennisetum acted to suppress
establishment of native plants after a
fire (Takeuchi 1991, p. 6).
For many decades, fires have affected
rare or endangered species and their
habitats on Molokai, Lanai, and Maui
(Gima 1998, in litt.; Hamilton 2009, in
litt.; Honolulu Advertiser 2010, in litt.;
Pacific Disaster Center 2011, in litt.).
These three islands experienced
approximately 1,290 brush fires
between 1972 and 1999 that burned a
total of 64,250 ac (26,000 ha) (County of
Maui 2009, ch. 3, p. 3; Pacific Disaster
Center 2011, in litt.). Between 2000 and
2003, the annual number of wildfires on
these islands jumped from 118 to 271;
of these, several burned more than 5,000
ac (2,023 ha) each (Pacific Disaster
Center 2011, in litt.). On Molokai,
between 2003 and 2004, three wildfires
each burned 10,000 ac (4,050 ha)
(Pacific Disaster Center 2011, in litt.).
From August through early September
2009, a wildfire burned approximately
8,000 ac (3,237 ha), including 600 ac
(243 ha) of the remote Makakupaia
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section of the Molokai Forest Reserve, a
small portion of The Nature
Conservancy’s (TNC’s) Kamakou
Preserve, and encroached on Onini
Gulch, Kalamaula, and Kawela
(Hamilton 2009, in litt.). Species at risk
because of wildfire on Molokai include
the plants Joinvillea ascendens ssp.
ascendens, Nothocestrum latifolium,
Portulaca villosa, Ranunculus
mauiensis, Schiedea pubescens, and
Solanum nelsonii, and the yellow-faced
bees Hylaeus anthracinus, H. facilis, H.
hilaris, and H. longiceps.
Several wildfires have occurred on
Lanai in the last decade. In 2006, a
wildfire burned 600 ac (243 ha) between
Manele Road and the Palawai Basin,
about 3 mi (4 km) south of Lanai City
(The Maui News 2006, in litt.). In 2007,
a brush fire at Mahana burned about 30
ac (12 ha), and in 2008, another 1,000
ac (405 ha) were burned by wildfire in
the Palawai Basin (The Maui News
2007, in litt.; KITV Honolulu 2008, in
litt.). Species at risk because of wildfire
on Lanai include Exocarpos menziesii,
Nothocestrum latifolium, Portulaca
villosa, Schidea pubescens; and the
yellow-faced bees Hylaeus anthracinus,
H. assimulans, H. facilis, H. hilaris, and
H. longiceps.
On west Maui, wildfires burned more
than 8,650 ac (3,501 ha) between 2007
and 2010 (Honolulu Advertiser 2010, in
litt.; Shimogawa 2010, in litt.). These
fires encroached into the West Maui
Forest Reserve, on the ridges of Olowalu
and Kealaloloa, which is habitat for
several endangered plants. In 2007, on
east Maui, a fire consumed over 600 ac
(240 ha), increasing invasion of the area
by nonnative plants (Pinus spp.) (Pacific
Disaster Center 2007, in litt.; The Maui
News 2011, in litt.). Species at risk
because of wildfire on west and east
Maui include the plants Festuca
hawaiiensis, Joinvillea ascendens ssp.
ascendens, Nothocestrum latifolium,
Ochrosia haleakalae, Portulaca villosa,
Ranunculus mauiensis, Sanicula
sandwicensis, Schiedea pubescens,
Sicyos macrophyllus, and Solanum
nelsonii, and the yellow-faced bees
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, and H. longiceps.
Several recent fires on Oahu in the
Waianae Mountain range have affected
rare and endangered species. Between
2004 and 2005, wildfires burned more
than 360 ac (146 ha) in Honouliuli
Preserve, habitat of more than 90 rare
and endangered plants and animals
(TNC 2005). In 2006, a fire at Kaena
Point State Park burned 60 ac (24 ha),
and encroached on endangered plants in
Makua Military Training Area. In 2007,
there was a significant fire at
Kaukonahua that crossed 12 gulches,
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eventually encompassing 5,655 ac
(2,289 ha) that negatively affected eight
endangered plant species and their
habitat (Abutilon sandwicense, Bonamia
menziesii, Colubrina oppositifolia,
Eugenia koolauensis, Euphorbia
haeleeleana, Hibiscus brackenridgei ssp.
mokuleianus, Nototrichium humile, and
Schiedea hookeri) (U.S. Army Garrison
2007b, Appendices pp. 1–5). This fire
provided ingress for nonnative
ungulates (cattle, goats, and pigs) into
previously undisturbed areas, and
opened dense native vegetation to the
invasive grass Urochloa maxima
(Panicum maximum, guinea grass), also
a food source for cattle and goats. The
grass was observed to generate blades
over 2 ft (0.6 m) in length only 2 weeks
following the fire (U.S. Army Garrison
2007b, Appendices pp. 1–5). In 2009,
two smaller fires burned 200 ac (81 ha)
at Manini Pali (Kaena Point State Park)
and almost 4 ac (1.5 ha) at Makua Cave.
Both of these fires burned into area
designated as critical habitat, although
no individual plants were directly
affected (U.S. Army Natural Resource
Program 2009, Appendix 2, 17 pp.).
Most recently, in 2014, two fires
affected native forest, one in the Oahu
Forest National Wildlife Refuge (350 ac,
140 ha), on the leeward side of the
Koolau Mountains (DLNR 2014, in litt.),
and one above Makakilo, in the Waianae
Mountains, just below Honouliuli FR,
that burned more than 1,000 ac (400 ha)
(KHON 2014, in litt.). The Makakilo fire
took over 2 weeks to contain. Species at
risk because of wildfire on Oahu
include the plants Joinvillea ascendens
ssp. ascendens, Nothocestrum
latifolium, Portulaca villosa,
Ranunculus mauiensis, and Sicyos
lanceoloideus, and the yellow-faced
bees Hylaeus anthracinus, H.
assimulans, H. facilis, H. kuakea, H.
longiceps, and H. mana.
In 2012, on Kauai, a wildfire that was
possibly started by an unauthorized
camping fire burned 40 ac (16 ha) in the
Na Pali-Kona Forest Reserve on Milolii
Ridge, forcing closure of a hiking trail.
Fortunately, several endangered and
threatened plants in the adjacent Kula
NAR were not impacted (KITV 2012, in
litt.). The same year, another wildfire
burned over 650 ac (260 ha) on Hikimoe
Ridge, and threatened the Puu Ka Pele
section of Waimea Canyon State Park
(Hawaii News Now 2012, in litt.; Star
Advertiser 2012, in litt.). Species at risk
of because wildfire on Kauai include the
plants Joinvillea ascendens ssp.
ascendens, Labordia lorenciana,
Nothocestrum latifolium, Ranunculus
mauiensis, Santalum involutum, and
Sicyos lanceoloideus.
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In the driest areas on the island of
Hawaii, wildfires are exacerbated by the
uncontrolled growth of nonnative
grasses such as Cenchrus setaceus (Fire
Science Brief 2009, in litt.). Since its
introduction to the island in 1917, this
grass now covers more than 200 square
mi (500 square km) of the leeward areas
of the island (Joint Fire Science Brief
(JFSB) 2009, in litt.). In the past 50
years, three wildfires on the leeward
side encompassed a total of 30,000 ac
(12,140 ha) (JFSB 2009, in litt.). These
wildfires traveled great distances at
rates of 4 to 8 miles per hour (mph) (7
to 12 kilometers per hour (kph)),
burning 2.5 ac (1 ha) to 6 ac (2.5 ha) per
minute (the equivalent of 6 to 8 football
fields per minute) (Burn Institute 2009,
p. 4). Between 2002 and 2003, three
successive lava-ignited wildfires in the
east rift zone of Hawaii Volcanoes
National Park affected native forests in
lowland dry, lowland mesic, and
lowland wet ecosystems (JFSB 2009, p.
3), cumulatively burning an estimated
11,225 ac (4,543 ha) (Wildfire News,
June 9, 2003; JFSP 2009, p. 3). These
fires destroyed over 95 percent of the
canopy cover and encroached upon
forest areas that were previously
thought to have low susceptibility to
wildfires. After the fires, nonnative
ferns were observed in higher elevation
rainforest where they had not
previously been seen, and were believed
to inhibit the recovery of the native
Metrosideros polymorpha (ohia) trees
(JFSP 2003, pp. 1–2). Nonnative grasses
invaded the burn area, increasing the
risk of fire encroaching into the
surrounding native forest (Ainsworth
2011, in litt.). Extreme drought
conditions also contributed to the
number and intensity of wildfires on
Hawaii Island (Armstrong and Media
2010, in litt.; Loh 2010, in litt.). This
‘‘extreme’’ drought classification for
Hawaii was recently lifted to
‘‘moderate’’; however, drier than
average conditions persist, and another
extreme drought event may occur
(NOAA 2015, in litt.). In addition, El
˜
Nino conditions in the Pacific (see
‘‘Climate Change’’ under Factor E,
below), a half-century of decline in
annual rainfall, and intermittent dry
spells have contributed to the
conditions favoring wildfires in all the
main Hawaiian Islands (Marcus 2010, in
litt.). Species at risk because of wildfire
on Hawaii Island include the plants
Exocarpos menziesii, Festuca
hawaiiensis, Joinvillea ascendens ssp.
ascendens, Ochrosia haleakalae,
Portulaca villosa, Ranunculus
mauiensis, Sanicula sandwicensis,
Sicyos macrophyllus, and Solanum
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nelsonii, and the yellow-faced bee
Hylaeus anthracinus.
In summary, fire is a threat to 14 plant
species and their habitat (Exocarpos
menziesii, Festuca hawaiiensis,
Joinvillea ascendens ssp. ascendens,
Labordia lorenciana, Nothocestrum
latifolium, Ochrosia haleakalae,
Portulaca villosa, Ranunculus
mauiensis, Sanicula sandwicensis,
Santalum involutum, Schiedea
pubescens, Sicyos lanceoloideus, S.
macrophyllus, and Solanum nelsonii),
and all seven yellow-faced bees because
these species and their habitat are
located in or near areas that were
burned previously, or in areas
considered at risk because of fire due to
the cumulative and compounding
effects of drought and the presence of
highly flammable nonnative grasses.
Habitat Destruction and Modification by
Hurricanes
Ten of the 11 ecosystems (all except
the anchialine pool ecosystem) where
these species occur are at risk of habitat
destruction and modification by
hurricanes. Hurricanes exacerbate the
impacts of other threats such as habitat
destruction and modification by
ungulates and competition with
nonnative plants. By destroying native
vegetation, hurricanes open the forest
canopy, modify the availability of light,
and create disturbed areas conducive to
invasion by nonnative pest species (see
‘‘Habitat Destruction and Modification
by Nonnative Plants’’, above) (Asner
and Goldstein 1997, p. 148; Harrington
et al. 1997, pp. 539–540). In addition,
hurricanes adversely affect native
Hawaiian stream habitat by defoliating
and toppling vegetation, thus loosening
the surrounding soil and increasing
erosion. Along with catastrophic
flooding, this soil and vegetative debris
can be washed into streambeds (by
hurricane-induced rain or subsequent
rain storms), resulting in the scouring of
stream bottoms and channels (Polhemus
1993a, 88 pp.). Natural disasters such as
hurricanes can be particularly
devastating to Hawaiian plant and
animal species that persist in low
numbers and in restricted ranges
(Mitchell et al. 2005, p. 4–3).
Hurricanes affecting Hawaii were only
rarely reported from ships in the area
from the 1800s until 1949. Between
1950 and 1997, 22 hurricanes passed
near or over the Hawaiian Islands, 5 of
which caused serious damage (Businger
1998, pp. 1–2). In November 1982,
Hurricane Iwa struck the Hawaiian
Islands with wind gusts exceeding 100
miles per hour (mph) (160 kilometers
per hour (kmh)), causing extensive
damage, especially on the islands of
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Kauai, Niihau, and Oahu (Businger
1998, pp. 2, 6). Many forest trees were
destroyed (Perlman 1992, pp. 1–9),
which opened the canopy and
facilitated the invasion of native forest
by nonnative plants (Kitayama and
Mueller-Dombois 1995, p. 671).
Hurricanes therefore exacerbate the
threats posed by nonnative plants, as
described in ‘‘Habitat Destruction and
Modification by Nonnative Plants,’’
above. In September 1992, Hurricane
Iniki, a category 4 hurricane with
maximum sustained winds of 130 mph
(209 kmh, 113 knots), passed directly
over the island of Kauai and close to the
island of Oahu, causing significant
damage to Kauai and along Oahu’s
southwestern coast (Blake et al. 2007,
pp. 20, 24). Biologists documented
damage to the habitat of six endangered
plant species on Kauai, and one plant
on Oahu. Polhemus (1993a, pp. 86–87)
documented the extirpation of the
scarlet Kauai damselfly (Megalagrion
vagabundum) (a species related to M.
xanthomelas), from the entire
Hanakapiai Stream system on the island
of Kauai as a result of the impacts of
Hurricane Iniki. Damage by future
hurricanes will further alter the
remaining native-plant dominated
habitat for rare plants and animals in
native ecosystems of Kauai, Oahu, and
other Hawaiian Islands (Bellingham et
al. 2005, p. 681) (see ‘‘Climate Change’’
under Factor E. Other Natural or
Manmade Factors Affecting Their
Continued Existence, below).
In summary, hurricanes exacerbate
other habitat threats, such as
competition with nonnative plants, as
well as result in direct habitat
destruction. This is a particular problem
for the plant Pritchardia bakeri, the
band-rumped storm-petrel, the
orangeblack Hawaiian damselfly, and all
seven yellow-faced bees.
Habitat Modification and Destruction
Due to Landslides, Rockfalls, Treefall,
Flooding, Erosion, Drought, and
Tsunamis
Habitat destruction and modification
by landslides, rockfalls, treefall,
flooding, erosion, and drought (singly or
in combination) is a threat to all 11
ecosystems in which these species
occur. Landslides, rockfalls, treefall,
flooding, and erosion change native
plant and animal communities by
destabilizing substrates, damaging or
destroying individual plants, and
altering hydrological patterns. In the
open sea near Hawaii, rainfall averages
25 to 30 inches (in) (630 to 760
millimeters (mm)) per year, yet the
islands may receive up to 15 times this
amount in some places, caused by
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orographic features (topography)
(Wagner et al. 1999, adapted from Price
(1983) and Carlquist (1980), pp. 38–39).
During storms, rain may fall at rates of
3 in (76 mm) per hour or more, and
sometimes may reach nearly 40 in
(1,000 mm) in 24 hours, resulting in
destructive flash-flooding in streams
and narrow gulches (Wagner et al. 1999,
adapted from Price (1983) and Carlquist
(1980), pp. 38–39). Due to the steep
topography in many mountainous areas
on the Hawaiian Islands, disturbance
caused by introduced ungulates
exacerbates erosion and increases the
potential for landslides, rockfalls, or
flooding, which in turn damages or
destroys native plants and disturbs
habitat of the band-rumped storm-petrel
(see Table 2). These events could
eliminate one or more isolated
occurrences of species that persist in
low numbers and a limited geographic
range, resulting in reduced redundancy
and resilience of the species.
Landslides, rockfalls, treefall,
flooding, and erosion are threats to 20
plant species (Cyanea kauaulaensis,
Cyclosorus boydiae, Deparia kaalaana,
Dryopteris glabra var. pusilla, Gardenia
remyi, Joinvillea ascendens ssp.
ascendens, Kadua fluviatilis, K.
haupuensis, Labordia lorenciana,
Lepidium orbiculare, Phyllostegia
brevidens, P. helleri, P. stachyoides,
Portulaca villosa, Pseudognaphalium
sandwicensium var. molokaiense,
Ranunculus hawaiensis, R. mauiensis,
Sanicula sandwicensis, Schiedea
pubescens, and Solanum nelsonii) and
to the band-rumped storm-petrel and
the orangeblack Hawaiian damselfly.
Landslides, rockfalls, and erosion can
directly affect nests and nesting habitat
of the band-rumped storm-petrel.
Destabilization of cliff habitat leads to
additional landslides and alteration of
hydrological patterns, affecting the
availability of soil moisture. Landslides
also destroy and modify riparian and
stream habitat by direct physical
damage, and create disturbed areas
leading to invasion by nonnative plants,
as well as damaging or destroying plants
directly. Kadua haupuensis, Labordia
lorenciana, Lepidium orbiculare,
Phyllostegia brevidens, and P. helleri are
known only from a few individuals in
single occurrences on cliffs or steepwalled stream valleys, and one
landslide could extirpate a species by
direct destruction. Monitoring data
presented by the Plant Extinction
Prevention Program (PEPP) and
botanical surveys suggest that flooding
is a likely threat to eight plant species,
Cyanea kauaulaensis, Cyclosorus
boydiae, Deparia kaalaana, Labordia
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lorenciana, Phyllostegia stachyoides,
Sanicula sandwicensis, Schiedea
pubescens, and Solanum nelsonii, as
some individuals occur on stream banks
(Wood et al. 2007, p. 198; PEPP 2011,
pp. 162–164; Oppenheimer and Lorence
2012, pp. 20–21; PEPP 2013, p. 54; PEPP
2014, pp. 95, 142). The naiad life stage
of the orangeblack Hawaiian damselfly
would be destroyed by flooding if an
individual is carried out of suitable
habitat or into areas occupied by
nonnative fish.
Drought is reported to be a threat to
10 plants (Cyclosorus boydiae, Deparia
kaalaana, Huperzia stemmermanniae,
Phyllostegia stachyoides, Ranunculus
hawaiensis, R. mauiensis, Sanicula
sandwicensis, Schiedea pubescens,
Sicyos lanceoloideus, and Solanum
nelsonii), the orangeblack Hawaiian
damselfly (directly or by desiccation of
streams and ponds), and all seven
yellow-faced bees (Magnacca 2007, pp.
181, 183; Polhemus 2008, p. 26; Chu et
al. 2010, pp. 4887, 4891, 4898; PEPP
2011, pp. 162–164; Fortini et al. 2013,
p. 2; PEPP 2013, p. 177; PEPP 2014, pp.
140–142, 154–156, 162, 166–167).
Between 1860 and 2002, there were 49
periods of drought on Oahu, 30 periods
of drought on Molokai, Lanai, and Maui,
and at least 18 serious or severe drought
events on Hawaii Island (Giambelluca et
al. 1991, pp. 3–4; Hawaii Commission
on Water Resource Management
(CWRM) 2009, in litt.; Hawaii Civil
Defense 2011, pp. 14–1–14–12). The
most severe drought events over the past
15 years were associated with the El
˜
Nino phenomenon (Hawaii Civil
Defense 2011, p. 14–3). In 1998, the city
of Hilo had the lowest January total
rainfall (0.014 in) ever observed for any
month since records have been kept,
with average rainfall being almost 10 in
for January (Hawaii Civil Defense 2011,
p. 14–3). Currently, the State remains
under abnormally dry to moderate
drought conditions, with the onset of
˜
another El Nino event (U.S. Drought
Monitor 2015, in litt., National Weather
Service 2015, in litt.). Drought events
dry up streams, irrigation ditches, and
reservoirs, and deplete groundwater
supplies (Hawaii CWRM 2009, in litt.).
Recent episodes of drought have driven
axis deer farther into forested areas in
search of food, increasing their negative
impacts on native vegetation from
herbivory, bark stripping, and trampling
(see Factor C. Disease or Predation,
below) (Waring 1996, in litt;
Nishibayashi 2001, in litt.). Drought
events could eliminate one or more
isolated populations of a species that
currently persists in low numbers and a
limited geographic range, resulting in
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reduced redundancy and resilience of
the species or extinction.
Tsunamis destroy and modify habitat
for species in Northwestern Hawaiian
Islands and in low-lying coastal areas of
the main Hawaiian Islands. Tsunamis in
Hawaii are caused by earthquakes,
submarine landslides, and volcanic
eruptions that may occur within the
archipelago or in distant parts of the
Pacific. These events disturb the ocean’s
surface, and gravity combined with the
water’s motion produces a series of
long-period waves that travel quickly
and can reach heights of 32 ft (10 m) or
more when reaching land. Major
tsunamis occur worldwide about once
every 10 years, on average, and almost
60 percent of those occur in the Pacific
Ocean (Pacific Tsunami Warning
Center, https://ptwc.weather.gov/ptwc/
faq.php#8, accessed June 2016). In 2011,
a tsunami caused by an earthquake in
Japan reached Hawaii and the
Northwestern Hawaiian Islands. This
tsunami swept over Midway Atoll’s
Eastern Island and Kure Atoll’s Green
Island, where it inundated plants,
spread plastic debris, killed thousands
of seabirds, and destroyed seabird
nesting areas as it traveled about 500 ft
(150 m) inland (DOFAW 2011, in litt.;
Starr 2011, in litt.; USFWS 2011, in
litt.). This threat could occur at any time
and negatively affect occurrences and
habitat of the plant Solanum nelsonii
and the yellow-faced bees Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, and H. longiceps.
Habitat Destruction and Modification by
Water Extraction
Freshwater habitats on all the main
Hawaiian Islands have been severely
altered and degraded because of past
and present land and water management
practices, including agriculture, urban
development, and development of
ground water, perched aquifer, and
surface water resources (Harris et al.
1993, p. 11; Meier et al. 1993, p. 181).
Extensive modification of lentic
(standing water) habitat in the Hawaiian
Islands began about 1100 A.D. with a
rapid increase in the human population
(Harris et al. 1993, p. 9; Kirch 1982, pp.
5–6). Hawaiians cultivated Colocasia
esculenta (kalo, taro) by creating
shallow, walled ponds, or loi, in
marshes and riparian areas (Meier et al.
1993, p. 181; Handy and Handy 1972, p.
58). By 1778, virtually all valley bottoms
with permanent stream flow and most
basin marshes were converted to taro
cultivation (Handy and Handy 1972, pp.
396, 411). Hawaiians also modified
wetlands by constructing fishponds,
many of which were primarily fresh
water, fed by streams or springs (Meier
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et al. 1993, p. 181). Despite this habitat
modification by early Hawaiians, many
areas of extensive marshland remained
intact and were used by the native
damselflies. Over time, however, many
of the wetlands formerly used for taro
were drained and filled for dry-land
agriculture or development (Stone 1989,
p. 129; Meier et al. 1993, pp. 181–182).
In addition, marshes are slowly filled
and converted to meadow habitat due to
increased sedimentation resulting from
increased storm water runoff from
upslope development and blockage of
downslope drainage (Wilson Okamoto
and Associates, Inc. 1993, p. 3–5).
Presently the most significant threat to
the remaining natural ponds and
marshes in Hawaii, habitat for the
orangeblack Hawaiian damselfly, is the
nonnative grass species Urochloa
mutica. This sprawling, perennial grass
was first observed on Oahu in 1924, and
now occurs on all the main islands
(O’Connor 1999, p. 1504). This species
forms dense, monotypic stands that can
completely eliminate any open water by
layering of its trailing stems (Smith
1985, p. 186).
Similar to the loss of wetlands in
Hawaii, the loss of streams has been
significant and began with the early
Hawaiians who modified stream
systems by diverting water to irrigate
taro. However, these Hawaiian-made
diversions were closely regulated and
were not permitted to take more than
half the stream flow, and were typically
used to flood taro loi only periodically
(Handy and Handy 1972, pp. 58–59).
The advent of sugarcane plantations in
1835 led to more extensive stream
diversions. These systems were
typically designed to tap water at upper
elevation sources (above 980 ft (300 m))
by means of concrete weirs. All or most
of the stream flow was diverted into
fields or reservoirs (Takasaki et al. 1969,
p. 65; Harris et al. 1993, p. 10). By the
1930s, major water diversions had been
developed on all the main islands, and
currently one-third of Hawaii’s
perennial streams are diverted (Harris et
al. 1993, p. 10). In addition to diverting
water for agriculture and domestic water
supply, streams have been diverted for
use in producing hydroelectric power
(Hawaii Stream Assessment 1990, p.
96). Surface flow has also been diverted
into channels, and the perched aquifers
which fed the streams have been tapped
by means of tunnels (Stearns and
Vaksvik 1935, pp. 365, 378–434; Stearns
1985, p. 291, 301–303). Many of these
aquifers are the sources of springs,
which contribute flow to streams. The
draining of these aquifers causes springs
to become dry (Stearns and Vaksvik
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1935, pp. 380, 388; USGS 2000, in litt.).
Most remaining streams that are not
already diverted have been, and
continue to be, seriously degraded by
the activities of feral ungulates and by
nonnative plants. Channelization has
not been restricted to lower reaches, and
it results in the loss of riparian
vegetation, increasing flow velocity,
illumination, and water temperature
(Parrish et al. 1984, pp. 83–84). These
conditions make the channels
unsuitable as habitat for the orangeblack
Hawaiian damselfly.
Water extraction (e.g., withdrawal of
subsurface fresh water for development
and human use) from underground fresh
water sources increases salinity levels of
anchialine pools and negatively affect
the anchialine pool shrimp, Procaris
hawaiana, which relies on the delicate
balance of mixohaline (brackish water)
habitats (Conry 2012, in litt.; National
Park Service 2016, in litt.). Water
extraction also negatively affects the
plant Cyclosorus boydiae and the
orangeblack Hawaiian damselfly by
degrading or destroying habitat for these
species (Harris et al. 1993, pp. 9–13;
Medeiros et al. 1993, p. 88; Meier et al.
1993, pp. 181–183; Palmer 2003, p. 88).
Habitat Destruction and Modification by
Climate Change
Climate change affects the habitat of
the 49 species. Discussion of climate
change impacts is included in our
complete discussion of climate change
under Factor E. Other Natural or
Manmade Factors Affecting Their
Continued Existence, below.
Summary of Factor A
Destruction and modification of the
habitat of each of the 49 species
addressed in this rule is occurring
throughout the entire range of each of
the species. These impacts include the
effects of agriculture and urban
development, introduced ungulates,
nonnative plants, fire, hurricanes,
landslides, rockfalls, treefall, flooding,
erosion, drought, tsunamis, and water
extraction.
Habitat destruction and modification
by agriculture and urban development is
an ongoing and serious threat to the
plant Cyclosorus boydiae, the
orangeblack Hawaiian damselfly, the
anchialine pool shrimp Procaris
hawaiana, and the yellow-faced bees
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, and H. longiceps.
Conversion of wetland and other aquatic
habitat (i.e., water extraction) for
agriculture and urban development is
ongoing, is expected to continue into
the future, and affects the orangeblack
Hawaiian damselfly by removing habitat
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required for hunting and breeding.
Water extraction affects the orangeblack
Hawaiian damselfly because it (1)
Reduces the amount and distribution of
stream habitat; (2) reduces stream flow
and habitat; and (3) leads to an increase
in water temperature, which causes
physiological stress to the damselfly
naiads. Water extraction affects the
delicate balance of the anchialine pool
ecosystem, including salinity and biota,
negatively affecting the anchialine pool
shrimp, Procaris hawaiana. Loss of
stream-course habitat affects Cyclosorus
boydiae because this is the only habitat
where this riparian species occurs.
The threat of habitat destruction and
modification by ungulates is ongoing as
ungulates currently occur in all
ecosystems on which these species
depend except the anchialine pool
system. Introduced ungulates pose a
threat to 37 of the 39 plants (except for
Cyanea kauaulaensis and Hypolepis
hawaiiensis var. mauiensis), and 9 of
the 10 animal species (all except for the
anchialine pool shrimp) in this rule that
occur in these 10 ecosystems because
ungulates: (1) Directly affect the species
by trampling and grazing (see Factor C
discussion, below); (2) increase soil
disturbance and erosion; (3) create open,
disturbed areas conducive to nonnative
plant invasion by dispersing fruits and
seeds, which results in conversion of a
native-dominated plant community to a
nonnative-dominated plant community;
and (4) increase marsh and stream
disturbance and sedimentation, which
negatively affects these aquatic habitats.
Habitat destruction and modification
by nonnative plants is a serious and
ongoing current threat to all 39 plant
species because nonnative plants: (1)
Adversely affect microhabitat by
modifying the availability of light; (2)
alter soil-water regimes; (3) modify
nutrient cycling processes; (4) alter fire
ecology, leading to incursions of firetolerant nonnative plant species into
native habitat; (5) outcompete, and
possibly directly inhibit (through
allelopathy) the growth of native plant
species; and (6) alter habitat and
substrate such that erosion leading to
rockfalls and landslides may increase.
Each of these processes can convert
native-dominated plant communities to
nonnative plant communities (Cuddihy
and Stone 1990, p. 74; Vitousek 1992,
pp. 33–35).
The threat of habitat destruction and
modification by fire to 14 plant species
(Exocarpos menziesii, Festuca
hawaiiensis, Joinvillea ascendens ssp.
ascendens, Labordia lorenciana,
Nothocestrum latifolium, Ochrosia
haleakalae, Portulaca villosa,
Ranunculus mauiensis, Sanicula
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sandwicensis, Santalum involutum,
Schiedea pubescens, Sicyos
lanceoloideus, S. macrophyllus, and
Solanum nelsonii) and all seven yellowfaced bee species is serious and ongoing
because fires occur frequently and
damage and destroy native vegetation,
including dormant seeds, seedlings, and
juvenile and adult plants, including
host plants for the bees. Many
nonnative, invasive plants, particularly
fire-tolerant grasses, create more
destructive fires, invade burned areas,
and can out-compete native plants and
inhibit their regeneration (D’Antonio
and Vitousek 1992, pp. 70, 73–74;
Tunison et al. 2002, p. 122). Successive
fires that burn farther and farther into
native habitat destroy the ecosystem and
its components upon which these
species depend.
Habitat destruction and modification
by natural disasters such as hurricanes
represent a serious threat to the plant
Pritchardia bakeri, the band-rumped
storm-petrel, the orangeblack Hawaiian
damselfly, and all seven yellow-faced
bee species. Hurricanes open the forest
canopy, modifying available light and
creating disturbed areas that are
conducive to invasion by nonnative
plants (Asner and Goldstein 1997, p.
148; Harrington et al. 1997, pp. 346–
347). The discussion under ‘‘Habitat
Destruction and Modification by
Nonnative Plants’’ provides additional
information related to canopy gaps, light
availability, and the establishment of
nonnative plant species. In addition,
hurricanes cause mortality of birds,
including adults and chicks drowned
when nest sites are flooded (Schreiber
2002, p. 186; Hass et al. 2012, pp. 252–
253). Hurricanes also destroy nesting
habitat, a particular problem for species
like storm-petrels that return to the
same nest site each year (Schreiber
2002, p. 186). These hurricane impacts
are likely for the band-rumped stormpetrel. Finally, hurricanes can alter and
directly damage streams and wetlands
used by the orangeblack Hawaiian
damselfly (Polhemus 1993a, pp. 86–87).
The impacts from hurricanes can be
particularly devastating to these species
because they persist in low numbers in
restricted ranges and are therefore less
resilient to such disturbances. A single
destructive hurricane holds the
potential of driving to extinction species
that persist as one or several small,
isolated populations.
Landslides, rockfalls, treefalls,
flooding, and erosion (singly or
combined) are a threat to 20 plant
species (Cyanea kauaulaensis,
Cyclosorus boydiae, Deparia kaalaana,
Dryopteris glabra var. pusilla, Gardenia
remyi, Joinvillea ascendens ssp.
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ascendens, Kadua fluviatilis, K.
haupuensis, Labordia lorenciana,
Lepidium orbiculare, Phyllostegia
brevidens, P. helleri, P. stachyoides,
Portulaca villosa, Pseudognaphalium
sandwicensium var. molokaiense,
Ranunculus hawaiensis, R. mauiensis,
Sanicula sandwicensis, Schiedea
pubescens, and Solanum nelsonii), the
band-rumped storm-petrel, and the
orangeblack Hawaiian damselfly by
destabilizing substrates, damaging and
killing individuals, altering hydrological
patterns, and destroying or modifying
habitat—all resulting in changes to
native plant and animal communities.
Drought is a threat to 10 plant species
(Cyclosorus boydiae, Deparia kaalaana,
Huperzia stemmermanniae, Phyllostegia
stachyoides, Ranunculus hawaiensis, R.
mauiensis, Sanicula sandwicensis,
Schiedea pubescens, Sicyos
lanceoloideus, and Solanum nelsonii),
the orangeblack Hawaiian damselfly
(directly or by desiccation of streams
and ponds), and all seven yellow-faced
bee species (and the host plants upon
which all seven yellow-faced bees
depend).
Habitat destruction and modification
by over-washing of low-lying areas by
tsunamis is a threat to coastal species,
including Solanum nelsonii, Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, and H. longiceps.
Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
Plants
We are not aware of any threats to the
39 plant species that would be
attributed to overutilization for
commercial, recreational, scientific, or
educational purposes.
Animals
Anchialine Pool Shrimp
Illegal collection is a threat to the
anchialine pool shrimp Procaris
hawaiana because of inadequate
monitoring and enforcement at the
pools where this species occurs. All
terrestrial and aquatic invertebrates
(including anchialine pool shrimp) are
protected under (1) the State of Hawaii
Revised Statutes (1993) chapter 195D–4f license; and (2) DLNR chapter 124:
Indigenous Wildlife, Endangered and
Threatened Wildlife, and Introduced
Wild Birds. Collection of plants and
animals is prohibited in the State
Natural Area Reserves (NARs) AhihiKinau (Maui) and Manuka (Hawaii
Island), but enforcement of prohibitions
is insufficient to prevent illegal
collecting at these remote sites.
Collection is not prohibited in State
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Parks or City and County property
where some anchialine pools occur, and
is not expressly prohibited at Lua O
Palahemo (Hawaii Island), and thus no
regulatory protection of these shrimp
exists at the remaining five anchialine
pools outside of Manuka NAR that are
known to contain P. hawaiana. A Native
Invertebrate Research and Collecting
permit issued by DLNR’s Division of
Forestry and Wildlife is required to
collect anchialine pool shrimp for
research or commercial purposes, and
the commercial market is supported by
legal, permitted collection. We expect
that permit holders, whether they are
collecting for scientific or commercial
purposes, adhere to the conditions of
their permit and do not pose a threat to
P. hawaiana. However, we consider
illegal collection of this anchialine pool
shrimp, P. hawaiana, to be an ongoing
threat because, despite the prohibition
on collecting within the NARs and the
permitting process for collection
elsewhere, collection can occur at any
time owing to insufficient patrolling or
other monitoring or enforcement at the
pools where P. hawaiana occurs.
Factor C. Disease or Predation
Disease
We are not aware of any current
threats to the 49 species that would be
attributable to disease. Disease may be
a potential threat to the yellow-faced
bee Hylaeus anthracinus, as pathogens
carried by nonnative bees, wasps, and
ants could be transmitted through
shared food sources (Graham 2015, in
litt.); however, we have no evidence of
this type of disease transmission at this
time.
Predation
Hawaii’s plants and animals evolved
in nearly complete isolation from
continental influence. Successful,
natural colonization of these remote
volcanic islands is infrequent, and many
organisms never succeeded in
establishing populations. As an
example, Hawaii lacks native ants and
conifers, has very few families of birds,
and has only had two native species of
land mammal, both insectivorous bats
(Loope 1998, p. 748; Ziegler 2002, pp.
244–245). In the absence of grazing or
browsing mammals, plants that became
established did not need mechanical or
chemical defenses against mammalian
herbivory such as thorns, prickles, and
toxins. Because the evolutionary
pressure to either produce or maintain
such defenses was lacking, Hawaiian
plants either lost or never developed
these adaptations (Carlquist 1980, p.
173). Likewise, native Hawaiian birds
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and insects experienced no evolutionary
pressure to develop defense
mechanisms against mammalian or
invertebrate predators that were not
historically present on the islands. The
native flora and fauna are thus
particularly vulnerable to the impacts of
introduced nonnative species, as
discussed below.
asabaliauskas on DSK3SPTVN1PROD with RULES
Introduced Ungulates
In addition to the habitat impacts
discussed above (see ‘‘Habitat
Destruction and Modification by
Introduced Ungulates’’ under Factor A.
The Present or Threatened Destruction,
Modification, or Curtailment of Their
Habitat or Range), grazing and browsing
(predation) by introduced ungulates are
a threat to the following 27 plant species
in this proposal (see Table 2, above):
Asplenium diellaciniatum (black-tailed
deer); Calamagrostis expansa (pigs),
Cyclosorus boydiae (pigs), Deparia
kaalaana (pigs), Exocarpos menziesii
(goats, mouflon, sheep), Festuca
hawaiiensis (goats, sheep), Gardenia
remyi (pigs, goats, axis deer), Huperzia
stemmermanniae (goats, axis deer,
cattle), Joinvillea ascendens ssp.
ascendens (pigs, goats, axis deer, blacktailed deer), Kadua fluviatilis (pigs,
goats), Labordia lorenciana (goats,
black-tailed deer), Microlepia strigosa
var. mauiensis (pigs), Myrsine fosbergii
(pigs, goats), Nothocestrum latifolium
(pigs, goats, axis deer, black-tailed deer,
mouflon, cattle), Ochrosia haleakalae
(goats, cattle), Phyllostegia brevidens
(pigs), P. stachyoides (pigs, goats),
Portulaca villosa (goats, axis deer,
mouflon), Pseudognaphalium
sandwicensium var. molokaiense (axis
deer), Ranunculus hawaiensis (pigs,
mouflon, cattle), R. mauiensis (pigs,
goats, axis deer, black-tailed deer,
cattle), Sanicula sandwicensis (goats),
Santalum involutum (goats), Schiedea
pubescens (axis deer, cattle), Sicyos
lanceoloideus (goats, black-tailed deer),
S. macrophyllus (mouflon, cattle), and
Solanum nelsonii (axis deer, cattle).
Feral Pigs
We have direct evidence of ungulate
damage to some of the 39 plant species,
but for many, due to their remote
locations or lack of study, ungulate
damage is presumed based on the
known presence of these introduced
ungulates in the areas where these
species occur and the results of studies
involving similar species or ecosystems
conducted in Hawaii and elsewhere
(Diong 1982, p. 160; Mueller-Dombois
and Spatz, 1975, pp. 1–29; Hess 2008,
4 pp.; Weller et al. 2011, p. 8). For
example, in a study conducted by Diong
(1982, p. 160) on Maui, feral pigs were
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observed browsing on young shoots,
leaves, and fronds of a wide variety of
plants, of which over 75 percent were
endemic species. A stomach-content
analysis in this study showed that most
of the pigs’ food source consisted of the
endemic Cibotium (hapuu, tree fern).
Pigs were observed to fell native plants
and remove the bark from standing
plants of species in the genera Cibotium,
Clermontia, Coprosma, Hedyotis
[Kadua], Psychotria, and Scaevola,
resulting in larger trees and shrubs
dying after a few months of repeated
feeding (Diong 1982, p. 144). Beach
(1997, pp. 3–4) found that feral pigs in
Texas spread disease and parasites, and
their rooting and wallowing behavior
led to spoilage of watering holes and
loss of soil through leaching and
erosion. Rooting activity by pigs also
decreased the survivability of some
plant species through disruption at root
level of mature plants and seedlings
(Beach 1997, pp. 3–4; Anderson et al.
2007, in litt.). In Hawaii, pigs dig up
forest ground cover consisting of
delicate and rare species of orchids,
ferns, mints, lobeliads, and other taxa,
including their roots, tubers, and
rhizomes (Stone and Anderson 1988, p.
137). The following plants are
particularly at risk of herbivory by feral
pigs: Calamagrostis expansa on Maui
and Hawaii Island (HBMP 2010);
Cyclosorus boydiae on Oahu (HBMP
2010); Deparia kaalaana on Maui
(HBMP 2010); Gardenia remyi on
Hawaii Island (PEPP 2011, pp. 113–114;
PEPP 2012, p. 102), west Maui (HBMP
2010), Molokai (HBMP 2010) and Kauai
(HBMP 2010); Joinvillea ascendens ssp.
ascendens on Hawaii Island (PEPP
2011, pp. 120–121; PEPP 2012, p. 113;
HBMP 2010), Kauai (PEPP 2014, p. 109;
HBMP 2010), Maui (HBMP 2010),
Molokai (HBMP 2010), and Oahu
(HBMP 2010); Kadua fluviatilis on
Kauai (HBMP 2010) and Oahu (HBMP
2010); Microlepia strigosa var.
mauiensis on Maui (Bily 2009, in litt.;
Oppenheimer 2007, in litt.); Myrsine
fosbergii on Kauai (HBMP 2010);
Nothocestrum latifolium on Maui (PEPP
2011, p. 140; HBMP 2010) and Molokai
(HBMP 2010); Phyllostegia brevidens on
Maui and Hawaii Island (PEPP 2014, p.
36); P. stachyoides on Molokai (PEPP
2014, pp. 140–141); Ranunculus
hawaiensis on Hawaii Island (HBMP
2010); and R. mauiensis on Kauai (PEPP
2011, p. 161; PEPP 2013, p. 177; PEPP
2014, p. 156; HBMP 2010), Maui (PEPP
2011, p. 144; PEPP 2013, pp. 177–178;
PEPP 2014, p. 155; HBMP 2010), and
Molokai (HBMP 2010). Feral pigs occur
in 10 of the 11 ecosystems (all except
anchialine pool) discussed here; the
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results of the studies described above
suggest that foraging by pigs can directly
damage and destroy these plants
through herbivory. Feral pigs may also
consume native host plants of the
yellow-faced bees Hylaeus anthracinus,
H. assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana.
Feral Goats
Feral goats are able to forage in
extremely rugged terrain and are
instrumental in the decline of native
vegetation in many areas of the
Hawaiian Islands (Cuddihy and Stone
1990, p. 64; Clarke and Cuddihy 1980,
p. C–20; van Riper and van Riper 1982,
pp. 34–35; Tomich 1986, pp. 153–156).
Feral goats consume a variety of plants
for food and have been observed to
browse on (but are not limited to) native
plant species in the following genera:
Argyroxiphium, Canavalia,
Chamaesyce, Erythrina, Plantago,
Schiedea, and Stenogyne (Cuddihy and
Stone 1990, p. 64; Warren 2004, p. 462;
Wood 2007, pers. comm.). A study
conducted on the island of Hawaii
demonstrated that native Acacia koa
seedlings are unable to survive due to
browsing and grazing by goats (Spatz
and Mueller-Dombois 1973, p. 874). If
goats remained in the area in high
numbers, mature trees eventually died
and with them the root systems that
supported suckers and vegetative
reproduction. When feral goats were
excluded by fences for 3 years, there
was a positive height-growth response
of A. koa suckers (Spatz and MuellerDombois 1973, p. 873). Another study at
Puuwaawaa on Hawaii Island
demonstrated that prior to management
actions in 1985, regeneration of endemic
shrubs and trees in a goat-grazed area
was almost totally lacking, contributing
to the invasion of forest understory by
exotic grasses and weeds. After the
removal of goats, A. koa and native
Metrosideros seedlings were observed
germinating by the thousands (HDLNR
2002, p. 52). Based on these studies, and
other comparisons of fenced and
unfenced areas, it is clear that goats
devastate native Hawaiian ecosystems
(Loope et al. 1988, p. 277). Because feral
goats occur in 10 of the 11 ecosystems
(all except anchialine pool) discussed in
this proposal, the results of the studies
described above indicate that goats
likely also alter these ecosystems and
directly damage or destroy native
plants. Browsing or grazing by feral
goats poses a particular threat to the
following plant species: Exocarpos
menziesii on Hawaii Island (NTBG
Herbarium Database 2014, in litt.),
Festuca hawaiiensis on Hawaii Island
(Wood 2001b, in litt.), Gardenia remyi
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asabaliauskas on DSK3SPTVN1PROD with RULES
on Kauai (PEPP 2011, p. 114; PEPP
2013, p. 107; Kishida 2011, in litt.),
Huperzia stemmermanniae on Hawaii
Island (HBMP 2010), Joinvillea
ascendens ssp. ascendens on Kauai
(PEPP 2010, p. 80), Kadua fluviatilis on
Kauai (HBMP 2010), Labordia
lorenciana on Kauai (PEPP 2011, p. 124;
PEPP 2013, p. 126), Myrsine fosbergii on
Kauai (HBMP 2010), Nothocestrum
latifolium on Maui (HBMP 2010),
Ochrosia haleakalae on Maui and
Hawaii Island (HBMP 2010),
Phyllostegia stachyoides on Molokai
(HBMP 2010), Portulaca villosa on
Hawaii Island (PEPP 2012, p. 140),
Ranunculus mauiensis on Kauai and
Maui (PEPP 2011, p. 161; PEPP 2012, p.
144; PEPP 2013, pp. 177–178; PEPP
2014, pp. 155–156; Kishida 2011, in
litt.), Sanicula sandwicensis on Maui
(PEPP 2011, p. 163), and Sicyos
lanceoloideus on Kauai (PEPP 2012, p.
154; PEPP 2013, p. 189). In addition,
browsing by feral goats may also damage
or destroy native host plants of the
yellow-faced bees Hylaeus anthracinus,
H. assimulans, H. facilis, and H. hilaris.
Axis Deer
Axis deer are known to consume a
wide range of forage items throughout
their native range and in areas where
they have been introduced (Anderson
1999, p. 3). Although they prefer to
graze on grass, axis deer have been
documented to eat over 75 species of
plants, including all plant parts
(Anderson 1999, p. 3). They exhibit a
high degree of opportunism regarding
their choice of forage, and consume
progressively less palatable plants until
no edible vegetation remains (Dinerstein
1987, in Anderson 1999, p. 5; Medeiros
2010, pers. comm.). Axis deer on Maui
follow a cycle of grazing and browsing
in open lowland grasslands during the
rainy season (November through March)
and then migrating to the lava flows of
montane mesic forest during the dry
summer months to graze and browse on
many native plant species, for example,
Abutilon menziesii (kooloaula, listed
endangered), Erythrina sandwicensis
(wiliwili), and Sida fallax (Medeiros
˜
2010, pers. comm.). During the El Nino
drought cycles from 1988 through 2001,
Maui experienced an 80 to 90 percent
decline in native shrub species caused
by axis deer browsing on and girdling
young saplings (Medeiros 2010, pers.
comm.). On Lanai, grazing by axis deer
has been reported as a major threat to
the endangered Gardenia brighamii
(nanu), and Swedberg and Walker
(1978, in Anderson 2003, pp. 124–25)
reported that the native plants
Osteomeles anthyllidifolia (uulei) and
Leptecophylla tameiameiae (pukiawe)
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comprised more than 30 percent of axis
deer rumen volume. During the driest
summer months, axis deer are observed
in coastal areas in search of food
(Medeiros 2010, pers. comm.). Because
axis deer occur in 10 of the 11
ecosystems on Molokai, Lanai, and
Maui (all except anchialine pool), the
results from the studies above, in
addition to direct observations from
field biologists, suggest that axis deer
can also alter these ecosystems and
directly damage or destroy native
plants. Browsing or grazing by axis deer
poses a threat to the following plant
species: Gardenia remyi on Molokai
(HBMP 2010), Huperzia
stemmermanniae on Maui (HBMP
2010), Joinvillea ascendens ssp.
ascendens on Maui (PEPP 2014, pp.
108–109), Nothocestrum latifolium on
Lanai (PEPP 2012, p. 129), Portulaca
villosa on Lanai (HBMP 2010),
Pseudognaphalium sandwicensium var.
molokaiense on Molokai (Wood 2005c,
in litt.; Kallstrom 2008, in litt.; MNTF
2010), Ranunculus mauiensis on Maui
(PEPP 2013, p. 178; PEPP 2014, pp.
154–155), Schiedea pubescens on
Molokai and Lanai (Wood 2004, in litt.;
Rowland 2006, in litt.; Oppenheimer
2001, in litt.), and Solanum nelsonii on
Molokai (PEPP 2012, p. 156; PEPP 2013,
pp. 190–191; PEPP 2014, p. 167). Axis
deer may also damage or destroy native
host plants of the yellow-faced bees
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, and H. longiceps.
Black-Tailed Deer
Black-tailed deer are extremely
adaptable, and in their native range
(U.S. Pacific coast) inhabit every
principal ecosystem including open
grasslands, agricultural land, shrubland,
woodland, mountain forests, semideserts, and high mountain ecosystems
(NRCS 2005, in litt.). Their home range
size varies in the continental United
States, but has been estimated to from
1 to 4 sq mi (2.5 to 10 km) and
sometimes as large as 30 sq mi (78 sq
km), with adults defending small areas
when caring for fawns (NRCS 2005, in
litt.). We do not know their home range
size on Kauai; however, the island is
only 562 sq mi (1,456 sq km) in size.
Black-tailed deer are primarily
browsers, but as they have a smaller
rumen compared to other browsers in
relation to their body size, they must
select the most nutritious plants and
parts of plants (Mule Deer Foundation
2011, in litt.). Their diet consist of a
diversity of living, wilted, dry, or
decaying vegetation, including leaves,
needles, succulent stems, fruits, nuts,
shrubs, herbaceous undergrowth,
domestic crops, and grasses (NRCS
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67837
2005, in litt.). Black-tailed deer consume
native vegetation on the island of Kauai
(van Riper and van Riper 1982, pp. 42–
43; Stone 1985, pp. 262–263; Tomich
1986, pp. 132–134; Cuddihy and Stone
1990, p. 67). In the 1990s, it was
estimated there were about 350 animals
in and near Waimea Canyon; however,
in 2013, the population was estimated
to be 1,000 to 1,200 animals in public
hunting areas (not including private
lands), and was expanding into the
southern and eastern sections of the
island (Mule Deer Working Group 2013,
in litt.). According to State records,
black-tailed deer are feeding largely on
the introduced species Psidium
cattleianum and Rubus rosifolius, as
well as the native species Alyxia stellata
(maile), Dodonaea viscosa (aalii),
Dianella sandwicensis (ukiuki),
Coprosma sp. (pilo), and Acacia koa
(Cuddihy and Stone 1990, p. 67).
Browsing by black-tailed deer is a threat
to the Kauai plant species Asplenium
diellaciniatum, Joinvillea ascendens
ssp. ascendens, Labordia lorenciana,
Nothocestrum latifolium, Ranunculus
mauiensis, and Sicyos lanceoloideus.
Mouflon and Sheep
Mouflon, feral domestic sheep, and
mouflon-sheep hybrids browse native
vegetation on Lanai and Hawaii Island.
Domestic sheep have been raised on
Kauai, Lanai, Kahoolawe, and Hawaii,
but today sheep farming only occurs on
Hawaii Island on Mauna Kea and
Hualalai (Pratt and Jacobi in Pratt et al.
2009, p. 151). Sheep browse (eating
shoots, leaves, flowers, and bark) on the
native Sophora chrysophylla (mamane),
the primary food source of the
endangered forest bird, the palila
(Loxioides bailleui) (Scowcroft and
Sakai 1983, p. 495). Feral sheep
reductions were initiated in palila
habitat; however, even after most were
removed, tree bark stripping continued
and some mamane populations did not
recover (Pratt and Jacobi in Pratt et al.
2009, p. 151). On Hawaii Island,
vegetation browsing by mouflon led to
the decline of the largest population of
the endangered Argyroxiphium kauense
(kau silversword, Mauna Loa
silversword, or ahinahina), reducing it
from a ‘‘magnificent population of
several thousand’’ (Degener et al. 1976,
pp. 173–174) to fewer than 2,000
individuals in a period of 10 years
(unpublished data in Powell 1992, p.
312). Mamane is also preferred browse
for mouflon, and according to Scowcroft
and Sakai (1983, p. 495), mouflon eat
the shoots, leaves, flowers, and bark of
this species. Mouflon are also reported
to strip bark from native koa trees and
to seek out the native plants Geranium
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asabaliauskas on DSK3SPTVN1PROD with RULES
cuneatum (hinahina) and Silene
hawaiiensis, and Lanai occurrences of
Gardenia brighamii (Benitez et al. 2008,
p. 57; Mehrhoff 1993, p. 11). While
mouflon were introduced to Lanai and
Hawaii Island as game mammals, a
private game ranch on Maui has added
mouflon to its stock, and it is likely that
over time some individuals may escape
(Hess 2010, pers. comm.; Kessler 2010,
pers. comm.). Browsing and grazing by
mouflon, feral domestic sheep, and
mouflon-sheep hybrids poses a threat to
the plant species Exocarpos menziesii
on Hawaii Island (Keitt and Island
Conservation 2008, pp. 90, 92; NPS
2013, pp. i, 124); Festuca hawaiiensis on
Hawaii Island (Oppenheimer 2001, in
litt.; HBMP 2007, in litt.); Nothocestrum
latifolium on Lanai (PEPP 2012, p. 129);
Portulaca villosa on Lanai (HBMP
2010); Ranunculus hawaiensis on
Hawaii Island (HBMP 2010); and Sicyos
macrophyllus on Hawaii Island (HBMP
2010). Because feral sheep and mouflon
occur in all of the described ecosystems
except for the anchialine pool
ecosystem, the data from studies above
suggest that in addition to consuming
the host plants of the yellow-faced bees
Hylaeus anthracinus and H. assimulans
on Lanai, herbivory by feral sheep and
mouflon also may consume host plants
of the other species on Lanai: H. facilis,
H. hilaris, and H. longiceps.
Feral Cattle
Grazing by cattle is considered one of
the most important factors in the
destruction of Hawaiian forests
(Baldwin and Fagerlund 1943, pp. 118–
122). Feral cattle are currently found
only on the islands of Molokai, Maui,
and Hawaii (Tomich 1986, pp. 140–144;
de Sa et al. 2013, 29 pp.). Cattle
consume tree seedlings and browse
saplings (Cuddihy 1984, p. 16). In
Hawaii Volcanoes National Park
(Hawaii Island), Cuddihy reported that
there were twice as many native plant
species as nonnatives in areas that had
been fenced to exclude cattle (Cuddihy
1984, pp. 16, 34). Loss of the native
sandalwood forest on Lanai is attributed
to cattle (Skottsberg 1953 in Cuddihy
1984, p. 16). Browsing and grazing by
feral cattle poses a threat to the
following plant species: Huperzia
stemmermanniae on Maui and Hawaii
Island (Medeiros et al. 1996b, p. 96);
Nothocestrum latifolium on Molokai
and Maui (HBMP 2010); Ochrosia
haleakalae on Maui (HBMP 2010);
Ranunculus hawaiensis on Hawaii
Island (HBMP 2010); R. mauiensis on
Maui and Hawaii Island (PEPP 2012, p.
144; PEPP 2013, p. 178; PEPP 2014, pp.
154–155; HBMP 2010); Schiedea
pubescens on Maui (Wood 2005d, in
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litt.; HBMP 2010); Sicyos macrophyllus
on Hawaii Island (PEPP 2010, p. 111;
HBMP 2010); and Solanum nelsonii on
Molokai (Wood 1999, in litt.; HBMP
2010). Because feral cattle occur in 6 of
the 11 ecosystems (lowland dry,
lowland mesic, lowland wet, montane
wet, montane mesic, and subalpine) in
which these species occur on Molokai,
Maui, and Hawaii Island, the results
from the studies cited above, in addition
to direct observations from field
biologists, indicate that grazing by feral
cattle can directly damage or destroy
these plants.
Blackbuck
The blackbuck antelope (Antelope
cervicapra) is an endangered species
from India brought to a private game
reserve on Molokai about 15 years ago
from an Indian zoo (Kessler 2010, pers.
comm.). According to Kessler (2010,
pers. comm.), a few individuals escaped
captivity and established a wild
population of unknown size on the low,
dry plains of western Molokai.
Blackbuck primarily use grassland
habitat for grazing. In India, foraging
consumption and nutrient digestibility
are high in the moist winter months and
low in the dry summer months (Jhala
1997, pp. 1348, 1351). Although most
plant species are grazed intensely when
they are green, some are grazed only
after they are dry (Jhala 1997, pp. 1348,
1351). Because the foraging dynamics of
blackbuck antelope in Hawaii and
possible habitat effects are unknown at
this time, we do not currently consider
this ungulate a threat to the four native
plant species known from dry areas on
Molokai: Gardenia remyi, Nothocestrum
latifolium, Portulaca villosa, and
Pseudognaphalium sandwicensium var.
molokaiense, or to the yellow-faced bees
Hylaeus anthracinus, H. facilis, H.
hilaris, and H. longiceps (which rely on
host plants that ungulates consume).
Other Introduced Vertebrates
Rats
Three species of introduced rats occur
in the Hawaiian Islands. Studies of
Polynesian rat (Rattus exulans) DNA
suggest they first appeared in the
islands along with emigrants from the
Marquesas Islands (French Polynesia) in
about 400 A.D., with a second
introduction around 1100 A.D. (Ziegler
2002, p. 315). The black rat (R. rattus)
and the Norway rat (R. norvegicus)
arrived in the islands more recently, as
stowaways on ships sometime in the
late 19th century (Atkinson and
Atkinson 2000, p. 25). The Polynesian
rat and the black rat are primarily found
in rural and remote areas of Hawaii, in
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dry to wet habitats, while the Norway
rat is typically found in urban areas or
agricultural fields (Tomich 1986, p. 41).
The black rat is widely distributed
throughout the main Hawaiian Islands
and can be found in a range of
ecosystems and as high as 9,000 ft
(2,700 m), but it is most common at lowto mid-elevations (Tomich 1986, pp. 38–
40). Sugihara (1997, p. 194) found both
the black and Polynesian rats up to
7,000 ft (2,000 m) on Maui, but found
the Norway rat only at lower elevations.
Rats are omnivorous and eat almost any
type of food (Nelson 2012, in litt.). Rats
occur in 7 of the 11 ecosystems (coastal,
lowland mesic, lowland wet, montane
wet, montane mesic, montane dry, and
wet cliff), and predation or herbivory by
rats is a threat to 19 plants
(Calamagrostis expansa (Maui and
Hawaii Island; HBMP 2010), Cyanea
kauaulaensis (Maui; PEPP 2012, pp. 71–
72; PEPP 2014, p. 73), Dryopteris glabra
var. pusilla (Kauai; Wood 2015, in litt.);
Gardenia remyi (Kauai, Molokai, Maui,
and Hawaii Island; Wood 2004, in litt.;
HBMP 2010); Joinvillea ascendens ssp.
ascendens (Kauai, Oahu, Molokai, Maui,
and Hawaii Island; PEPP 2014, p. 109),
Kadua haupuensis (Kauai; Lorence et al.
2010, p. 140), Labordia lorenciana
(Kauai; Wood et al. 2007, p. 198),
Ochrosia haleakalae (Maui;
Oppenheimer 2015, in litt.), Phyllostegia
helleri (Kauai; HBMP 2010), P.
stachyoides (Molokai, Maui, and Hawaii
Island; PEPP 2012, p. 133; PEPP 2013,
pp. 158–159; PEPP 2014, pp. 140–142),
Pritchardia bakeri (Oahu; Hodel 2012,
pp. 42, 73), Ranunculus mauiensis
(Kauai, Oahu, Molokai, Maui, and
Hawaii Island; HBMP 2010), Sanicula
sandwicensis (Maui and Hawaii Island;
PEPP 2012, p. 148), Santalum
involutum (Kauai; Harbaugh et al. 2010,
pp. 835–836), Schiedea diffusa ssp.
diffusa (Molokai, Maui; HBMP 2010), S.
pubescens (Molokai, Lanai, and Maui;
Wood 2005d, in litt.; HBMP 2010),
Sicyos macrophyllus (Maui and Hawaii
Island; Pratt 2008, in litt.), Solanum
nelsonii (NWHI, Niihau, Molokai, Maui,
and Hawaii Island; PEPP 2012, p. 156;
PEPP 2014, p. 167), and Wikstroemia
skottsbergiana (Kauai; Mitchell et al.
2005, in litt.)) and to the band-rumped
storm-petrel (Lehua, Kauai, Maui,
Kahoolawe, Lanai, and Hawaii Island;
Pyle and Pyle 2009, in litt.).
Rat Impacts on Plants: Rats affect
native plants by eating fleshy fruits,
seeds, flowers, stems, leaves, roots, and
other plant parts (Atkinson and
Atkinson 2000, p. 23), and by stripping
bark and cutting small branches (twig
cutting) in search of moisture and
nutrients, with detrimental impacts to
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plants’ vigor and regeneration (Abe and
Umeno 2011, pp. 27–39; Nelson 2012,
pp. 1–4, 8–9). Studies in New Zealand
have demonstrated that differential
regeneration as a consequence of rat
predation alters species composition of
forested areas (Cuddihy and Stone 1990,
pp. 68–69). Rats have caused declines or
even the total elimination of island
plant species (Campbell and Atkinson
1999 in Atkinson and Atkinson 2000, p.
24). In the Hawaiian Islands, rats may
consume as much at 90 percent of the
seeds produced by some native plants,
and in some cases prevent regeneration
of forest species completely (Cuddihy
and Stone 1990, pp. 68–69). Hawaiian
plants with fleshy fruit, such as Cyanea
and Pritchardia, are particularly
susceptible to rat predation (Cuddihy
and Stone 1990, pp. 67–69). Predation
of seeds by rats poses a serious and
ongoing threat to all the Hawaiian
Pritchardia palms, including P. bakeri,
because rats are able to consume every
seed in a fruiting stalk, preventing
successful reproduction (Hodel 2012,
pp. 42, 73). Fossil pollen records
indicate that Pritchardia palms were
once among the dominant species of
coastal, lowland, and interior forests of
Hawaii (Burney et al. 2001, pp. 630–
631; Chapin et al. 2007, p. 21); today,
complete coverage by all age classes of
Pritchardia occurs only on small islets
currently unoccupied by rats (Athens
2009, p. 1498). Because rats occur in
seven of the ecosystems in which these
species occur, the results from these
studies, in addition to direct
observations by field biologists, suggest
that predation by rats can directly
damage or destroy native plants.
Rat Impacts on the Band-Rumped
Storm-Petrel: Introduced predators are
the most serious threat facing the bandrumped storm-petrel. Rats occur on all
the main Hawaiian Islands, and
populations are also high on Lehua;
however, attempts to control rats on
Lehua are ongoing (Parkes and Fisher
2011, 48 pp.). Ground-, crevice-, and
burrow-nesting seabirds, as well as their
eggs and young, are highly susceptible
to predation by rats; storm-petrels are
the most susceptible of seabirds to rat
predation and have experienced
population-level impacts and
extirpation as a result (Simons 1984, p.
1073; Jones et al. 2008, pp. 20–21).
Evidence from the islands of Hawaii and
Maui show that the Hawaiian petrel, a
much larger seabird that nests in some
of the same areas as the band-rumped
storm-petrel, suffers huge losses to
introduced predators (Johnston 1992, in
litt.; Hodges and Nagata 2001, pp. 308–
310; Hu et al. 2001, p. 234). The effects
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of introduced predators on the breeding
success of the band-rumped stormpetrel are probably similar to the
documented effects on the breeding
success of Hawaiian petrels because
these birds are similarly vulnerable.
Population modeling showed that
consistent predation of Hawaiian
petrels, where reproductive success was
reduced to 35 percent and adult survival
was 80 percent, could drive a
population to extinction in 20 to 30
years (Simons 1984, pp. 1071–1073). Rat
bones were collected from a bandrumped storm-petrel nest on a sheer
cliff on Kauai, and two live rats were
observed moving along small rock
ledges in the same area (Wood et al.
2002, p. 8), demonstrating that even
remote and otherwise inaccessible nest
sites are not safe from these predators.
Because rats are present in all three
ecosystems in which the band-rumped
storm-petrel occurs (coastal, dry cliff,
and wet cliff), predation by rats likely
results in decreases in the numbers and
populations of the band-rumped stormpetrel. We do not anticipate a reduction
of this threat in the near future.
Barn Owl Impacts on the Band-Rumped
Storm-Petrel
Two species of owls, the native pueo
(Asio flammeus sandwichensis) and the
introduced barn owl (Tyto alba), are
known to prey on native Hawaiian
birds. For example, between 1996 and
1998, 10 percent of nest failures of the
puaiohi (small Kauai thrush, Myadestes
palmeri), an endangered forest bird, on
Kauai were attributed to owls
(Snetsinger et al. 1994, p. 47; Snetsinger
et al. 2005, pp. 72, 79). The bandrumped storm petrel only comes to land
after dark, and likely avoids predation
by the pueo, which hunts in daylight
(Hawaii DOFAW 2005). The nonnative
barn owl, however, is a nocturnal
hunter and may prey on the stormpetrel. Barn owls were introduced to
Kauai, Oahu, Molokai, and Hawaii
Island between 1958 and 1963, to
control rats and mice in sugar cane
plantations, and now they occur on all
of the main islands (USFWS 2013, p. 9).
Barn owls are well-known predators of
storm-petrels and other seabirds on
islands (LeCorre and Jouventin 1997, p.
215; Velarde et al. 2007, p. 286; Guerra
et al. 2014, p. 182; Ringler et al. 2015,
p. 79). Direct impacts of barn owls on
band-rumped storm-petrels in Hawaii
are not well documented, but evidence
and numerous anecdotal reports exist of
barn owls preying on seabirds in the
main Hawaiian islands, including the
threatened Newell’s shearwater and
endangered Hawaiian petrel, and
including on Kauai and Lehua, where
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band-rumped storm-petrels are known
to nest (summarized in USFWS 2013,
pp. 11–12). Because barn owls occur
throughout the range of the bandrumped storm-petrel in Hawaii, they are
likely to be predators of these seabirds.
Cat Impacts on the Band-Rumped
Storm-Petrel
Cats (Felis catus) were introduced to
Hawaii in the early 1800s, and are
present on all the main Hawaiian
Islands (Tomich 1986, p. 101). Cats are
notorious for their predation on birds
(Tomich 1986, p. 102). Native
mammalian carnivores are absent from
oceanic islands because of their low
dispersal ability, but once introduced,
are significant predators on seabird
colonies and terrestrial birds that have
no innate defenses against predation by
these animals (Scott et al. 1986, p. 363;
Ainley et al. 1997, p. 24; Ziegler 2002,
p. 243; Hess and Banko 2006, in litt.;
Nogales et al. 2013, p. 804). Cats may
have contributed to the extinction of the
Hawaiian rail (Porzana sandwichensis)
(Stone 1985 in Stone and Scott 1985, p.
266). Although cats are more common at
lower elevations, there are populations
in areas completely isolated from
human presence, including montane
forests and alpine areas of Maui and
Hawaii Island (Lindsey et al. in Pratt et
al. 2009, p. 277; Scott et al. 1986, p.
363). Examination of the stomach
contents of feral cats at Hakalau Forest
NWR (Hawaii Island) found native and
introduced birds to be the most common
prey item (Banko et al. 2004, p. 162).
Cats are believed to prey on roosting or
incubating adult band-rumped stormpetrels and young, as evidenced by
carcasses found in Hawaii Volcanoes
National Park depredated by cats (Hu,
pers. comm. in Slotterback 2002, in litt.;
Hess et al. 2008, pp. 11, 14). Predation
by cats is well known for the
endangered Hawaiian petrel, which has
some accessible and well-studied
nesting areas; this species shares lifehistory and evolutionary traits with the
band-rumped storm-petrel that make
both vulnerable to nonnative
mammalian predators. We expect the
band-rumped storm-petrel to experience
impacts of cat predation similar to those
documented in the Hawaiian petrel. On
Mauna Loa (Hawaii Island), feral cats
were major predators of Hawaiian
petrels (Hu et al. 2001, p. 234), and on
Haleakala (Maui), almost half of the
known mortalities of Hawaiian petrels
between 1964 and 1996 were attributed
to cats (Natividad Hodges and Nagata
2001, p. 312; Hu et al. 2001, p. 234).
Population modeling of the Hawaiian
petrel indicated that the petrel
population would be unable to
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withstand any level of predation for
long, and even with seemingly low
levels of predation, the petrel
population would be reduced by half in
fewer than 30 years (Simons 1984, p.
1073). The band-rumped storm petrel is
smaller in size than the petrel, but also
nests in burrows and rock-crevices,
lacks co-evolved predator avoidance
behavior, and has a lengthy incubation
and fledgling period, making this
species highly vulnerable to predation
by introduced mammals. Because feral
cats occur in all four ecosystems in
which the band-rumped storm petrel
occurs, they are likely to be significant
predators of these birds.
Mongoose Impacts on the Band-Rumped
Storm-Petrel
The small Indian mongoose
(Herpestes auropunctatus) was
introduced to Hawaii in 1883, to control
rodents in sugar cane plantations
(Tomich 1986, pp. 95–96). This species
quickly became widespread on Oahu,
Molokai, Maui, and Hawaii Island, from
sea level to elevations as high as 7,000
ft (2,130 m) (Tomich 1986, pp. 93–94).
Mongooses have been sighted, and two
captured, on Kauai, but it is still
uncertain if the species is established
there or how large populations might be
(Kauai Invasive Species Committee
2013, in litt.; The Garden Island 2012,
in litt.; Hess et al. in Pratt et al. 2009,
p. 429). Mongooses are omnivorous, are
known to prey on Hawaiian birds and
their eggs, and are considered a likely
factor in the decline of the endangered
Hawaiian goose (nene, Branta
sandvicensis) (Tomich 1986, p. 97).
They are known or suspected predators
on other Hawaiian birds, including the
Hawaiian crow (alala, Corvus
hawaiiensis), Hawaiian duck (koloa,
Anas wyvilliana), Hawaiian coot (alae
keokeo, Fulica alai), Hawaiian stilt (aeo,
Himantopus mexicanus knudseni),
Hawaiian gallinule (ula, Gallinula
chloropus sandvicensis), Hawaiian
petrel, and Newell’s shearwater. Bird
extinctions in other areas are attributed
to mongooses, such as the loss of the
barred-wing rail (Nesoclopeus
poecilopterus) in Fiji, and the Jamaica
petrel (Pterodroma caribbaea) (Hays and
Conant 2007, p. 6). Birds extirpated
from islands occupied by mongooses
retain their populations on islands
known to be mongoose-free (Hays and
Conant 2007, p. 7). In Hawaii,
mongooses occur in habitat types where
they are not found within their natural
range, and they have no predators and
few communicable diseases or parasites.
Because mongooses occur in all four
ecosystems in which the band-rumped
storm petrel occurs, they are likely to be
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significant predators of the bandrumped storm-petrel.
Nonnative Fish Impacts on the
Orangeblack Hawaiian Damselfly
Predation by nonnative fishes is a
threat to the orangeblack Hawaiian
damselfly. Similar to the aquatic insects,
Hawaii has a low diversity of freshwater
fishes, with only five native species in
two families (gobies (Gobiidae) and
sleepers (Eleotridae)) that occur on all
the main islands (Devick 1991, p. 196).
Information on these five species
indicates that the Hawaiian damselflies
probably experienced limited natural
predation pressure from these native
fishes (Kido 1997, p. 493; Englund 1999,
p. 236). Conversely, fish predation has
been an important factor in the
evolution of behavior in damselfly
naiads in continental systems (Johnson
1991, p. 13). Some species of
damselflies, including the native
Hawaiian species, are not adapted to
coexist with some fish species, and are
found only in bodies of water without
fish (Henrikson 1988, pp. 179–180;
McPeek 1990a, pp. 92–93). The naiads
of these species tend to occupy more
exposed positions and engage in
conspicuous foraging behavior that
makes them susceptible to predation by
fishes (Macan 1977, p. 47; McPeek
1990b, p. 1722). The introduction of
nonnative fishes has been implicated in
the extirpation of a species related to the
orangeblack Hawaiian damselfly, the
endangered Pacific Hawaiian damselfly
(Megalagrion pacificum), from Oahu,
Kauai, and Lanai, and from many
streams on the remaining islands where
it occurs (Moore and Gagne 1982, pp.
1–4). More than 70 species of fish have
been introduced into Hawaiian
freshwater habitats (Devick 1991, p. 189;
Englund and Eldredge in Staples and
Cowie 2001, p. 32; Englund 2004, in
litt., p.27). The impact of fish
introductions prior to 1900 cannot be
assessed because this predates the
initial collection of damselflies in
Hawaii (Perkins 1913, p. clxxvi). In
1905, two species, the mosquito fish
(Gambusia affinis) and the sailfin molly
(Poecilia latipinna), were introduced for
biological control of mosquitoes (Van
Dine 1907, pp. 6–9). In 1922, three
additional species were established for
mosquito control, the green swordtail
(Xiphophorus helleri), the moonfish
(Xiphophorus maculatus), and the
guppy (Poecilia reticulata). By 1935, the
orangeblack Hawaiian damselfly was
found only in waters without
introduced fishes (Williams 1936, p.
289; Zimmerman 1948b, p. 341;
Polhemus 1993b, p. 591; Englund 1998,
p. 235). Beginning about 1980, a large
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number of new fish introductions began
in Hawaii, originating primarily from
the aquarium fish trade (Devick 1991, p.
189). This recent wave of fish
introductions on Oahu corresponded
with the drastic decline and range
reduction of other Hawaiian damselfly
species: The endangered oceanic
Hawaiian damselfly (Megalagrion
oceanicum), the endangered crimson
Hawaiian damselfly (M. leptodemas),
and the endangered blackline Hawaiian
damselfly (M. nigrohamatum
nigrolineatum). Currently, these
damselflies are found only in drainages
or higher parts of stream systems where
nonnative fish are not yet established
(Englund and Polhemus 1994, pp. 8–9;
Englund 2004, in litt., p. 27). In
summary, Hawaiian damselflies evolved
with few, if any, predatory fishes and
the lack of defensive behavior makes the
orangeblack Hawaiian damselfly
particularly vulnerable to, and are
threatened by, predation by nonnative
fish.
Nonnative Fish Impacts on the
Anchialine Pool Shrimp
In Hawaii, the introduction of
nonnative fishes into anchialine pools
and the ensuing predation by nonnative
fishes is considered the greatest threat to
native shrimp within anchialine pool
systems (Bailey-Brock and Brock 1993,
p. 354). These impacts are discussed
further under Factor E. Other Natural or
Manmade Factors Affecting Their
Continued Existence, below.
Bullfrog Impacts on the Orangeblack
Hawaiian Damselfly
Native to the eastern United States
and the Great Plains region, the bullfrog
(Rana catesbeiana, Lithobates
catesbeiana), was first introduced to
Hawaii in 1899, to help control insects,
and has become established on all the
main Hawaiian Islands (Bryan 1931, pp.
62–63; Bury and Whelan 1985, p. 1;
Lever 2003, p. 203). The bullfrog is
flexible in both habitat and food
requirements (McKeown 1996, pp. 24–
27; Bury and Whelan 1984, pp. 3–7;
Lever 2003, pp. 203–204), and can
utilize any water source within a
temperature range of 60 to 75 degrees
Fahrenheit (°F) (16 to 24 degrees Celsius
(°C)) (DesertUSA 2008). Englund et al.
(2007, pp. 215, 219) found a strong
correlation between the presence of
bullfrogs and the absence of Hawaiian
damselflies in their study of streams on
all the main Hawaiian Islands. Because
bullfrogs are omnivorous feeders and
occur in the same habitat as the
orangeblack Hawaiian damselfly, we
consider predation by bullfrogs a threat
to the orangeblack Hawaiian damselfly.
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Introduced Invertebrates
Slugs
Herbivory by nonnative slugs is a
threat to 10 of the 39 plant species
(Cyanea kauaulaensis (Maui); Deparia
kaalaana (Maui), Dryopteris glabra var.
pusilla (Kauai), Hypolepis hawaiiensis
var. mauiensis (Maui), Ochrosia
haleakalae (Maui, Hawaii Island),
Phyllostegia brevidens (Maui), P.
stachyoides (Molokai, Maui),
Ranunculus mauiensis (Kauai, Maui),
Schiedea diffusa ssp. diffusa (Maui),
and S. pubescens (Molokai, Maui))
through mechanical damage,
destruction of plant parts, and mortality
(see Table 2) (Joe 2006, p. 10; HBMP
2010; PEPP 2011, pp. 149, 170; PEPP
2012, pp. 71–72, 117–118, 133, 144–
145, 153; PEPP 2013, pp. 54, 67, 91,
125–126, 158–159, 177–178, 185;
Oppenheimer and Bustamente 2014, p.
106; PEPP 2014, pp. 73, 112–114, 136,
141–142, 154–156, 159, 162–163). Slugs
are known to damage individuals of
Cyanea and Cyrtandra species in the
wild (Wood 2001, in litt.; Sailer and
Kier 2002, in litt.; PEPP 2007, p. 38;
PEPP 2008, pp. 23, 29, 52–53, 57).
Information in the U.S. Army’s 2005
‘‘Status Report for the Makua
Implementation Plan’’ indicates that
herbivory by slugs can be a threat to all
species of Cyanea, and can result in up
to 80 percent seedling mortality (U.S.
Army Garrison 2005, p. 3–51). Slug
damage has also been reported on other
Hawaiian plants including
Argyroxiphium grayanum (greensword),
Alsinidendron sp., Hibiscus sp.,
Schiedea kaalae (maolioli), Solanum
sandwicense (popolo aiakeakua), and
Urera sp. (Gagne 1983, pp. 190–191;
Sailer 2006, pers. comm. in Joe 2006,
pp. 28–34). Joe and Daehler (2008, p.
252) found that native Hawaiian plants
are more vulnerable to slug damage than
nonnative plants. In particular, they
found that individuals of the
endangered plants Cyanea superba and
Schiedea obovata had 50 percent higher
mortality when exposed to slugs as
compared to individuals that were
within exclosures without slugs.
Because slugs are reported in five
ecosystems (lowland mesic, lowland
wet, montane wet, montane mesic, and
wet cliff) on all the main Hawaiian
Islands, the data from the studies cited
above, in addition to direct observations
by field biologists, indicate that slugs
can directly damage or destroy native
plants.
Black Twig Borers
The black twig borer (Xylosandrus
compactus) is known to infest a wide
variety of common plant taxa, including
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rare native plant species (Davis 1970, p.
39; Extension Entomology and US–
CTAHR Integrated Pest Management
Program 2006, p. 1). This insect pest
burrows into branches, introduces a
pathogenic fungus as food for its larvae,
and lays its eggs (Davis 1970, p. 39).
Twigs, branches, and entire plants can
be damaged or killed from an infestation
(Extension Entomology and UH–CTAHR
Integrated Pest Management Program
2006, in litt.). On the Hawaiian Islands,
the black twig borer has many hosts,
disperses easily, and is probably present
at most elevations up to 2,500 ft (762 m)
(Howarth 1985, pp. 152–153). The black
twig borer is reported as a threat to
Labordia lorenciana and Nothocestrum
latifolium (Ching-Harbin 2015, in litt.;
Kishida 2015, in litt.).
Backswimmers
Backswimmers are aquatic true bugs
(Heteroptera) in the family
Notonectidae, so called because they
swim upside down. Backswimmers are
voracious predators and frequently feed
on prey much larger than themselves,
such as tadpoles, small fish, and other
aquatic invertebrates including
damselfly naiads (Borror et al. 1989, p.
296; Zalom 1978, p. 617).
Backswimmers (several species) were
introduced in recent times. Buenoa
pallipes (NCN) has been recorded from
Hawaii Island, Oahu, Maui, and Kauai
(Zimmerman 1948, pp. 232–233; Larsen
1996, p. 40). This species is found in
streams and can be abundant in lowland
ponds and reservoirs. It feeds on any
suitably sized insect, including
damselfly naiads (Zalom 1978, p. 617).
Two additional species of
backswimmers have become established
in Hawaii, Anisops kuroiwae (NCN) on
Maui and Lanai, and Notonecta indica
(NCN) on Hawaii Island, Oahu, and
Maui (Larsen 1996, pp. 39–40).
Predation by backswimmers is a threat
to the orangeblack Hawaiian damselfly
(Haines 2015, in litt.).
Ants
At least 47 species of ants are known
to be established in the Hawaiian
Islands (Hawaii Ants 2008, 11 pp.). No
native ant species occur in Hawaii, and
the native yellow-faced bee species in
Hawaii evolved in the absence of
predation pressure from ants. Ants are
known to prey upon Hawaiian yellowfaced bee (Hylaeus) species, with
observations of drastic reductions in
yellow-faced bee populations in antinfested areas (Medeiros et al. 1986, pp.
45–46; Reimer 1994, p. 17; Stone and
Loope 1987, p. 251; Cole et al. 1992, pp.
1313, 1317, 1320). The presence of ants
in nearly all of the low-elevation habitat
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sites currently and historically occupied
by yellow-faced bee species may
preclude these species’ recovery in
some of these areas (Reimer 1994, pp.
17–18; Daly and Magnacca 2003, pp. 9–
10). Although the primary impact of
ants on Hawaii’s native invertebrate
fauna is via predation, they also
compete for nectar (Reimer 1994, p. 17;
Howarth 1985, p. 155; Hopper et al.
1996, p. 9; Holway et al. 2002, pp. 188,
209; Daly and Magnacca 2003, p. 9;
Lach 2008, p. 155) and nest sites
(Krushelnycky et al. 2005, pp. 6–7).
Some ant species may affect yellowfaced bee species indirectly as well, by
consuming seeds of native host plants,
thereby reducing the plants’ recruitment
and fecundity (Bond and Slingsby 1984,
p. 1031). The threat of ant predation on
the yellow-faced bees is amplified by
the fact that most ant species have
winged reproductive adults and can
quickly expand their range by
establishing new colonies in suitable
habitat (Staples and Cowie 2001, p. 55).
In addition, these attributes allow some
ants to destroy otherwise geographically
isolated populations of native
arthropods (Nafus 1993, pp. 19, 22–23).
Several studies suggest a serious
ecosystem-level effect of invasive ants
on plant pollination (Krushelnycky et
al. 2005, p. 9; Lach 2008, p. 155). Where
ranges overlap, ants compete with
native pollinators such as yellow-faced
bees and preclude them from
pollinating native plants (Howarth 1985,
p. 157), potentially leading to a decrease
in availability of the bees’ native plant
food sources. Lach (2008, p. 155) found
that yellow-faced bees that regularly
consume pollen from flowers of
Metrosideros polymorpha (ohia) were
entirely absent from trees with flowers
visited by the ant Pheidole
megacephala.
The four most aggressive ant species
in Hawaii are the big-headed ant
(Pheidole megacephala), the yellow
crazy ant (Anoplolepis gracilipes), the
tropical fire ant (Solenopsis geminata),
and S. papuana (NCN). The big-headed
ant is native to central Africa and was
first reported in Hawaii in 1879
(Krushelnycky et al. 2005, p. 24). This
species occurs from coastal to mesic
habitat up to 4,000 ft (1,220 m) in
elevation. With few exceptions, native
insects have been eliminated in habitats
where the big-headed ant is present
(Perkins 1913, p. xxxix; Gagne 1979, p.
81; Gillespie and Reimer 1993, p. 22).
Native habitat of the yellow crazy ant is
not known, but it is speculated the
species originated in West Africa
(MacGown 2015, in litt.). It occurs in
low to mid elevations (less than 2,000
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ft (600 m)) in rocky areas of moderate
annual rainfall (less than 100 in (250
cm)) (Reimer et al. 1990, p. 42).
Although surveys have not been
conducted to ascertain this species’
presence in each of the known habitats
occupied by the seven yellow-faced
bees, we know that the yellow crazy ant
occurs adjacent to some of the identified
populations sites based upon
observations of their expanding range
and their preference for coastal and dry
forest habitat (as indicated where the
species is most commonly collected)
(Antweb 2015, in litt.; Magnacca and
King 2013, pp. 13–14). Direct
observations indicate that Hawaiian
arthropods are susceptible to predation
by this ant species. Gillespie and Reimer
(1993, pp. 21, 26) and Hardy (1979, pp.
37–38) documented the complete
elimination of native spiders from mesic
and dry forests after they were invaded
by the big-headed ant and the yellow
crazy ant. Lester and Tavite (2004, p.
291) found that the yellow crazy ant in
the atolls of Tokelau (Central Polynesia)
form very high densities in a relatively
short period of time with locally serious
consequences for invertebrate diversity.
Densities of 3,600 individuals collected
in pitfall traps within a 24-hour period
were observed, as well as predation on
invertebrates ranging from crabs to other
ant species. Results from these and
other studies (Reimer et al. 1990, p. 47)
indicate that yellow crazy ants have the
potential as predators to profoundly
affect endemic insect fauna in areas they
occupy. We believe that the yellow
crazy ant is a threat to populations of
the Hawaiian yellow-faced bees in areas
within their range.
Solenopsis papuana, native to the
Pacific region but not to Hawaii, is the
only abundant, aggressive ant that has
invaded intact mesic and wet forest, as
well as coastal and lowland dry
ecosystems. First detected in 1967, this
species occurs from sea level to over
3,600 ft (1,100 m) on all of the main
Hawaiian Islands, and is still expanding
its range (Reimer et al. 1990, p. 42;
Reimer 1993, p. 14). Studies have been
conducted that suggest a negative effect
of this ant species on indigenous
invertebrates (Gillespie and Reimer
1993, p. 21). Although surveys have not
been conducted to ascertain the
presence of S. papuana in each of the
known ecosystems occupied by the
seven yellow-faced bees, because of the
expanding range of this introduced ant
species, and its widespread occurrence
in coastal to wet habitats, it is a possible
threat to all known populations of the
seven yellow-faced bees.
Solenopsis geminata is also
considered a significant threat to native
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invertebrates in Hawaii (Wong and
Wong 1988, p. 171). Found in drier
areas of all the main Hawaiian Islands,
it displaced Pheidole megacephala as
the dominant ant in some localities
more than 20 years ago (Wong and
Wong 1988, p. 175). Known to be a
voracious predator, this ant species was
documented to significantly increase
native fruit fly mortality in field studies
in Hawaii (Wong and Wong 1988, p.
175). Solenopsis geminata is included
among the eight species ranked as
having the highest potential risk to New
Zealand species in a detailed pest risk
assessment for the country (GISD 2011,
in litt.), and is included as one of the
five ant species listed among the ‘‘100
of the World’s Worst Invaders’’
(Manaaki Landcare Research 2015, in
litt.). In addition to predation, S.
geminata workers tend honeydewproducing members of the Homoptera
suborder, especially mealybugs, which
can affect plants directly and indirectly
through the spread of disease (Manaaki
Landcare Research 2015, in litt.).
Although surveys have not been
conducted to ascertain the presence of
S. geminata in each of the known seven
yellow-faced bees’ habitat sites, because
of its expanding range and widespread
presence, S. geminata is a threat to all
known populations of the seven yellowfaced bees.
Although we have no direct
information that correlates the decrease
in populations of the seven yellow-faced
bees in this final rule due to the
establishment of nonnative ants,
predation of and competition with other
yellow-faced bee species by ants has
been documented, resulting in clear
reductions in or absence of populations
(Magnacca and King 2013, p. 24). We
expect similar predation impacts to the
seven yellow-faced bees to continue as
a result of the widespread presence of
ants throughout the Hawaiian Islands,
their highly efficient and non-specific
predatory behavior, and their ability to
quickly disperse and establish new
colonies. Therefore, we conclude that
predation by nonnative ants represents
a serious threat to the continued
existence of the seven yellow-faced
bees, now and into the future.
Wasps
Predation by the western yellow
jacket wasp (Vespula pensylvanica) is a
serious and ongoing threat to the seven
yellow-faced bees (Gambino et al. 1987,
p. 170; Wilson et al. 2009, pp. 1–5). The
western yellow jacket is a social wasp
species native to mainland North
America. It was first reported on Oahu
in the 1930s (Sherley 2000, p. 121), and
an aggressive race became established in
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1977 (Gambino et al. 1987, p. 170). In
temperate climates, the western yellow
jacket wasp has an annual life cycle, but
in Hawaii’s tropical climate, colonies of
this species persist year round, allowing
growth of large populations (Gambino et
al. 1987, p. 170) and thus a greater
impact on prey populations. Most
colonies occur between 2,000 and 3,500
ft (600 and 1050 m) in elevation
(Gambino et al. 1990, p. 1088), although
they can also occur at sea level. The
western yellow jacket wasp is known to
be an aggressive, generalist predator and
has been documented preying upon
Hawaiian yellow-faced bee species
(Gambino et al. 1987, p. 170; Wilson et
al. 2009, p. 2). It has been suggested that
the western yellow jacket wasp may
compete for nectar with native
Hawaiian invertebrates, but we have no
information to suggest this represents a
threat to the seven yellow-faced bees.
Predation by the western yellow jacket
wasp is a significant threat to the seven
yellow-faced bee species because of the
wasps’ presence in habitat combined
with the small number of occurrences
and small population sizes of the
Hawaiian yellow-faced bees.
Summary of Factor C
We are unaware of any information
that indicates that disease is a threat to
the 39 plant species. We are also
unaware of any information that
indicates that disease is a threat to the
band-rumped storm-petrel, the
orangeblack Hawaiian damselfly, or the
anchialine pool shrimp, Procaris
hawaiana. It has been suggested that
transmission of disease from alien
invertebrates by mutual flower
visitation is a threat to the seven yellowfaced bees (Hylaeus spp.), but we
currently have no evidence that this is
occurring.
We consider predation and herbivory
by one or more of the nonnative animal
species (pigs, goats, axis deer, blacktailed deer, sheep, mouflon, cattle, rats,
barn owls, cats, mongooses, fish, slugs,
ants, black twig borers, and wasps) to
pose an ongoing threat to 35 of the 39
plant species and to all 10 animal
species throughout their ranges for the
following reasons:
(1) Observations and reports have
documented that pigs, goats, axis deer,
black-tailed deer, sheep, mouflon, and
cattle browse 27 of the 39 plant species,
in addition to other studies
demonstrating the negative impacts of
ungulate browsing on native plant
species of the islands. If the numbers
and range of blackbuck antelope
increase, their browsing will be a threat
to native plants that occur on Molokai,
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including host plants for the yellowfaced bees.
(2) Nonnative rats and slugs (either
singly or combined) cause mechanical
damage to plants and destruction of
plant parts (branches, flowers, fruits,
and seeds), and are considered a threat
to 22 of the 39 plant species.
(3) Rats also prey upon adults,
juveniles, and eggs of the band-rumped
storm-petrel, and are linked with the
dramatic decline of many closely related
bird species. Because rats are found in
all of the ecosystems in which the bandrumped storm-petrel occurs, we
consider predation by rats to be a
serious and ongoing threat.
(4) Barn owls and cats have
established populations in the wild on
all the main islands, and mongooses
have established populations on all the
main islands except for Kauai. All of
these nonnative animals are known to
prey on ground- and burrow-nesting
seabirds. Predation by these animals is
a serious and ongoing threat to the
band-rumped storm-petrel.
(5) The absence of Hawaiian
damselflies (including the orangeblack
Hawaiian damselfly) from ponds, pools,
and other aquatic habitat on the main
Hawaiian Islands is strongly correlated
with the presence of predatory
nonnative fish; numerous observations
and reports suggest nonnative predatory
fishes eliminate native damselflies from
these habitats. Accordingly, predation
by nonnative fishes is a serious and
ongoing threat to the orangeblack
Hawaiian damselfly. Predation by
bullfrogs, backswimmers, and Jackson’s
chameleons, and competition with
caddisflies are threats to the orangeblack
Hawaiian damselfly.
(6) Once introduced to anchialine
pools, nonnative fish, through predation
and competition for food sources,
directly affect anchialine pool shrimp,
including Procaris hawaiana, and also
disrupt anchialine pool ecology.
(7) Damage and destruction by the
black twig borer is a threat to two plant
species, Labordia lorenciana and
Nothocestrum latifolium.
(8) Predation by nonnative ants and
wasps poses a threat to all seven yellowfaced bees.
These threats are serious and ongoing,
act in concert with other threats to the
species, and are expected to continue or
increase in magnitude and intensity into
the future without effective management
actions to control or eradicate them. The
effects of the combined threats suggest
the need for immediate implementation
of recovery and conservation methods.
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Factor D. The Inadequacy of Existing
Regulatory Mechanisms
Overview
Currently, no existing Federal, State,
or local laws, treaties, or regulations
specifically conserve or protect 48 of the
49 species (except the band-rumped
storm-petrel by the Migratory Bird
Treaty Act (MBTA; 16 U.S.C. 703–712)),
or adequately address the threats to any
of the 49 species (see Table 2). There are
a few small programs and organizations
that conduct vegetation monitoring and
nonnative species and predator control,
but these activities are nonregulatory,
and neither continuation of these
conservation efforts nor funding for
them is guaranteed.
Federal laws pertaining to the 49
species addressed here include
Executive Order (E.O.) 13112, the
MBTA, the Lacey Act (16 U.S.C. 3371–
3378; 18 U.S.C. 42–43), the Federal
Noxious Weed List (7 CFR 360.200), and
the Convention on International Trade
in Endangered Species of Wild Fauna
and Flora (CITES). The U.S. Department
of Agriculture (USDA) inspects
propagative and restricted plant
materials and animals, and implements
‘‘Special Local Needs’’ rules for
pesticide use, but only on a species-byspecies basis. The Department of
Homeland Security-Customs and Border
Protection (CBP) is responsible for
inspecting commercial, private, and
military vessels and aircraft, and related
cargo and passengers arriving from
foreign locations. However, CBP focuses
on quarantine issues involving nonpropagative plant materials; wooden
packing materials, timber, and products;
internationally regulated commercial
species under CITES; and federally
listed noxious plants, seeds, soils, and
pests of concern to the continental
United States, such as pests to mainland
U.S. forests and agriculture.
Hawaii State law regarding natural
resource protections include those
under Hawaii revised statutes (HRS):
Plant and nondomestic animal
quarantine and microorganism import
(HRS 11–3–150A) and noxious weed
control (HRS 11–3–152); flood control
(HRS 12–2), water and land
development (HRS 12–174), and State
water code (HRS 12–2–174D); wildlife
(general wildlife, hunting, game birds,
game mammals, and wild birds and
other wildlife) (HRS 12–4–183D);
aquatic resources and wildlife-alien
aquatic organisms (HRS 12–5–187A);
general and miscellaneous, invasive
species council (HRS 12–6–194);
conservation of aquatic life, wildlife,
and land plants (HRS 12–6–195D); and
Natural Area Reserves (NARs) (HRS 12–
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6–195). These laws are interpreted and
implemented under Hawaii
administrative rules (HAR). Applicable
HARs include: Noxious weed rules
(HAR 4–6–68); plant and nondomestic
animal quarantine, microorganism
import rules (HAR 4–6-ch 71A, 71C),
and plant intrastate rules (HAR4–6–72);
rules regulating game mammal hunting
(HAR 13–5–2-ch 123; indigenous
wildlife, endangered and threatened
wildlife, and introduced wild birds
(HAR 13–5–2-ch 124); protection of
instream uses of water (HAR 13–7-ch
169), and NARs system (HAR 13–9-ch
208–210).
Private and local programs that
provide protections, and that help to
implement Federal and State
environmental regulations, laws, and
rules for one or more of the 49 species,
include the Hawaii Invasive Species
Committee (HISC), the Coordinating
Group on Alien Pest Species (CGAPS),
and the Hawaii Association of
Watershed Partnerships (HAWP). In
addition, the Plant Extinction Protection
Program (PEPP) was created to protect
Hawaii’s rare plant species in need of
immediate conservation efforts, by
monitoring, propagating, outplanting,
and providing some protection from
threats.
We discuss Federal and State
regulatory mechanisms, along with
agencies and groups authorized to
implement them, and the coordination
between them, below.
Federal Regulatory Mechanisms
On February 3, 1999, Executive Order
(E.O.) 13112 was signed establishing the
National Invasive Species Council
(NISC). This E.O. requires that a Council
of Departments dealing with invasive
species be created to prevent the
introduction of invasive species;
provide for their control; and minimize
the economic, ecological, and human
health impacts that invasive species
cause. Invasive species include aquatic
plant and animal species, terrestrial
plants and animal species, and plant
and animal pathogens. This E.O. was
reviewed in 2005 (NISC 2005). NISC
uses a cooperative approach to enhance
the Federal Government’s response to
the threat of invasive species, and
emphasizes prevention, early detection
and rapid response, and sharing of
information. See our discussion below
concerning the Hawaii Invasive Species
Committee (HISC) regarding the
effectiveness of this law.
The MBTA is the domestic law that
implements the United States’
commitment to four international
conventions (with Canada, Japan,
Mexico, and Russia) for the protection
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of shared migratory bird resources. The
MBTA regulates most aspects of take,
possession, transport, sale, purchase,
barter, export, and import of migratory
birds and prohibits the killing,
capturing, and collecting of individuals,
eggs, and nests, unless such action is
authorized by permit. While the MBTA
does prohibit actions that directly kill a
covered species, unlike the Endangered
Species Act (Act), it does not prohibit
habitat modification that indirectly kills
or injures a covered species, affords no
habitat protection when the birds are
not present, and provides only very
limited mechanisms for addressing
chronic threats to covered species, such
as nonnative predators.
The Lacey Act authorizes the
Secretary of the Interior to list as
‘‘injurious’’ any wildlife deemed to be
harmful to human beings, to the
interests of agriculture, horticulture,
forestry, or to wildlife or the wildlife
resources of the United States. The
Service inspects arriving wildlife
products, and enforces the injurious
wildlife provisions of the Lacey Act.
Among other provisions, the Lacey Act
prohibits importation of injurious
mammals, birds, fish, amphibians and
reptiles listed in the Lacey Act or which
are declared by the Secretary of the
Interior through regulation to be
injurious to human beings, agriculture,
horticulture, forestry or wildlife;
however, these prohibitions do not
apply to plants and organisms other
than those listed or designated by
regulations as injurious wildlife
(USFWS 2016, in litt.). The current list
of animals considered as ‘‘injurious
wildlife’’ is provided at 50 CFR part 16.
The list includes fruit bats, mongoose,
European rabbits and hares, wild dogs,
rats or mice, raccoon dogs, brushtail
possum (the species introduced to New
Zealand), starlings, house sparrows,
mynas, dioch, Java sparrows, red
whiskered bulbuls, walking catfish,
mitten crabs, zebra mussels, fish in the
snakehead family, four species of carp,
salmonids, brown tree snakes, and
pythons (USFWS 2012, 50 CFR part 16).
The Lacey Act requires declarations of
importation only for formal entries (i.e.,
commercial shipments), but not for
informal entries (i.e., personal
shipments) (USDA–APHIS 2015, in
litt.). Additionally, a species may still be
imported or transported across State
lines while it is being considered for
addition to the list of ‘‘injurious
wildlife’’ (Fowler et al. 2007, pp. 357–
358). Mongoose, rabbits, rats, mice,
house sparrows, mynas, Java sparrows,
and red whiskered bulbuls are already
established in Hawaii, are difficult and
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costly to control, or are not controlled
at all. None of the aquatic species on the
injurious species list is present in
Hawaii.
The continued spread of injurious
species indicates the limited
effectiveness of the Lacey Act in
preventing introductions of such species
to the State (Fowler et al. 2007, p. 357).
As an example of continued
introduction of nonnative species in
Hawaii, opossums (Didelphis
virginiana) have been found in shipping
containers on Oahu in 2005, 2011, and
most recently in 2015 (Star Advertiser
2015b, in litt). This species is not
included on the Lacey Act’s list of
injurious wildlife. Opossums are
omnivorous scavengers, consuming a
wide variety of food items including
insects, small vertebrates, bird eggs,
slugs and snails, and fruits and berries
(Oregon Department of Fish and
Wildlife 2015, in litt.; Clermont College
2015, in litt.). If opossums were to
establish wild populations in Hawaii,
their predation on ground-nesting
seabirds, insects, and snails could
negatively affect the band-rumped storm
petrel, the orangeblack Hawaiian
damselfly, one or more of the 39 plants,
and endangered snail species.
The Department of AgricultureAnimal and Plant Health Inspection
Service-Plant Protection and Quarantine
(USDA–APHIS–PPQ) inspects
propagative plant material, provides
identification services for arriving
plants and animals, conducts pest risk
assessments, and other related matters,
but focuses on pests of wide concern
across the United States (HDOA 2009, in
litt.). The USDA–APHIS–PPQ’s
Restricted Plants List restricts the
import of a limited number of noxious
weeds. If not specifically prohibited,
current Federal regulations allow plants
to be imported from international ports
with some restrictions. The Federal
Noxious Weed List (see 7 CFR 360.200;
USDA 2012) includes more than 100 of
the many globally known invasive
plants, 21 of which are already
established in Hawaii. Plants on the list
do not require a weed risk assessment
prior to importation from international
ports.
A local organization (under the
Institute of Pacific Islands ForestryUSFS), Pacific Island Ecosystems at Risk
(PIER) has compiled a complete list of
those plant species that are a threat to
ecosystems in the Pacific Islands, and
those that are potentially invasive and
are present in the Pacific Region, along
with a weed-risk assessment for most of
them (https://www.hear.org/pier/, last
updated May 15, 2013). There are over
1,000 plant species on the PIER list,
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and, in our proposed rule (80 FR 58820,
September 30, 2015; see pp. 58869–
58881), we discuss 114 of these invasive
plants that currently have the greatest
impacts on the 49 species. In addition,
the USDA–APHIS–PPQ is in the process
of finalizing rules to include a weed risk
assessment for plants newly imported to
Hawaii (and that may not yet appear on
the PIER list).
Water extraction is a threat to the
plant species (Cyclosorus boydiae), the
orangeblack Hawaiian damselfly, and
the anchialine pool shrimp Procaris
hawaiana. The U.S. Army Corps of
Engineers (COE) has regulatory
jurisdiction under section 404 of the
Clean Water Act (33 U.S.C. 1251 et seq.)
for activities that would result in a
discharge of dredged or fill material into
waters of the United States; however, in
issuing permits for such activities, the
COE does not typically establish
minimal instream flow standards (IFS)
as a matter of policy (U.S. Army 1985,
RGL 85–6).
State Regulatory Mechanisms
The Hawaii Endangered Species law
(HRS 195D) prohibits take, possession,
sale, transport or commerce in
designated species. This includes
aquatic as well as terrestrial animal
species, and terrestrial plants (not
freshwater or marine plants). This State
law also recognizes as endangered or
threatened those species determined to
be endangered or threatened pursuant to
the Act. This Hawaii law states that a
threatened species (under the Act) or an
indigenous species may be determined
to be an endangered species under State
law. Protection of these species is under
the authority of Hawaii’s Department of
Land and Natural Resources, and under
administrative rule (HAR 13–5–2-Ch
124). Although this State law can
address threats such as habitat
modification, light attraction, and line
collision through HCPs that address the
effects of individual projects or
programs, it does not address the
pervasive threats to the 49 species posed
by nonnative predators and feral
ungulates.
The importation of nondomestic
animals, including aquatic species and
microorganisms, is regulated by a
permit system (HAR 4–71) managed
through the Hawaii Department of
Agriculture (HDOA). In addition,
transport of plants and plant parts
between Hawaiian Islands is managed
through the HDOA (HAR 4–6–72), but
only for those species that have already
been determined to be pest species. The
objective of these administrative rules is
to implement the requirements of HRS
11–3–150A. The list of nondomestic
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animals (HAR 4–71) is defined by
providing a list of those animals
considered domestic: dog, cat, horse, ass
(burro or donkey), cattle and beefalo,
sheep, goat, swine, pot-bellied pig,
alpaca, llama, rabbit, chicken, turkeys,
pigeons, ducks, geese, and their hybrids.
Examples of regulated pests are listed at
HAR 4–72, including nonnative insects,
slugs, insects, plants, and plant viruses
that can damage or harm commercial
crops. The HDOA’s Board of Agriculture
maintains lists of nondomestic animals
that are prohibited from entry, animals
without entry restrictions, or those that
require a permit for import and
possession. The HDOA requires a
permit to import animals, and
conditionally approves entry for
individual possession, businesses (e.g.,
pets and resale trade, retail sales, and
food consumption), or institutions.
However, habitat destruction and
modification, and predation, by feral
domestic animals (such as goat and cats,
respectively) are two primary threats to
the 49 species not addressed by the
HDOA prohibitions and permitting
process.
The State of Hawaii allows
importation of most plant taxa, with
limited exceptions, if shipped from
domestic ports (HLRB 2002; USDA–
APHIS–PPQ 2010; CGAPS 2009).
Hawaii’s plant import rules (HAR 4–70)
regulate the importation of 13 plant taxa
of economic interest, including
pineapple, sugarcane, palms, and pines.
Certain horticultural crops (e.g.,
orchids) may require import permits
and have pre-entry requirements that
include treatment or quarantine or both
either prior to or following entry into
the State.
Critical biosecurity gaps include
inadequate staffing, facilities, and
equipment for Federal and State
inspectors devoted to invasive species
interdiction (HLRB 2002; USDA–
APHIS–PPQ 2010; CGAPS 2009). In
recognition of these gaps, a State law
has been passed that allows the HDOA
to collect fees for quarantine inspection
of freight entering Hawaii (Act 36 (2011)
HRS 150A–5.3). Legislation enacted in
2011 (H.B. 1568) requires commercial
harbors and airports to provide
biosecurity and inspection facilities to
facilitate the movement of cargo through
ports. This bill is a significant step
toward optimizing biosecurity capacity
in the State, but only time will
determine its effectiveness (Act 2011
(11)). We believe there is a need for all
civilian and military port and airport
operations and construction to make
biosecurity concerns a core objective.
As an example, the threat of
introduction of nonnative species is
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evidenced by the 2013 discovery of
presence of the nonnative coconut
rhinoceros beetle (CRB, Oryctes
rhinoceros), which quickly spread from
its known point of introduction across
the island of Oahu in a few months
(HISC 2014, in litt. + maps; HDOA 2014,
in litt.). The CRB is considered one of
the most damaging insects to coconut
and African oil palm trees in southern
and southeast Asia, as well as the
western Pacific Islands, and could
devastate populations of native and
nonnative palm trees in Hawaii (GiblinDavis 2001 in HISC 2014, in litt.). A
rapid response team headed by HDOA
(with USDA, University of Hawaii, U.S.
Navy, and other partners) has set up
pheromone traps island-wide, and
capture and range delineation efforts are
ongoing, along with funding for support
services to control the CRB (HISC 2014,
in litt.). However, existing regulatory
mechanisms did not prevent the
introduction of this pest species into
Hawaii. These regulatory mechanisms,
such as HAR 71A and HAR 71C
(regarding release of nonnative species)
and H.B. 1568 (pertaining to State law
to enforce biosecurity measures),
therefore, are inadequate to prevent
introduction of nonnative species.
Efforts to control the CRB continue, but
whether those efforts will be effective is
unknown at this time.
Hawaii’s noxious weed law was
enacted to prevent the introduction and
transport of noxious weeds or their
seeds or vegetative reproductive parts
into any area that is reasonably free of
those noxious weeds (HRS 11–3–152),
and it states that the Hawaii Department
of Agriculture shall take necessary
measures to restrict the introduction
and establishment of specific noxious
weeds in such areas. Hawaii
administrative rule (HAR 4–6–68)
further defines the term ‘‘noxious weed’’
and the criteria for designation of plants
as such and criteria for designation of a
noxious weed ‘‘free area.’’ The list of
noxious weeds, compiled in 1992,
consists of 79 plant species, 49 of which
were not yet established in Hawaii.
Since that time, 20 species on the list
have become established in Hawaii:
Bocconia frutescens (plume poppy),
Cereus uruguayanus (spiny tree cactus),
Chromolaena odorata (siamweed),
Cortaderia jubata (Andean pampas
grass), Cytisus scoparius (Scotch
broom), Hyptis suaveolens (wild
spikenard), Malachra alceifolia
(malachra), Melastoma spp. (melastoma;
two species now established, M.
candidum and M. sanguineum),
Miconia spp. (miconia; M. calvescens
now on four islands), Passiflora
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pulchella (wingleaf passionfruit), Piper
aduncum (spiked pepper), Prosopis
juliflora (algarroba), Pueraria
phaseoloides (tropical kudzu), Rubus
sieboldii (Molucca raspberry), Senecio
madagascariensis (fireweed), Solanum
elaeagnifolium (silverleaf nightshade),
Solanum robustum (shrubby
nightshade), Solanum torvum
(turkeyberry), and Spartium junceum
(Spanish broom). Thus, despite State
legislation and regulations addressing
invasive and noxious species, their
entry into the State continues.
The State manages the use of surface
and ground water resources through the
Commission on Water Resource
Management (CWRM), as mandated by
the State Water Code (HRS 174, HAR
13–168–196). The State considers all
natural flowing surface water (streams,
springs, seeps) as State property (HRS
174C), and the DLNR has management
responsibility for the aquatic organisms
in these waters (HRS Annotated 1988,
Title 12 1992 Cumulative Supplement).
In Hawaii, instream flow is regulated by
establishing standards on a stream-bystream basis. The standards currently in
effect represent flow conditions in 1987
(status quo), the year the administrative
rules (State Water Code) were adopted
(HRS 174C–71, HAR title 13, ch 169–
44–49). Because of the complexity of
establishing instream flow standards
(IFS) for 376 perennial streams, the
Commission retains interim IFS at status
quo levels as set in 1987 (CWRM 2009,
in litt.; CRWM 2014, in litt). In the
Waiahole Ditch Combined Contested
Hearing on Oahu (1991–2006), the
Hawaii Supreme Court determined that
status quo interim IFS were not
adequate, and required the Commission
to reassess the IFS for Waiahole Ditch
and other streams Statewide (Cast No.
CCH–OA95–1; Maui Now.com, in litt.).
The Commission has been gathering
information to fulfill this requirement
since 2006, but no IFS
recommendations have been made to
date (CWRM 2008, p. 3–153; CRWM
2014, in litt.).
In addition, in the Hawaii Stream
Assessment Report (HDLNR 1990;
prepared in coordination with the
National Park Service (NPS)), the
Commission identified high-quality
rivers and streams (and portions thereof)
that may be placed within a Wild and
Scenic River System. This report ranked
70 out of 176 analyzed rivers and
streams as outstanding high-quality
habitat, and recommended that streams
meeting certain criteria be protected
from further development (HDLNR
1990, pp. xxi–xxiv). However, there is
no mechanism within the State’s Water
Code to designate and set aside these
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streams, or to identify and protect
stream habitat. Accordingly, damselfly
populations (including the orangeblack
Hawaiian damselfly) are at risk of
continued loss of habitat.
Hawaii’s DLNR Division of Aquatic
Resources (DAR) is responsible for
conserving, protecting, and enhancing
the State’s renewable resources of
aquatic life and habitat (HDAR 2015, in
litt.; DLNR–DAR 2003, p. 3–13). The
release of live nonnative fish or other
nonnative aquatic life into any waters of
the State is prohibited (HRS 187A–6.5),
and DAR has the authority to seize,
confiscate, or destroy as a public
nuisance any of these prohibited species
(HRS 187A–2; HRS 187A–6.5).
However, the DAR recognizes that
nonnative species continue to enter the
State and move between islands (DLNR–
DAR 2003, p. 2–12).
There are no existing regulatory
mechanisms that specifically protect
Hawaii’s anchialine pools (habitat for
the anchialine pool shrimp Procaris
hawaiana and the orangeblack Hawaiian
damselfly); however 2 anchialine pools
on Maui and 12 anchialine pool on
Hawaii Island are located within State
NARs. State NARs were created to
preserve and protect samples of
Hawaii’s ecosystems and geological
formations, and are monitored.
Designation as a State NAR prohibits the
removal of any native organism and the
disturbance of pools (HAR 13–209–4).
Though signs are posted at NARs to
notify the public that anchialine pools
are off-limits to bathers, off-road vehicle
use around the pools, and other
activities, the anchialine pools are in
remote areas and the State does not have
sufficient funding to effectively enforce
those restrictions.
Nonnative ungulates pose threats of
habitat destruction and modification
and predation (herbivory) to 37 of the 39
plants species, and of habitat
destruction and modification to 9 of the
10 animals in this rule (see Table 2).
The State provides opportunities to the
public to hunt game mammals
(ungulates including feral pigs and
goats, axis deer, black-tailed deer, and
mouflon, sheep and mouflon-sheep
hybrids) on 91 State-designated public
hunting areas (within 45 units) on all
the main Hawaiian Islands except
Kahoolawe and Niihau (HAR–DLNR
2010, 13–123; HDLNR 2009b, pp. 25–
30). On Niihau, public hunting
opportunities are managed by a private
business (Niihau Safaris Inc. 2015, in
litt.). The State’s management objectives
for game mammals range from
maximizing public hunting
opportunities (i.e., ‘‘sustained yield’’) in
some areas to removal by State staff or
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their designees from other areas (HAR–
DLNR 2010, p. 12–123; HDLNR 2009b,
pp. 25–30). Thirty of the 39 plant
species, the band-rumped storm-petrel,
the orangeblack Hawaiian damselfly,
and three yellow-faced bees have
populations in areas where habitat is
used for game enhancement and game
populations are maintained at levels for
public hunting (Holmes and Joyce 2009,
4 pp.; HAR–DLNR 2010, p. 12–123;
HBMP 2010). Public hunting areas are
defined, but not fenced, and game
mammals have unrestricted access to
most areas across the landscape,
regardless of underlying land-use
designation. While fences are sometimes
built to protect areas from game
mammals, the current number and
locations of fences are not adequate to
prevent habitat destruction and
modification for 46 of the 49 species.
One additional State regulation (HRS
12–183D) was enacted recently to
prevent intra-island transport of axis
deer only. There are no other State
regulations than those described above
that address protection of the species
and their habitat from feral ungulates.
Under statutory authorities provided
by HRS title 12, subtitle 4, 183D
Wildlife, the DLNR maintains HAR ch
124 (2014), which defines ‘‘injurious
wildlife’’ as ‘‘any species or subspecies
of animal except game birds and game
mammals which is known to be harmful
to agriculture, aquaculture, indigenous
wildlife or plants, or constitute a
nuisance or health hazard and is listed
in the exhibit entitled Exhibit 5, Chapter
13–124, List of Species of Injurious
Wildlife in Hawaii.’’ Under HAR 13–
124–3(d), ‘‘no person shall, or attempt
to: (1) Release injurious wildlife into the
wild; (2) Transport them to island or
locations within the State where they
are not already established and living in
a wild state; and (3) Export any such
species or the dead body or parts
thereof, from the State. Permits for these
actions may be considered on a case-bycase basis.’’ This law was enacted after
an incident in 2012 of interisland
transport of axis deer (for hunting
purposes) to Hawaii Island, which was
without axis deer previously.
Local Mechanisms
Local biologists and botanists
recognize the urgent need to address the
importation of nonnative, invasive
species, and are working to implement
actions required; however, their funding
is not guaranteed. We discuss the four
primary groups below.
In 1995, the Coordinating Group on
Alien Pest Species (CGAPS), a
partnership of managers from Federal,
State, County, and private agencies and
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organizations involved in invasive
species work in Hawaii, was formed in
an effort to coordinate policy and
funding decisions, improve
communication, increase collaboration,
and promote public awareness (CGAPS
2009). This group facilitated the
formation of the Hawaii Invasive
Species Council (HISC), which was
created by gubernatorial executive order
in 2002, to coordinate local initiatives
for the prevention of introduction and
for control of invasive species by
providing policy-level direction and
planning for the State departments
responsible for invasive species issues
(CGAPS 2009). In 2003, the Governor
signed into law Act 85, which conveys
statutory authority to the HISC to
continue to coordinate approaches
among the various State and Federal
agencies, and international and local
initiatives, for the prevention and
control of invasive species (HDLNR
2003, p. 3–15; HISC 2009, in litt.; HRS
194–2). Some of the recent priorities for
the HISC include interagency efforts to
control nonnative species such as the
plants Miconia calvescens (miconia) and
Cortaderia spp. (pampas grass), coqui
frogs (Eleutherodactylus coqui), the
CRB, and ants (HISC 2009, 2013, and
2015, in litt.; OISC 2015, in litt.; https://
dlnr.hawaii.gov/hisc). Budget cuts
beginning in 2009 restricted State
funding support of HISC, resulting in a
serious setback of conservation efforts
(HISC 2009; HISC 2015).
The Hawaii Association of Watershed
Partnerships comprises 11 separate
partnerships on six Hawaiian Islands.
These partnerships are voluntary
alliances of public and private
landowners, ‘‘committed to the common
value of protecting forested watersheds
for water recharge, conservation, and
other ecosystem services through
collaborative management’’ (https://
hawp.org/partnerships). Funding for the
partnerships is provided through a
variety of State and Federal sources,
public and private grants, and in-kind
services provided by the partners and
volunteers. However, budget cuts of 40
to 60 percent have occurred since 2009,
with serious impacts to the positive
contributions of these groups to
implementing the laws and rules that
can protect and control threats to one or
more of the 49 species.
Another group was established to
coordinate State and Federal agency
efforts in the protection of rare endemic
plant species in the State and Guam and
the Commonwealth of the Northern
Mariana Islands (CNMI), Hawaii’s Plant
Extinction Prevention Program (PEPP).
This program identifies and supports
the ‘‘rarest of the rare’’ plant species in
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need of immediate conservation efforts.
The goal of PEPP is to prevent the
extinction of plant species that have
fewer than 50 individuals remaining in
the wild.
These four partnerships, CGAPS,
HISC, HAWP, and PEPP, are stop-gap
measures that attempt to address issues
that are not resolved by individual State
and Federal agencies. The capacity of
State and Federal agencies and their
nongovernmental partners in Hawaii to
provide sufficient inspection services,
enforce regulations, and mitigate or
monitor the effects of nonnative species
is limited due to the large number of
taxa currently causing damage (CGAPS
2009). Many invasive, nonnative species
established in Hawaii currently have
limited but expanding ranges, and they
cause considerable concern. Resources
available to reduce the spread of these
species and counter their negative
effects are limited. Control efforts are
focused on a few invasive species that
cause significant economic or
environmental damage to commercial
crops and public and private lands.
Comprehensive control of an array of
nonnative species and management to
reduce disturbance regimes that favor
them remain limited in scope. If current
levels of funding and regulatory support
for control of nonnative species are
maintained, the Service expects existing
programs to continue to exclude, or, on
a very limited basis, control these
species only in the highest priority
areas. Threats from established
nonnative species are ongoing and are
expected to continue into the future.
As an example of current and future
challenges for biosecurity in Hawaii, a
strain of the plant rust Puccinia psidii
(ohia rust) was first noticed affecting
stands of the nonnative rose apple
(Syzygium jambos) and the native
Metrosideros (ohia) seedlings (both in
the plant family Myrtaceae) in nurseries
in 2005. Metrosideros spp. are a
dominant component of native forest in
Hawaii, providing watershed protection
and habitat for native wildlife. The
Hawaii Board of Agriculture
recommended a quarantine rule be
passed against the introduction of all
new strains of ohia rust (through
transmission on Myrtaceae species used
in the horticulture trade), to prevent
destruction of ohia forests and the risk
to agriculture and horticulture
industries (Environment Hawaii 2015,
pp. 1,8–9). However, the rule remains in
draft form and under review (HDOA
2015, in litt.), accessed August 1, 2016).
An example of the failure of biosecurity
in Hawaii and the speed with which a
new invader can cause widespread
destruction is the introduction of the
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gall wasp Quadrastichus erythrinae.
This highly destructive wasp was
detected in Taiwan in 2003. Despite
evidence of its rapid advance across the
Pacific Basin with concomitant loss of
populations of native and ornamental
trees in the genus Erythrina, this wasp
arrived and naturalized in Hawaii in
2005 (Gramling 2005, p. 1). The wasp
dispersed throughout the main
Hawaiian Islands within weeks, and as
a result, the endemic wiliwili, Erythrina
sandwicensis, was quickly devastated
(Rubinoff et al. 2010, p. 24).
On the basis of the information
provided above, existing State and
Federal regulatory mechanisms are not
preventing the introduction of
nonnative species and pathogens into
Hawaii via interstate and international
pathways, or via intrastate movement of
nonnative species between islands and
watersheds. Nor do these mechanisms
address the current threats posed to the
49 species by established nonnative
species. Therefore, State and Federal
regulatory mechanisms do not
adequately protect the 49 species, or
their habitats, from the threat of new
introductions of nonnative species or
the continued expansion of nonnative
species populations on and between
islands and watersheds. The impacts
from these threats are ongoing and are
expected to continue into the future.
Summary of Factor D
Existing State and Federal regulatory
mechanisms are not preventing the
introduction into Hawaii of nonnative
species or controlling the spread of
nonnative species between islands and
watersheds, or establishing or
maintaining instream flow standards.
Water extraction is a threat to one plant,
Cyclosorus boydiae, to the orangeblack
Hawaiian damselfly, and the anchialine
pool shrimp (Factor A). Habitat-altering
ungulates and nonnative plants (Factor
A) pose major ongoing threats to all 49
species addressed in this rule. Thirtyfive of the 39 plants and all 10 animals
experience the threat of predation or
herbivory by nonnative animals (Factor
C). The seven yellow-faced bees and the
orangeblack Hawaiian damselfly
experience competition with nonnative
insect species (Factor E). The
intentional or inadvertent introduction
of nonnative species and their spread
within Hawaii, and the damage caused
by existing populations of nonnative
species, continues despite existing
regulatory mechanisms designed to
address this threat (in all its
manifestations described above) to all
49 species. No existing regulatory
mechanisms effectively address
maintenance of instream flow, springs,
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seeps, and anchialine pools or address
the threats of water extraction and
stream modification for the anchialine
pool shrimp and orangeblack Hawaiian
damselfly. All of these threats are
ongoing and are expected to continue
into the future; therefore, we conclude
the existing regulatory mechanisms are
inadequate to reduce or eliminate these
threats to the 49 species.
Factor E. Other Natural or Manmade
Factors Affecting Their Continued
Existence
Other factors that pose a threat to
some or all of the 49 species include
artificial lighting and structures,
ingestion of marine debris and plastics,
dumping of trash and the introduction
of nonnative fish into anchialine pools,
recreational use of and sedimentation of
anchialine pools, low numbers of
individuals and populations,
hybridization, lack of or declining
regeneration, competition with
nonnative invertebrates, and loss of host
plants. Each threat is discussed in detail
below, along with identification of
which species are affected by these
threats. The impacts of climate change
to these species and their ecosystems
have the potential to exacerbate all of
the threats described below.
Artificial Lighting and Structures Effects
on the Band-Rumped Storm-Petrel
Artificial lights are a welldocumented threat to night-flying
seabirds such as petrels, shearwaters,
and storm-petrels (Croxall et al. 2012, p.
28). A significant impact to the bandrumped storm-petrel results from the
effects of artificial (night) lighting on
fledglings and, to a lesser degree, on
adults. Lighting of roadways, resorts,
ballparks, residences, and other
development, as well as on cruise ships
out at sea, both attracts and confuses
night-flying storm-petrels and other
seabirds (Harrison et al. 1990, p. 49;
Reed et al. 1985, p. 377; Telfer et al.
1987, pp. 412–413; Banko et al. 1991, p.
651). Storm-petrels use the night sky to
navigate and possibly to search for
bioluminescent marine prey (Telfer et
al. 1987, p. 412). Artificial lights can
attract night-flying seabirds and result
in ‘‘fallout’’ (birds becoming grounded)
when birds become confused and
exhaust themselves circling around
lights or collide with buildings,
powerlines, or other structures. Once
grounded, these birds are at risk of
predation or being run over by cars
(Reed et al. 1985, p. 377; Telfer et al.
1987, p. 410). Vulnerability to artificial
lighting varies among species and age
classes and is influenced by season,
lunar phase, and weather conditions.
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Young birds are more likely than adults
to become disoriented by manmade
light sources (Montevecchi 2006, pp.
101–102). Over a 12-year period (1978
to 1990), Harrison et al. (1990, p. 49)
reported that 15 band-rumped stormpetrels, 13 of which were fledglings,
were recovered on Kauai as a result of
fallout. Between 1991 and 2008, another
21 band-rumped storm-petrels were
collected on Kauai (Holmes and Joyce
2009, p. 2). Currently, fallout due to
light pollution is recorded almost
annually on Kauai (Kauai Island Utility
Cooperative 2015, in litt.). In addition,
band-rumped storm-petrels may be
attracted to lights at sea and collide with
boats; this source of injury and mortality
has been documented for other stormpetrel species (e.g., Black 2005, p. 67).
The actual extent of such loss and its
overall impact on the band-rumped
storm-petrel population in Hawaii is not
known because scavengers often prevent
the detection or recovery of the dead or
injured birds, and the scattered and
remote nesting areas of this species
preclude demographic monitoring to
quantify the impacts of this source of
mortality. However, given the probable
small total number of band-rumped
storm-petrels nesting in Hawaii and the
threats they face from nonnative
predators such as rats and cats, any
additional mortalities are likely to have
negative impacts on the population.
A related threat to seabirds in Hawaii,
including the band-rumped stormpetrel, is collision with structures such
as communication towers and utility
lines (Cooper and Day 1998, pp. 16–18;
Podolsky et al. 1998, pp. 23–33). Several
seabird species that nest in the
Hawaiian Islands, including the
Newell’s Townsend’s shearwater
(federally listed as threatened), the
Hawaiian petrel (federally listed as
endangered), and the band-rumped
storm-petrel, regularly commute
between inland nest sites and the ocean.
These birds commute at night, when
manmade obstacles such as
communication towers and utility lines
are difficult to see. They strike these
unseen obstacles, and often die or are
injured as a result. An early study
estimated that 340 Newell’s Townsend’s
shearwater fledglings die annually on
the eastern and southern shores of Kauai
as a result of collisions (Podolsky et al.
1998, p. 30); however, current analyses
for all seabirds on Kauai indicate the
number of collisions with utility lines is
much higher, over 2,000 strikes per year
(using site-specific strike rates), but
numbers of birds that hit utility lines is
site-dependent (Travers et al. 2014, pp.
19, 29–37; Service 2015, in litt., Slide
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21). Absent preventative measures, the
impact to the band-rumped storm-petrel
from artificial lighting and collisions
with structures is expected to increase
as the human population grows and
development continues on the Hawaiian
Islands.
Other Human Effects on the BandRumped Storm-Petrel
Other factors that may negatively
affect the band-rumped storm-petrel
include commercial fisheries
interactions and alteration of prey base
upon which the band-rumped stormpetrel depends. Commercial fisheries
are known to adversely affect certain
species of seabirds (Furness 2003, pp.
33–35). Seabirds are caught in fishing
gear and suffer mortality by drowning.
Seabirds also come into contact with
and consume deep-water fish to which
they would not normally have access,
and can become contaminated by high
levels of heavy metals in these fish
(Furness 2003, p. 34). Commercial
fisheries also cause depletion of small
pelagic schooling fish, a significant food
source for seabirds (Furness 2003, p.
34). The potential effects of these
activities have not been assessed for the
band-rumped storm-petrel; however,
storm-petrels have been observed to
attend fishing vessels (e.g., Yorio and
Caille 1999, p. 21; Yeh et al. 2013, p.
146), and the effects of fishery
interactions on this species are likely to
be similar to those documented for other
seabird species in the same order
(Procellariiformes or tubenoses;
albatrosses and petrels). In addition,
plastics and other debris in the open
ocean can be ingested accidentally by
band-rumped storm-petrels and pose a
threat to this species (Ryan 1989, p.
629). Although a study by Moser and
Lee (1992, p. 85) found no evidence of
plastic ingestion by band-rumped stormpetrels, the sample size was very small
(4 individuals) and inadequate to
conclusively determine whether this
species suffers from ingestion of
plastics. Other species of storm-petrels
have been documented to ingest plastics
(Bond and Lavers 2013, p. 3; Ryan 2015,
p. 20; Wilcox et al. 2015, p. 3), and
band-rumped storm-petrels are likely to
do so also. Many closely related
seabirds do suffer ill effects from
ingestion of plastics, including physical
damage to the digestive tract, effects of
toxins carried on the plastics, and
resulting mortality (Ryan 1989, pp. 623–
629; Tanaka et al. 2013, pp. 2–3).
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Effects of Recreational Use, and
Dumping of Trash and Nonnative Fish
Into Anchialine Pools
On Hawaii Island, it is estimated that
up to 90 percent of the anchialine pools
have been destroyed or altered by
human activities (Brock 2004, p. i). The
more recent human modification of
anchialine pools includes bulldozing
and filling of pools (Bailey-Brock and
Brock 1993, p. 354). Trampling damage
from use of anchialine pools for
swimming and bathing has been
documented (Brock 2004, pp. 13–17).
Historically, pools were sometimes
modified with stone walls and steps by
Hawaiians who used them for bathing.
There are no documented negative
impacts to pond biota as a result of this
activity; however, introduction of soaps
and shampoos is of concern (Brock
2004, p. 15).
The depressional features of
anchialine pools make them susceptible
to dumping. Refuse found in degraded
pools and pools that have been filled
with rubble have been dated to about
100 years old, and the practice of
dumping trash into pools continues
today (Brock 2004, p. 15). For example,
Lua O Palahemo (Hawaii Island) is
located approximately 560 ft (170 m)
from a sandy beach frequented by
visitors who fish and swim. There are
multiple dirt roads that surround the
pool making it highly accessible. Plastic
bags, paper, fishing line, water bottles,
soda cans, radios, barbed wire, and a
bicycle have been documented within
the pool (Kensley and Williams 1986,
pp. 417–418; Bozanic 2004, p. 1; Wada
2010, in litt.). Introduction of trash
involving chemical contamination into
anchialine pools, as has been observed
elsewhere on Hawaii Island (Brock
2004, pp. 15–16), drastically affects
water quality and results in local
extirpation of anchialine pool shrimp
species.
Anchialine pool habitats can
gradually disappear when wind-blown
materials accumulate through a process
known as senescence (Maciolek and
Brock 1974, p. 3; Brock 2004, pp. 11,
35–36). Conditions promoting rapid
senescence include an increased
amount of sediment deposition, good
exposure to light, shallowness, and a
weak connection with the water table,
resulting in sediment and detritus
accumulating within the pool instead of
being flushed away with tidal exchanges
and ground water flow (Maciolek and
Brock 1974, p. 3; Brock 2004, pp. 11,
35–36). Sedimentation degrades the
health of Hawaiian anchialine pool
systems in which the anchialine pool
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shrimp, Procaris hawaiana, and the
orangeblack Hawaiian damselfly occur.
In general, the accidental or
intentional introduction and spread of
nonnative fishes (bait and aquarium
fish) is considered the greatest threat to
anchialine pools in Hawaii (Brock 2004,
p. 16). Maciolek (1983, p. 612) found
that the abundance of shrimp in a given
population is indirectly related to
predation by fish. Lua O Palahemo is
vulnerable to the intentional dumping
of nonnative bait and aquarium fishes
because the area is accessible to vehicles
and human traffic, although due to its
remote location, it is not monitored
regularly by State agency staff. The
release of mosquito fish and tilapia into
the Waikoloa Anchialine Pond Preserve
(WAAPA) at Waikoloa, North Kona,
Hawaii, resulted in the infestation of all
ponds within an approximately 3-ha (8ac) area, which represented about twothirds of the WAAPA. Within 6 months,
all native hypogeal (subterranean)
shrimp species disappeared (Brock
2004, p. iii). Nonnative fish drive
anchialine species out of the lighted,
higher productivity portion of the pools,
into the surrounding water table bed
rock, subsequently leading to the
decimation of the benthic community
structure of the pool (Brock 2004, p. iii).
In addition, nonnative fish prey on and
exclude native hypogeal shrimp that are
usually a dominant and essential faunal
component of anchialine pool
ecosystems (Brock 2004, p. 16; BaileyBrock and Brock 1993, pp. 338–355).
The loss of the shrimp changes
ecological succession by reducing
herbivory of macroalgae, allowing an
overgrowth and change of pool flora.
This overgrowth changes the system
from clear, well-flushed basins to a
system characterized by heavy
sedimentation and poor water exchange,
which increases the rate of pool
senescence (Brock 2004, p. 16).
Nonnative fishes, unlike native fishes,
are able to complete their life cycles
within anchialine pool habitats, and
remain a permanent detrimental
presence in all pools in which they are
introduced (Brock 2004, p. 16). In
Hawaii, the most frequently introduced
fishes are those in the Poeciliidae family
(freshwater fish which bear live young)
and include mosquito fish, various
mollies (Poecilia spp.), and tilapia, that
prey on and exclude the herbivorous
aquatic animals upon which Procaris
hawaiana feeds. More than 90 percent
of the 600 to 700 anchialine habitats in
the State of Hawaii were degraded
between 1974 and 2004, due to the
introduction of nonnative fishes (Brock
2004, p. 24). According to Brock (2012,
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pers. comm.), sometime in the 1980s,
nonnative fishes were introduced into
Lua O Palahemo. It is our understanding
that the fish were subsequently removed
by illegal use of a fish poison (EPA
2007. pp. 22–23; Finlayson et al. 2010,
p. 2), and to our knowledge the pool is
currently free of nonnative fish;
however, nonnative fish could be
introduced into the pool at any time.
Low Numbers of Individuals and
Populations
Species that undergo significant
habitat loss and degradation, and other
threats resulting in population decline,
range reduction, and fragmentation, are
inherently highly vulnerable to
extinction because of localized
catastrophes such as hurricanes, floods,
rockfalls, landslides, treefalls, and
drought; climate change impacts;
demographic stochasticity; and the
increased risk of genetic bottlenecks and
´
inbreeding depression (Gilpin and Soule
1986, pp. 24–34). These conditions are
easily reached by island species and
especially species endemic to single
islands that face numerous threats such
as those described above (Pimm et al.
1988, p. 757; Mangel and Tier 1994, p.
607). Populations that have been
diminished and isolated by habitat loss,
predation, and other threats exhibit
reduced levels of genetic variability,
which diminishes the species’ capacity
to adapt to environmental changes,
thereby lessening the probability of
long-term persistence (Barrett and Kohn
1991, p. 4; Newman and Pilson 1997, p.
361). Very small, isolated plant
populations are also more susceptible to
reduced reproductive vigor due to
ineffective pollination, inbreeding
depression, and hybridization. This is
particularly true for functionally
unisexual plants like Myrsine fosbergii
of which some individuals are
functionally dioecious (male and female
flowers occur on separate individuals).
Isolated individuals have difficulty in
achieving natural pollen exchange,
which decreases the production of
viable seed. Populations are also
affected by demographic stochasticity,
through which populations are skewed
toward either male or female
individuals by chance. The problems
associated with small occurrence size
and vulnerability to random
demographic fluctuations or natural
catastrophes are further magnified by
interactions with other threats, such as
those discussed above (see Factor A and
Factor C, above).
Plants
The effects resulting from having a
reduced number of individuals and
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occurrences poses a threat to all 39
plant species addressed in this proposal.
We consider the following 19 species to
be especially vulnerable to extinction
due to threats associated with small
occurrence size or small number of
occurrences because:
• The only known occurrences of
Cyanea kauaulaensis, Labordia
lorenciana, Lepidium orbiculare, and
Phyllostegia helleri are threatened either
by landslides, rockfalls, treefalls,
drought, or erosion, or a combination of
these factors.
• Cyanea kauaulaensis, Cyrtandra
hematos, Gardenia remyi, Joinvillea
ascendens ssp. ascendens, Labordia
lorenciana, Nothocestrum latifolium,
and Ochrosia haleakalae numbers are
declining, and they have not been
observed regenerating in the wild.
• The only known wild individuals of
Cyperus neokunthianus, Kadua
haupuensis, and Stenogyne kaalae ssp.
sherffii are extirpated; there is one
remaining individual of Deparia
kaalaana, and only three individuals of
Phyllostegia brevidens. Kadua
haupuensis and Stenogyne kaalae ssp.
sherffii only exist in propagation.
• The following single-island
endemic species are known from fewer
than 250 individuals: Asplenium
diellaciniatum, Cyanea kauaulaensis,
Cyperus neokunthianus, Cyrtandra
hematos, Dryopteris glabra var. pusilla,
Hypolepis hawaiiensis var. mauiensis,
Kadua haupuensis, Labordia
lorenciana, Lepidium orbiculare,
Phyllostegia helleri, Pritchardia bakeri,
Santalum involutum, Stenogyne kaalae
ssp. sherffii, and Wikstroemia
skottsbergiana.
Animals
Like most native island biota, the
Hawaiian population of band-rumped
storm-petrel, the orangeblack Hawaiian
damselfly, the anchialine pool shrimp
(Procaris hawaiana), and the seven
yellow-faced bees are particularly
sensitive to disturbances due to their
diminished numbers of individuals and
populations, and small geographic
ranges.
The band-rumped storm-petrel is
represented in Hawaii by very small
numbers of populations, and perhaps
not more than a few hundred
individuals (Harrison et al. 1990, p. 49).
A single human-caused action such as
establishment of mongoose on Kauai, or
a hurricane during the breeding season,
could cause reproductive failure and the
mortality of a significant percentage of
the remaining individuals. Threats to
this species include habitat destruction
and modification, landslides and
erosion, hurricanes, predation, injury
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and mortality from lights and structures,
and other human factors (such as
commercial fisheries). The effects of
these threats are compounded by the
current low number of individuals and
populations of band-rumped stormpetrel.
We consider the orangeblack
Hawaiian damselfly vulnerable to
extinction due to impacts associated
with low numbers of individuals and
low numbers of populations because
this species is known from only five of
eight Hawaiian Islands (Hawaii Island,
Maui, Lanai, Molokai, and Oahu) where
it occurred historically, and because of
the current reduction in numbers on
each of those five islands. Jordan et al.
(2007, p. 247) conducted a genetic and
comparative phylogeography analysis (a
study of historical processes responsible
for genetic divergence within a species)
of four Hawaiian Megalagrion species,
including the orangeblack Hawaiian
damselfly. This analysis demonstrated
Megalagrion populations with low
genetic diversity are at greater risk of
decline and extinction than those with
high genetic diversity. The authors
found that low genetic diversity was
observed in populations known to be
bottlenecked or relictual (groups of
animals or plants that exist as a remnant
of a formerly widely distributed group),
including populations of the
orangeblack Hawaiian damselfly. The
following threats to this species have all
been documented: habitat destruction
and modification by agriculture and
urban development, droughts, floods,
and hurricanes; predation by nonnative
fish, backswimmers, bullfrogs, and
Jackson’s chameleons; competition with
caddisflies; and water extraction from
streams and ponds. The effects of these
threats are compounded by the current
low number of individuals and
populations of the orangeblack
Hawaiian damselfly.
We consider the anchialine pool
shrimp, Procaris hawaiana, vulnerable
to extinction due to impacts associated
with low numbers of individuals and
populations because this species is
known from only 25 of over 500
assessed anchialine pools on Hawaii
Island, and from only 2 anchialine pools
on Maui. Threats to P. hawaiana
include: Habitat destruction and
modification; agriculture and urban
development; commercial trade;
dumping of nonnative fish and trash
into anchialine pools; recreation; and
water extraction. The effects of these
threats are compounded by the low
number of individuals and populations
of P. hawaiana.
We consider the seven Hawaiian
yellow-faced bees vulnerable to
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extinction due to impacts associated
with low numbers of individuals and
populations. The seven yellow-faced
bee species currently occur in only 22
locations (with some overlap) on six
main Hawaiian Islands, and are
vulnerable to habitat change and
stochastic events due to low numbers
and occurrences (Daly and Magnacca
2003, p. 3; Magnacca 2007, p. 173).
Hylaeus anthracinus occurs in 15 total
locations from Hawaii Island, Maui,
Kahoolawe, Molokai, and Oahu, but has
not been recently observed in its last
known location on Lanai; H. assimulans
is found in 5 total locations on Maui,
Lanai, and Kahoolawe, but has not been
observed recently on Oahu or Molokai;
H. facilis is found in 2 total locations on
Oahu and Molokai, but has not been
observed recently from Lanai and Maui;
H. hilaris is known from one population
on Molokai and has not been observed
recently from Lanai and Maui; H.
kuakea is known from one small area on
Oahu; H. longiceps is known from 6
total locations on Maui, Lanai, Molokai,
and Oahu, but has not been collected
from several historical locations on
those islands; and H. mana is known
from 3 locations on Oahu. Threats to
these species include agriculture and
urban development; habitat destruction
and modification by nonnative
ungulates, nonnative plants, tsunamis,
fire, drought, and hurricanes; the effects
of climate change on habitat; loss of host
plants; and predation or competition by
nonnative ants, wasps, and bees. The
effects of these threats are compounded
by the low numbers of individuals and
populations of the seven yellow-faced
bees.
Hybridization
Natural hybridization is a frequent
phenomenon in plants and can lead to
the creation of new species (Orians
2000, p. 1949), or sometimes to the
decline of species through genetic
assimilation or ‘‘introgression’’
(Ellstrand 1992, pp. 77, 81; Levin et al.
1996, pp. 10–16; Rhymer and Simberloff
1996, p. 85). Hybridization, however, is
especially problematic for rare species
that come into contact with species that
are abundant or more common (Rhymer
and Simberloff 1996, p. 83). We
consider hybridization to be a threat to
Cyrtandra hematos, Microlepia strigosa
var. mauiensis, and Myrsine fosbergii
because it will lead to extinction of the
original genotypically distinct species
and varieties, as noted by biologists’
observations of occurrences (Kawelo
2009, in litt.; Ching Harbin 2015, in litt.;
Oppenheimer 2015, in litt.;).
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No Regeneration
Lack of, or low levels of, regeneration
(reproduction and recruitment) in the
wild has been observed, and is a threat
to seven plants: Cyanea kauaulaensis,
Cyrtandra hematos, Gardenia remyi,
Joinvillea ascendens ssp. ascendens,
Labordia lorenciana, Nothocestrum
latifolium, and Ochrosia haleakalae (see
Plants under ‘‘Low Numbers of
Individuals and Populations,’’ above).
The reasons for this are not well
understood; however, seed predation by
rats and ungulates, inbreeding
depression, and lack of pollinators are
thought to play a role (Wagner et al.
1999, p. 1451; Wood et al. 2007, p. 198;
HBMP 2010; Oppenheimer and Lorence
2012, pp. 20–21; PEPP 2010, p. 73; PEPP
2014, p. 34).
Competition With Nonnative
Invertebrates
There are 15 known species of
nonnative bees in Hawaii (Snelling
2003, p. 342), including two nonnative
Hylaeus species (Magnacca 2007, p.
188). Most nonnative bees inhabit areas
dominated by nonnative vegetation and
do not compete with Hawaiian bees for
foraging resources (Daly and Magnacca
2003, p. 13); however, the European
honey bee (Apis mellifera) is an
exception. This social species is often
very abundant in areas with native
vegetation and aggressively competes
with Hylaeus for nectar and pollen
(Hopper et al. 1996, p. 9; Daly and
Magnacca 2003, p. 13; Snelling 2003, p.
345). The European honey bee was first
introduced to the Hawaiian Islands in
1875, and currently inhabits areas from
sea level to the upper tree line boundary
(Howarth 1985, p. 156). Individuals of
the European honey bee have been
observed foraging on Hylaeus host
plants such as Scaevola spp. and
Sesbania tomentosa (ohai) (Hopper et
al. 1996, p. 9; Daly and Magnacca 2003,
p. 13; Snelling 2003, p. 345). Although
we lack information indicating
Hawaiian Hylaeus populations have
declined because of competition with
the European honey bee for nectar and
pollen, it does forage in Hylaeus habitat
and excludes Hylaeus species
(Magnacca 2007b, p. 188; Lach 2008, p.
155). Hylaeus species do not occur in
native habitat where there are large
numbers of European honey bee
individuals, but the impact of smaller,
more moderate populations is not
known (Magnacca 2007, p. 188).
Nonnative, invasive bees are widely
documented to decrease nectar volumes
and usurp native pollinators (Lach 2008,
p. 155). There are also indications that
populations of the European honey bee
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are not as vulnerable as Hylaeus species
to predation by nonnative ant species
(see Factor C. Disease or Predation,
above). As described above, Hylaeus
bees that collect pollen from flowers of
the native tree Metrosideros polymorpha
were absent from trees with flowers
visited by the big-headed ant, while
visits by the European honey bee were
not affected (Lach 2008, p. 155). As a
result, Lach (2008, p. 155) concluded
that the European honey bee may have
a competitive advantage over Hylaeus
species because it is not excluded by the
big-headed ant. Other nonnative bees
found in areas of native vegetation and
overlapping with native Hylaeus
population sites include Ceratina
species (carpenter bees), Hylaeus
albonitens (Australian colletid bees), H.
strenuus (NCN), and Lasioglossum
impavidum (NCN) (Magnacca 2007, p.
188; Magnacca and King 2013, pp. 19–
22).
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Loss of Host Plants Through
Competition
The seven yellow-faced bees are
dependent upon native flowering plants
for their food resources, pollen and
nectar, and for nesting sites. Introduced
invertebrates outcompete native
Hylaeus for use of host plants for pollen,
nectar, and nesting sites. This effect is
compounded by the impacts of
nonnative ungulates on native host
plants for Hylaeus (see discussion under
Factors A and C, above). Nonnative
plants are a threat to the seven yellowfaced bees and their host plants because
they (1) Degrade habitat and outcompete
native plants; (2) increase the intensity,
extent, and frequency of fire, converting
native shrubland and forest to land
dominated by nonnative grasses; and (3)
as a result of fire, cause the loss of the
native host plants upon which the
yellow-faced bees depend (Factor A).
Drought, fire, and water extraction lead
to loss of host plants within the known
ranges of populations of yellow-faced
bees, and are discussed under Factor A.
The Present or Threatened Destruction,
Modification, or Curtailment of Their
Habitat or Range, above.
Competition With Caddisflies
Caddisflies (Order Trichoptera), a
nonnative aquatic insect, were first
observed and identified in Hawaii in the
1940s (Flint et al. 2003, p. 31); several
species are established on all the main
Hawaiian Islands. They may have been
introduced inadvertently with aquarium
plants released into streams (Flint et al.
2003, p. 37). Stream sampling showed
that caddisflies accounted for 57 percent
of the stream benthos (flora and fauna
in stream sediment) in upper elevation
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Kauai streams (Englund et al. 2000, p.
23; Flint et al. 2003, p. 38), and
caddisflies now inhabit every Oahu
stream. Caddisflies compete with native
aquatic invertebrate for resources and
space (Haines 2015, in litt.), which may
reduce prey abundance for naiads of the
orangeblack Hawaiian damselfly. In
addition, caddisflies provide a food
source for introduced fish species,
contributing to successful establishment
of nonnative fish (Flint et al. 2003, p.
38), an additional threat to the
orangeblack Hawaiian damselfly.
Climate Change
Our analyses under the Act include
consideration of ongoing and projected
changes in climate, and the impacts of
global climate change and increasing
temperatures on Hawaii ecosystems are
the subjects of active research. Global
temperature has increased over the past
century, and particularly since the mid20th century (IPCC 2014, p. 5), and this
increase in temperature is correlated to
emissions of carbon dioxide and other
greenhouse gasses, which have
increased more since 1970 than in prior
periods (IPCC 2014, pp. 13–14).
Analysis of the historical record
indicates surface temperature in Hawaii
has been increasing since the early
1900s, with relatively rapid warming
over the past 30 years. The average
increase since 1975 has been 0.48 °F
(0.27 °C) per decade for annual mean
temperature at elevations above 2,600 ft
(800 m) and 0.16 °F (0.09 °C) per decade
for elevations below 800 m
(Giambelluca et al. 2008, pp. 3–4).
Relative to average global temperature
from 1986 to 2005, the average ambient
air temperature is likely to increase
globally by at least 0.5 to 4.7 °F (0.3 to
2.6 °C) by the year 2100 (IPCC 2013, p.
20). Based on models using climate data
downscaled for Hawaii, the ambient
temperature is projected to increase by
3.8 to 7.7 °F (2.1 to 4.3 °C), depending
upon elevation and the emission
scenario (Liao et al. 2015, p. 4344). On
the main Hawaiian Islands, predicted
changes associated with increases in
temperature include a shift in vegetation
zones upslope, a similar shift in animal
species’ ranges, changes in mean
precipitation with unpredictable effects
on local environments, increased
occurrence of drought cycles, and
increases in intensity and numbers of
hurricanes (Loope and Giambelluca
1998, pp. 514–515; U.S. Global Change
Research Program (US–GCRP) 2009, pp.
10, 12, 17–18, 32–33; Giambelluca 2013,
p. 6). Additionally, sea level is rising as
a result of thermal expansion of
warming ocean water; the melting of ice
sheets, glaciers, and ice caps; and the
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addition of water from terrestrial
systems (Climate Institute 2011, in litt.),
and sea-level rise negatively affects
species occurring in low-lying coastal
areas including Solanum nelsonii (Starr
2011, in litt.) and affects the stability of
anchialine pools systems that are habitat
for Procaris hawaiana (Sakihara 2015,
in litt.).
The forecast of changes in
precipitation is highly uncertain
because it depends, in part, on how the
˜
˜
El Nino–La Nina weather cycle (an
episodic feature of the oceanatmosphere system in the tropical
Pacific having important global
consequences for weather and climate)
might change (State of Hawaii 1998, pp.
2–10). The historical record indicates
that Hawaii tends to be dry (relative to
˜
a running average) during El Nino
˜
phases and wet during La Nina phases
(Chen and Chu 2005, pp. 4809–4810).
However, over the past century, the
Hawaiian Islands have experienced a
decrease in precipitation of just over 9
percent (US National Science and
Technology Council 2008, p. 61) and a
trend of decrease (from the long-term
mean) is evident in recent decades (Chu
and Chen 2005, pp. 4802–4803; Diaz et
al. 2005, pp. 1–3). Stream-gauge data
provide corroborating evidence of a
long-term decrease in precipitation and
stream flow on the Hawaiian Islands
(Oki 2004, p. 4). This long-term drying
trend, coupled with existing ditch
˜
diversions and periodic El Nino-caused
drying events, has created a pattern of
severe and persistent stream dewatering
events (Polhemus 2008, in litt., p. 26).
Models of future rainfall downscaled for
Hawaii generally project increasingly
wet windward slopes and mild to
extreme drying of leeward areas in
particular by the middle and end of the
21st century (Timm and Diaz 2009, p.
4262; Elison Timm et al. 2015, pp. 95,
103–105). Altered seasonal moisture
regimes can have negative impacts on
plant growth cycles and overall negative
impacts on native ecosystems (US–
GCRP 2009, pp. 32–33). Long periods of
decline in annual precipitation result in
a reduction of moisture availability, an
increase in drought frequency and
intensity, and a self-perpetuating cycle
of nonnative plant invasion, fire, and
erosion (US–GCRP 2009, pp. 32–33;
Warren 2011, pp. 221–226) (see ‘‘Habitat
Destruction and Modification by Fire,’’
above). Overall, the documented and
projected increase in variance of
precipitation events will change
patterns of water availability for the
species (Parmesan and Matthews 2006,
p. 340), changes that point to changes in
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plant communities as a consequence
over the coming decades.
Tropical cyclone frequency and
intensity are projected to change as a
result of increasing temperature and
changing circulation associated with
climate change over the next 100 to 200
years (Vecchi and Soden 2007, pp.
1068–1069, Figures 2 and 3; Emanuel et
al. 2008, p. 360, Figure 8; Yu et al. 2010,
p. 1371, Figure 14). In the central
Pacific, modeling projects an increase of
up to two additional tropical cyclones
per year in the main Hawaiian Islands
by 2100 (Murakami et al. 2013, p. 2,
Figure 1d). In general, tropical cyclones
with the intensities of hurricanes have
been an uncommon occurrence in the
Hawaiian Islands. From the 1800s until
1949, hurricanes were only rarely
reported from ships in the area. Between
1950 and 1997, 22 hurricanes passed
near or over the Hawaiian Islands, and
5 of these caused serious damage
(Businger 1998, in litt.). A recent study
shows that, with a possible shift in the
path of the subtropical jet stream
northward, away from Hawaii, more
storms will be able to approach and
reach the Hawaiian Islands from an
easterly direction, with Hurricane Iselle
in 2014 being an example (Murakami et
al. 2015, p. 751).
As described above (see Climate
Change Vulnerability Assessment for
the Hawaiian Plants, above, and Table
2), 27 of the 39 plant species in this
proposal were included in the recent
analysis of the vulnerability of Hawaiian
plants to climate changes conducted by
Fortini et al. (2013, 134 pp.). All 27
species scored as moderately to
extremely vulnerable, as did most other
species in the analysis that already are
considered to be of conservation
concern (because they face multiple
non-climate threats) (Fortini et al. 2013,
pp. 25, 37). The specific impacts of
climate change effects on the habitat,
biology, and ecology of individual
species are largely unknown and remain
a subject of study. However, in the
assessment of more than 1,000 Hawaiian
plants, including 319 already listed as
endangered or threatened, a strong
relationship emerged between climate
vulnerability scores and current threats
and conservation status (Fortini et al.
2013, p. 5). Therefore, we anticipate that
the 13 plant species not analyzed are
likely to be similarly vulnerable to
climate change effects. The projected
landscape- or island-scale changes in
temperature and precipitation, as well
as the potentially catastrophic impacts
of projected increases in storm
frequency and severity, also point to
likely adverse impacts of climate change
on all 10 of the animal species
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considered in this proposal because
they rely on abiotic conditions, such as
water temperature, or habitat elements,
such as host plants and prey species,
likely to be substantively altered by
climate change.
Although we lack information about
the specific effects of current and
projected climate change on these
species, we anticipate that increased
ambient temperature and hurricane
intensity, changing precipitation
patterns, and sea-level rise and
inundation will create additional
stresses on these species because they
are vulnerable to these disturbances. For
example, projected warmer
temperatures and increased storm
severity resulting from climate change
are likely to exacerbate other threats to
the species, such as by enhancing the
spread of nonnative invasive plants into
these species’ native ecosystems in
Hawaii. The drying trend, especially on
leeward sides of islands, creates suitable
conditions for increased invasion by
nonnative grasses and enhances the risk
of wildfire. Sea-level rise threatens
ecosystems and species nearest the
coast, including the anchialine pool
ecosystem.
The risk of extinction as a result of the
effects of climate change increases when
a species’ range and habitat
requirements are restricted, its habitat
decreases, and its numbers and number
of populations decline (IPCC 2014, pp.
14–15). The fragmented range,
diminished number of populations, and
low total number of individuals have
compromised the rangewide
redundancy and resilience of these 49
species. Therefore, we would expect
them to be particularly vulnerable to the
habitat impacts of the effects of climate
change (Loope and Giambelluca 1998,
pp. 504–505; Pounds et al. 1999, pp.
611–612; Still et al. 1999, p. 610;
Benning et al. 2002, pp. 14,246–14,248;
Giambelluca and Luke 2007, pp. 13–15).
Although we cannot predict the timing,
extent, or magnitude of specific impacts,
we do expect the effects of climate
change to exacerbate the current threats
to these species, such as habitat loss and
degradation.
In summary, based on the best
available information, we conclude that
climate change effects, including
increased inter-annual variability of
ambient temperature, precipitation, and
hurricanes, are likely to impose
additional stresses on all 11 ecosystems
and all of the 49 species we are listing
in this rule, thus exacerbating current
threats to these species. These 49
species all persist with small population
sizes and highly restricted or
fragmented ranges. They thus face
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increased immediate risk from
stochastic events such as hurricanes,
which can extinguish an important
proportion of the remaining individuals,
and from long-term, landscape-scale
environmental changes because reduced
populations often lack ecological or
genetic adaptive capacity (Fortini et al.
2013, pp. 3–5).
In addition to impacts resulting from
changes in terrestrial habitat and
disturbance regimes, climate change
affects aquatic habitat. For example,
physiological stress in the orangeblack
Hawaiian damselfly is caused by
increased water temperatures to which
the species is not adapted (Pounds et al.
1999, pp. 611–612; Still et al. 1999, p.
610; Benning et al. 2002, pp. 14246,
14248). All of these aspects of climate
change and their impacts on native
species and ecosystems will be
exacerbated by human demands on
Hawaii’s natural resources; for example,
decreased availability of fresh water will
magnify the impact of human water
consumption on Hawaii’s natural
streams and reservoirs (Giambelluca et
al. 1991, p. v). Climate change impacts
contribute to the multiple threats
affecting the status of all of these
species, and the effects of climate
change are projected to increase in the
future.
Summary of Factor E
We consider the threat from artificial
lighting and structures to be a serious
and ongoing threat to the band-rumped
storm-petrel in Hawaii because these
threats cause injury and mortality,
resulting in a loss of breeding
individuals and juveniles, and are
expected to continue into the future.
Injury or mortality or loss of food
sources caused by the activities of
commercial fisheries, and injury or
mortality resulting from ingestion of
plastics and marine debris, are likely to
contribute to further decline in the
Hawaiian population of the bandrumped storm-petrel.
We consider the threats from
recreational use of, and dumping of
trash and introduction of nonnative fish
into, the pools that support the
anchialine pool shrimp Procaris
hawaiana to be serious threats that have
the potential to occur at any time,
although their occurrence is not
predictable. The use of anchialine pools
for dumping of trash leads to
accelerated sedimentation in the pool,
exacerbating conditions leading to its
senescence. Changing the anchialine
pool system by dumping of trash,
introduction of nonnative fish, and
sedimentation also affects habitat for the
orangeblack Hawaiian damselfly. In
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addition, recreational use of off-road
vehicles contributes to increased
sedimentation in anchialine pools, and
has been noted to affect the habitat of
the orangeblack Hawaiian damselfly on
Lanai.
We consider the impacts from limited
numbers of individuals and populations
to be a serious and ongoing threat to all
39 plant species, and especially for the
following 19 plants: Asplenium
diellaciniatum Cyanea kauaulaensis,
Cyperus neokunthianus, Cyrtandra
hematos, Deparia kaalaana, Dryopteris
glabra var. pusilla, Gardenia remyi,
Hypolepis hawaiiensis var. mauiensis,
Joinvillea ascendens ssp. ascendens,
Kadua haupuensis, Labordia
lorenciana, Lepidium orbiculare,
Myrsine fosbergii, Phyllostegia
brevidens, P. helleri, Pritchardia bakeri,
Santalum involutum, Stenogyne kaalae
ssp. sherffii, and Wikstroemia
skottsbergiana, as low numbers and
small occurrences of these plants result
in greater vulnerability to stochastic
events and can result in reduced levels
of genetic variability leading to
diminished capacity to adapt to
environmental changes. Under these
circumstances, the likelihood of longterm persistence is diminished, and the
likelihood of extirpation or extinction is
increased. This threat applies to the
entire range of each of these species.
We also consider the impacts from
limited numbers of individuals and
populations to be a serious and ongoing
threat to the band-rumped storm-petrel,
the orangeblack Hawaiian damselfly, the
anchialine pool shrimp Procaris
hawaiana, and to the yellow-faced bees
(Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana). The threat
from limited numbers of individuals
and populations is ongoing and is
expected to continue into the future
because (1) A single catastrophic event
may result in extirpation of remaining
populations and extinction of these
species; (2) species with few known
occurrences are less resilient to threats
that might otherwise have a relatively
minor impact (on widely distributed
species); (3) these species experience
reduced reproductive vigor due to
inbreeding depression; and (4) they
experience reduced levels of genetic
variability leading to diminished
capacity to adapt to environmental
changes, thereby lessening the
probability of its long-term persistence.
The threat from hybridization is an
unpredictable but ongoing threat to
Cyrtandra hematos, Microlepia strigosa
var. mauiensis, and Myrsine fosbergii, as
has been observed at current
occurrences.
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We consider the threat to Cyanea
kauaulaensis, Cyrtandra hematos,
Gardenia remyi, Joinvillea ascendens
ssp. ascendens, Labordia lorenciana,
Nothocestrum latifolium, and Ochrosia
haleakalae from lack of regeneration to
be ongoing and to continue into the
future because the reasons for the lack
of recruitment in the wild are unknown
and uncontrolled, and any competition
from nonnative plants or habitat
modification by ungulates or fire, or
other threats could lead to the
extirpation of these species.
We consider the threat of competition
with nonnative invertebrates a serious
and ongoing threat to the yellow-faced
bees, Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana.
Nonnative wasps and bees are
aggressive and can prevent use of the
native host plants required for food and
nesting by all seven yellow-faced bees.
Competition with caddisflies is a threat
to the orangeblack Hawaiian damselfly.
Based on current and projected
changes in climate, increasing
temperature, changing precipitation
regimes, increases in storm severity, and
sea-level rise will likely exacerbate the
threats to these 49 species. The effects
of climate change on these species
include, but are not limited to,
physiological stress caused by increased
water or air temperature or lack of
moisture, the long-term destruction and
modification of habitat, increased
competition by nonnative species, and
changes in disturbance regimes that lead
to changes in habitat and direct
mortality of individuals (e.g., fire,
drought, flooding, and hurricanes).
Determination for 49 Species
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
CFR part 424, set forth the procedures
for adding species to the Federal Lists
of Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, we may list a species based on (A)
The present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; or (E)
other natural or manmade factors
affecting its continued existence. Listing
actions may be warranted based on any
of the above threat factors, singly or in
combination.
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to each of the 49
species. We find that all of these species
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face threats that are ongoing and are
expected to continue into the future
throughout their ranges. Habitat
destruction and modification by
agriculture and urban development, and
conversion of wetland habitat or water
extraction resulting from such activity,
is a threat to one plant, Cyclosorus
boydiae, and seven animals (the
orangeblack Hawaiian damselfly, the
anchialine pool shrimp (Procaris
hawaiana), Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H.
longiceps) (Factor A). Habitat
destruction and modification by
nonnative feral ungulates poses a threat
to 46 of the 49 species (except for
Cyanea kauaulaensis, Hypolepis
hawaiiensis var. mauiensis, and the
anchialine pool shrimp) (Factor A).
Habitat destruction and modification by
nonnative plants poses a threat to all 39
plant species and 9 of the 10 animals
(except for Procaris hawaiana) (Factor
A). Fourteen of the plant species
(Exocarpos menziesii, Festuca
hawaiiensis, Joinvillea ascendens ssp.
ascendens, Labordia lorenciana,
Nothocestrum latifolium, Ochrosia
haleakalae, Portulaca villosa,
Ranunculus mauiensis, Sanicula
sandwicensis, Santalum involutum,
Schiedea pubescens, Sicyos
lanceoloideus, S. macrophyllus, and
Solanum nelsonii) and all seven yellowfaced bees, are at risk of habitat
destruction and modification by fire.
Habitat loss and mortality resulting from
hurricanes is a threat to the plant
Pritchardia bakeri, the band-rumped
storm-petrel, the orangeblack Hawaiian
damselfly, and all seven yellow-faced
bees (Factor A). Twenty of the plant
species (Cyanea kauaulaensis,
Cyclosorus boydiae, Deparia kaalaana,
Dryopteris glabra var. pusilla, Gardenia
remyi, Joinvillea ascendens ssp.
ascendens, Kadua fluviatilis, K.
haupuensis, Labordia lorenciana,
Lepidium orbiculare, Phyllostegia
brevidens, P. helleri, P. stachyoides,
Portulaca villosa, Pseudognaphalium
sandwicensium var. molokaiense,
Ranunculus hawaiensis, R. mauiensis,
Sanicula sandwicensis, Schiedea
pubescens, and Solanum nelsonii), the
band-rumped storm-petrel, and the
orangeblack Hawaiian damselfly are
threatened by the destruction and
modification of their habitats from,
either singly or in combination,
landslides, rockfalls, treefalls, flooding,
or tsunamis (Factor A). Habitat loss or
degradation and loss of host plants,
mortality, and water extraction due to
drought is a threat to the plants
Cyclosorus boydiae, Deparia kaalaana,
Huperzia stemmermanniae, Phyllostegia
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stachyoides, Ranunculus hawaiensis, R.
mauiensis, Sanicula sandwicensis,
Schiedea pubescens, Sicyos
lanceoloideus, and Solanum nelsonii;
the orangeblack Hawaiian damselfly;
and all seven yellow-faced bees (Factor
A and Factor E). Unpermitted collection
for commercial purposes poses a serious
threat to the anchialine pool shrimp
Procaris hawaiana (Factor B). Predation
or herbivory is a serious and ongoing
threat to 35 of the 39 plant species (by
feral pigs, goats, axis deer, black-tailed
deer, cattle, sheep, mouflon, rats, slugs,
and the black twig borer), to the bandrumped storm petrel (by barn owls, cats,
rats, and mongoose), and to the seven
yellow-faced bees (by ants and wasps)
(Factor C). Predation by bullfrogs,
backswimmers, nonnative fish, and
Jackson’s chameleons is a threat to the
orangeblack Hawaiian damselfly (Factor
C). Predation by nonnative fish is a
threat to the anchialine pool shrimp
(Factor C). The existing regulatory
mechanisms do not adequately address
these threats to the 49 species (Factor
D). Injury and mortality caused by
artificial lighting and structures are
serious and ongoing threats to the bandrumped storm-petrel (Factor E). The
threats of injury or mortality, or loss of
food sources, caused by the activities of
commercial fisheries, and injury or
mortality resulting from ingestion of
plastics and marine debris, can
contribute to further decline of the
Hawaiian population of the bandrumped storm-petrel (Factor E).
Recreational use of, and dumping of
trash and nonnative fish into, anchialine
pools is a threat to the anchialine pool
shrimp and also to the orangeblack
Hawaiian damselfly that uses that
habitat (Factor E). Competition by ants,
wasps, and bees for the food and nesting
resources, including loss of native host
plants, is a threat to all seven yellowfaced bees. Competition with caddisflies
is a threat to the orangeblack Hawaiian
damselfly (Factor E). These threats are
exacerbated by these species’ inherent
vulnerability to extinction from
stochastic events at any time because of
their endemism, low numbers of
individuals and populations, and
restricted habitats. There are serious and
ongoing threats to all 49 species due to
factors associated with low numbers of
individuals and populations (Factor E).
The threat of low numbers to seven
plants (Cyanea kauaulaensis, Cyrtandra
hematos, Gardenia remyi, Joinvillea
ascendens ssp. ascendens, Labordia
lorenciana, Nothocestrum latifolium,
and Ochrosia haleakalae) is exacerbated
by lack of regeneration in the wild
(Factor E). Hybridization is a threat to
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three plant species, Cyrtandra hematos,
Microlepia strigosa var. mauiensis, and
Myrsine fosbergii (Factor E). The effects
of rising temperature and other aspects
of climate change are likely to
exacerbate many of these threats and
likely to pose threats to the 49 species
(Factor E).
The Act defines an endangered
species as any species that is ‘‘in danger
of extinction throughout all or a
significant portion of its range’’ and a
threatened species as any species ‘‘that
is likely to become endangered
throughout all or a significant portion of
its range within the foreseeable future.’’
We find that each of the endemic
Hawaiian species and the Hawaii DPS of
the band-rumped storm-petrel are
presently in danger of extinction
throughout their entire ranges. Based on
the immediacy, severity, scope, and
interaction of the threats described
above, such as the pervasive threats of
predation and habitat loss and
degradation posed by nonnative plants
and animals, a determination of
threatened status for any of these
species is not appropriate. Therefore, on
the basis of the best available scientific
and commercial information, we are
listing the following 49 species as
endangered in accordance with sections
3(6) and 4(a)(1) of the Act: the plants
Asplenium diellaciniatum,
Calamagrostis expansa, Cyanea
kauaulaensis, Cyclosorus boydiae,
Cyperus neokunthianus, Cyrtandra
hematos, Deparia kaalaana, Dryopteris
glabra var. pusilla, Exocarpos menziesii,
Festuca hawaiiensis, Gardenia remyi,
Huperzia stemmermanniae, Hypolepis
hawaiiensis var. mauiensis, Joinvillea
ascendens ssp. ascendens, Kadua
fluviatilis, Kadua haupuensis, Labordia
lorenciana, Lepidium orbiculare,
Microlepia strigosa var. mauiensis,
Myrsine fosbergii, Nothocestrum
latifolium, Ochrosia haleakalae,
Phyllostegia brevidens, Phyllostegia
helleri, Phyllostegia stachyoides,
Portulaca villosa, Pritchardia bakeri,
Pseudognaphalium sandwicensium var.
molokaiense, Ranunculus hawaiensis,
Ranunculus mauiensis, Sanicula
sandwicensis, Santalum involutum,
Schiedea diffusa ssp. diffusa, Schiedea
pubescens, Sicyos lanceoloideus, Sicyos
macrophyllus, Solanum nelsonii,
Stenogyne kaalae ssp. sherffii, and
Wikstroemia skottsbergiana; and the
following animals: the Hawaii DPS of
the band-rumped storm-petrel
(Oceanodroma castro), the orangeblack
Hawaiian damselfly (Megalagrion
xanthomelas), the anchialine pool
shrimp (Procaris hawaiana), and the
yellow-faced bees Hylaeus anthracinus,
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Hylaeus assimulans, Hylaeus facilis,
Hylaeus hilaris, Hylaeus kuakea,
Hylaeus longiceps, and Hylaeus mana.
Under the Act and our implementing
regulations, a species may warrant
listing if it is in danger of extinction or
likely to become so throughout all or a
significant portion of its range (SPR).
Under our SPR policy (79 FR 37578,
July 1, 2014), if a species is endangered
or threatened throughout a significant
portion of its range and the population
in that significant portion is a valid
DPS, we will list the DPS rather than the
entire taxonomic species or subspecies.
We have determined that the Hawaii
population of the band-rumped stormpetrel is a valid DPS, and we are listing
that DPS. Each of the other 48 species
endemic to the Hawaiian Islands that
we are listing in this rule is highly
restricted in its range, and the threats
occur throughout its range. Therefore,
we assessed the status of each species
throughout its entire range. In each case,
the threats to the survival of these
species occur throughout the species’
range and are not restricted to any
particular portion of that range.
Accordingly, our assessment and
determination applies to each species
throughout its entire range. Likewise,
we assessed the status of the Hawaii
DPS of the band-rumped storm-petrel
throughout the range of the DPS and
have determined that the threats occur
throughout the DPS and are not
restricted to any particular portion of
the DPS. Because we have determined
that these 48 species and one DPS are
endangered throughout all of their
ranges, no portion of their ranges can be
‘‘significant’’ for purposes of the
definitions of ‘‘endangered species’’ and
‘‘threatened species.’’ See the Final
Policy of Interpretation of the Phrase
‘‘Significant Portion of Its Range’’ in the
Endangered Species Act’s Definitions of
‘‘Endangered Species’’ and ‘‘Threatened
Species’’ (79 FR 37578, July 1, 2014).
Available Conservation Measures
Conservation measures provided to
species listed as endangered or
threatened species under the Act
include recognition, recovery actions,
requirements for Federal protection, and
prohibitions against certain practices.
Recognition through listing results in
public awareness and conservation by
Federal, State, and local agencies;
private organizations; and individuals.
The Act encourages cooperation with
the States and requires that recovery
actions be carried out for all listed
species. The protection required by
Federal agencies and the prohibitions
against certain activities are discussed,
in part, below.
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The primary purpose of the Act is the
conservation of endangered and
threatened species and the ecosystems
upon which they depend. The ultimate
goal of such conservation efforts is the
recovery of these listed species, so that
they no longer need the protective
measures of the Act. Subsection 4(f) of
the Act calls for the Service to develop
and implement recovery plans for the
conservation of endangered and
threatened species. The recovery
planning process involves the
identification of actions that are
necessary to halt or reverse the species’
decline by addressing the threats to its
survival and recovery. The goal of this
process is to restore listed species to a
point where they are secure, selfsustaining, and functioning components
of their ecosystems.
Recovery planning includes the
development of a recovery outline
shortly after a species is listed and
preparation of a draft and final recovery
plan. The recovery outline guides the
immediate implementation of urgent
recovery actions and describes the
process to be used to develop a recovery
plan. Revisions of the plan may be done
to address continuing or new threats to
the species, as new substantive
information becomes available. The
recovery plan identifies site-specific
management actions that set a trigger for
review of the five factors that control
whether a species remains endangered
or may be downlisted or delisted, and
methods for monitoring recovery
progress. Recovery plans also establish
a framework for agencies to coordinate
their recovery efforts and provide
estimates of the cost of implementing
recovery tasks. Recovery teams
(composed of species experts, Federal
and State agencies, nongovernmental
organizations, and stakeholders) are
often established to develop recovery
plans. When completed, the recovery
outline, draft recovery plan, and the
final recovery plan will be available on
our Web site (https://www.fws.gov/
endangered), or from our Pacific Islands
Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
Implementation of recovery actions
generally requires the participation of a
broad range of partners, including other
Federal agencies, States,
nongovernmental organizations,
businesses, and private landowners.
Examples of recovery actions include
habitat restoration (e.g., restoration of
native vegetation), research, captive
propagation and reintroduction, and
outreach and education. The recovery of
many listed species cannot be
accomplished solely on Federal lands
because their range may occur primarily
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or solely on non-Federal lands. To
achieve recovery of these species
requires cooperative conservation efforts
on all lands.
Following publication of this final
listing rule, funding for recovery actions
will be available from a variety of
sources, including Federal budgets,
State programs, and cost share grants for
non-Federal landowners, the academic
community, and nongovernmental
organizations. In addition, pursuant to
section 6 of the Act, the State of Hawaii
will be eligible for Federal funds to
implement management actions that
promote the protection or recovery of
the 49 species. Information on our grant
programs that are available to aid
species recovery can be found at: https://
www.fws.gov/grants.
Please let us know if you are
interested in participating in recovery
efforts for one or more of these 49
species. Additionally, we invite you to
submit any new information on these
species whenever it becomes available
and any information you may have for
recovery planning purposes (see FOR
FURTHER INFORMATION CONTACT).
Section 7(a) of the Act, as amended,
requires Federal agencies to evaluate
their actions with respect to any species
that is proposed or listed as endangered
or threatened and with respect to its
critical habitat, if any is designated.
Regulations implementing this
interagency cooperation provision of the
Act are codified at 50 CFR part 402.
Section 7(a)(2) of the Act requires
Federal agencies to ensure that activities
they authorize, fund, or carry out are not
likely to jeopardize the continued
existence of a listed species or destroy
or adversely modify its critical habitat.
If a Federal action may affect a listed
species or its critical habitat, the
responsible Federal agency must enter
into consultation with the Service.
Federal agency actions within the
species’ habitat that may require
conference or consultation or both as
described in the preceding paragraph
include, but are not limited to, actions
within the jurisdiction of the Natural
Resources Conservation Service (NRCS),
the U.S. Army Corps of Engineers, the
U.S. Fish and Wildlife Service, and
branches of the Department of Defense
(DOD). Examples of these types of
actions include activities funded or
authorized under the Farm Bill Program,
Environmental Quality Incentives
Program, Ground and Surface Water
Conservation Program, Clean Water Act
(33 U.S.C. 1251 et seq.), Partners for
Fish and Wildlife Program, and DOD
construction activities related to
training or other military missions.
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The Act and its implementing
regulations set forth a series of general
prohibitions and exceptions that apply
to endangered wildlife. The prohibitions
of section 9(a)(1) of the Act, codified at
50 CFR 17.21, make it illegal for any
person subject to the jurisdiction of the
United States to take (which includes
harass, harm, pursue, hunt, shoot,
wound, kill, trap, capture, or collect; or
to attempt any of these) endangered
wildlife within the United States or the
high seas. In addition, it is unlawful to
import; export; deliver, receive, carry,
transport, or ship in interstate or foreign
commerce in the course of commercial
activity; or sell or offer for sale in
interstate or foreign commerce any
listed species. It is also illegal to
possess, sell, deliver, carry, transport, or
ship any such wildlife that has been
taken illegally. Certain exceptions apply
to employees of the Service, the
National Marine Fisheries Service, other
Federal land management agencies, and
State conservation agencies.
We may issue permits to carry out
otherwise prohibited activities
involving endangered wildlife under
certain circumstances. Regulations
governing permits are codified at 50
CFR 17.22. With regard to endangered
wildlife, a permit may be issued for
scientific purposes, to enhance the
propagation or survival of the species,
or for incidental take in connection with
otherwise lawful activities. There are
also certain statutory exemptions from
the prohibitions, which are found in
sections 9 and 10 of the Act.
With respect to endangered plants,
prohibitions outlined at 50 CFR 17.61
make it illegal for any person subject to
the jurisdiction of the United States to
import or export, transport in interstate
or foreign commerce in the course of a
commercial activity, sell or offer for sale
in interstate or foreign commerce, or to
remove and reduce to possession any
such plant species from areas under
Federal jurisdiction. In addition, for
endangered plants, the Act prohibits
malicious damage or destruction of any
such species on any area under Federal
jurisdiction, and the removal, cutting,
digging up, or damaging or destroying of
any such species on any other area in
knowing violation of any State law or
regulation, or in the course of any
violation of a State criminal trespass
law. Exceptions to these prohibitions
are outlined at 50 CFR 17.62.
We may issue permits to carry out
otherwise prohibited activities
involving endangered plants under
certain circumstances. Regulations
governing permits are codified at 50
CFR 17.62. With regard to endangered
plants, the Service may issue a permit
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authorizing any activity otherwise
prohibited by 50 CFR 17.61 for scientific
purposes or for enhancing the
propagation or survival of endangered
plants.
It is our policy, as published in the
Federal Register on July 1, 1994 (59 FR
34272), to identify to the maximum
extent practicable at the time a species
is listed, those activities that would or
would not constitute a violation of
section 9 of the Act. The intent of this
policy is to increase public awareness of
the effect of a listing on proposed and
ongoing activities within the range of a
listed species. Based on the best
available information, activities that
may potentially result in a violation of
section 9 of the Act include but are not
limited to:
(1) Unauthorized collecting, handling,
possessing, selling, delivering, carrying,
or transporting of the species, including
import or export across State lines and
international boundaries, except for
properly documented antique
specimens of these taxa at least 100
years old, as defined by section
100(h)(1) of the Act;
(2) Activities that take or harm the
band-rumped storm-petrel, the
orangeblack Hawaiian damselfly, the
anchialine pool shrimp (Procaris
hawaiana), and the seven yellow-faced
bees by causing significant habitat
modification or degradation such that it
causes actual injury by significantly
impairing essential behavior patterns.
This may include introduction of
nonnative species that compete with or
prey upon the 10 animal species or the
unauthorized release of biological
control agents that attack the life stage
of any of these 10 species; and
(3) Damaging or destroying any of the
39 plant species in violation of the
Hawaii State law prohibiting the take of
listed species.
Common name
Questions regarding whether specific
activities would constitute a violation of
section 9 of the Act should be directed
to the Pacific Islands Fish and Wildlife
Office (see FOR FURTHER INFORMATION
CONTACT). Requests for copies of the
regulations concerning listed species
and general inquiries regarding
prohibitions and permits may be
addressed to the U.S. Fish and Wildlife
Service, Pacific Region, Ecological
Services, Endangered Species Permits,
Eastside Federal Complex, 911 NE. 11th
Avenue, Portland, OR 97232–4181
(telephone 503–231–6131; facsimile
503–231–6243).
Required Determinations
National Environmental Policy Act (42
U.S.C. 4321 et seq.)
We have determined that
environmental assessments and
environmental impact statements, as
defined under the authority of the
National Environmental Policy Act
(NEPA; 42 U.S.C. 4321 et seq.), need not
be prepared in connection with listing
a species as an endangered or
threatened species under the
Endangered Species Act. We published
a notice outlining our reasons for this
determination in the Federal Register
on October 25, 1983 (48 FR 49244).
References Cited
A complete list of references cited in
this rulemaking is available on the
Internet at https://www.regulations.gov
under Docket No. FWS–R1–ES–2015–
0125 and upon request from the Pacific
Islands Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
Authors
The primary authors of this final rule
are the staff members of the Pacific
Islands Fish and Wildlife Office.
List of Subjects in 50 CFR Part 17
Endangered and threatened species,
Exports, Imports, Reporting and
recordkeeping requirements,
Transportation.
Regulation Promulgation
Accordingly, we amend part 17,
subchapter B of chapter I, title 50 of the
Code of Federal Regulations, as follows:
PART 17—ENDANGERED AND
THREATENED WILDLIFE AND PLANTS
1. The authority citation for part 17
continues to read as follows:
■
Authority: 16 U.S.C. 1361–1407; 1531–
1544; and 4201–4245, unless otherwise
noted.
2. Amend § 17.11(h), the List of
Endangered and Threatened Wildlife, as
follows:
■ a. By adding an entry for ‘‘Stormpetrel, band-rumped (Hawaii DPS)’’ in
alphabetical order under BIRDS;
■ b. By adding entries for ‘‘Bee, yellowfaced’’ (Hylaeus anthracinus), ‘‘Bee,
yellow-faced’’ (Hylaeus assimulans),
‘‘Bee, yellow-faced’’ (Hylaeus facilis),
‘‘Bee, yellow-faced’’ (Hylaeus hilaris),
‘‘Bee, yellow-faced’’ (Hylaeus kuakea),
‘‘Bee, yellow-faced’’ (Hylaeus
longiceps), ‘‘Bee, yellow-faced’’
(Hylaeus mana), and ‘‘Damselfly,
orangeblack Hawaiian’’ (Megalagrion
xanthomelas) in alphabetical order
under INSECTS; and
■ c. By adding an entry for ‘‘Shrimp,
anchialine pool’’ (Procaris hawaiana)
before the entry for ‘‘Shrimp, anchialine
pool’’ (Vetericaris chaceorum) under
CRUSTACEANS.
The additions read as follows:
■
§ 17.11 Endangered and threatened
wildlife.
*
Scientific name
Where listed
*
Oceanodroma castro ................
*
*
U.S.A. (HI) ................................
*
*
(h) * * *
Status
*
*
Listing citations and applicable
rules
BIRDS
*
*
Storm-petrel, band-rumped (Hawaii DPS).
asabaliauskas on DSK3SPTVN1PROD with RULES
*
*
INSECTS
Bee, yellow-faced ......................
Hylaeus anthracinus .................
Wherever found ........................
E
Bee, yellow-faced ......................
Hylaeus assimulans .................
Wherever found ........................
E
Bee, yellow-faced ......................
Hylaeus facilis ..........................
Wherever found ........................
E
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Common name
Scientific name
Where listed
Bee, yellow-faced ......................
Hylaeus hilaris ..........................
Wherever found ........................
E
Bee, yellow-faced ......................
Hylaeus kuakea ........................
Wherever found ........................
E
Bee, yellow-faced ......................
Hylaeus longiceps ....................
Wherever found ........................
E
Bee, yellow-faced ......................
Hylaeus mana ..........................
Wherever found ........................
E
*
*
Damselfly, orangeblack Hawaiian.
*
Megalagrion xanthomelas ........
*
*
Wherever found ........................
E
*
CRUSTACEANS
*
*
*
Shrimp, anchialine pool .............
*
*
*
*
Procaris hawaiana ....................
*
*
3. Amend § 17.12(h), the List of
Endangered and Threatened Plants, as
follows:
■ a. By adding entries for Calamagrostis
expansa, Cyanea kauaulaensis, Cyperus
neokunthianus, Cyrtandra hematos,
Exocarpos menziesii, Festuca
hawaiiensis, Gardenia remyi, Joinvillea
ascendens ssp. ascendens, Kadua
fluviatilis, Kadua haupuensis, Labordia
lorenciana, Lepidium orbiculare,
Myrsine fosbergii, Nothocestrum
latifolium, Ochrosia haleakalae,
Phyllostegia brevidens, Phyllostegia
■
Scientific name
Status
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*
*
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*
*
*
*
Wherever found ........................
*
*
*
*
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E
*
helleri, Phyllostegia stachyoides,
Portulaca villosa, Pritchardia bakeri,
Pseudognaphalium sandwicensium var.
molokaiense, Ranunculus hawaiensis,
Ranunculus mauiensis, Sanicula
sandwicensis, Santalum involutum,
Schiedea diffusa ssp. diffusa, Schiedea
pubescens, Sicyos lanceoloideus, Sicyos
macrophyllus, Solanum nelsonii,
Stenogyne kaalae ssp. sherffii, and
Wikstroemia skottsbergiana in
alphabetical order under FLOWERING
PLANTS; and
Listing citations and applicable
rules
*
*
b. By adding entries for Asplenium
diellaciniatum, Cyclosorus boydiae,
Deparia kaalaana, Dryopteris glabra var.
pusilla, Huperzia stemmermanniae,
Hypolepis hawaiiensis var. mauiensis,
and Microlepia strigosa var. mauiensis
in alphabetical order under FERNS AND
ALLIES.
The additions read as follows:
■
§ 17.12
*
Endangered and threatened plants.
*
*
(h) * * *
Common name
Where listed
Status
*
*
Calamagrostis expansa .............
*
Maui reedgrass .........................
*
*
Wherever found ........................
E
*
*
Cyanea kauaulaensis ................
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Cyperus neokunthianus ............
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Cyrtandra hematos ....................
*
Haiwale .....................................
*
*
Wherever found ........................
E
*
*
Listing citations and applicable
rules
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Common name
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*
*
Exocarpos menziesii .................
*
Heau .........................................
*
*
Wherever found ........................
E
Festuca hawaiiensis ..................
No common name ....................
Wherever found ........................
E
*
*
Gardenia remyi ..........................
*
Nanu .........................................
*
*
Wherever found ........................
E
*
Joinvillea ascendens ssp.
ascendens.
*
Ohe ...........................................
*
*
Wherever found ........................
E
*
*
Kadua fluviatilis .........................
*
Kamapuaa ................................
*
*
Wherever found ........................
E
Kadua haupuensis ....................
No common name ....................
Wherever found ........................
E
*
*
Labordia lorenciana ...................
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Lepidium orbiculare ...................
*
Anaunau ...................................
*
*
Wherever found ........................
E
*
*
Myrsine fosbergii .......................
*
Kolea ........................................
*
*
Wherever found ........................
E
*
*
Nothocestrum latifolium .............
*
Aiea ..........................................
*
*
Wherever found ........................
E
*
*
Ochrosia haleakalae .................
*
Holei .........................................
*
*
Wherever found ........................
E
*
*
Phyllostegia brevidens ..............
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Phyllostegia helleri ....................
asabaliauskas on DSK3SPTVN1PROD with RULES
Scientific name
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Phyllostegia stachyoides ...........
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Portulaca villosa ........................
*
Ihi ..............................................
*
*
Wherever found ........................
E
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*
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Scientific name
Common name
Where listed
*
*
Pritchardia bakeri ......................
*
Baker’s loulu .............................
*
*
Wherever found ........................
E
*
Pseudognaphalium
sandwicensium var.
molokaiense.
*
Enaena .....................................
*
*
Wherever found ........................
E
*
*
Ranunculus hawaiensis ............
*
Makou .......................................
*
*
Wherever found ........................
E
Ranunculus mauiensis ..............
Makou .......................................
Wherever found ........................
E
*
*
Sanicula sandwicensis ..............
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Santalum involutum ...................
*
Iliahi ..........................................
*
*
Wherever found ........................
E
*
*
Schiedea diffusa ssp. diffusa ....
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Schiedea pubescens .................
*
Maolioli .....................................
*
*
Wherever found ........................
E
*
*
Sicyos lanceoloideus .................
*
Anunu .......................................
*
*
Wherever found ........................
E
Sicyos macrophyllus .................
Anunu .......................................
Wherever found ........................
E
*
*
Solanum nelsonii .......................
*
Popolo ......................................
*
*
Wherever found ........................
E
*
*
Stenogyne kaalae ssp. sherffii ..
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Wikstroemia skottsbergiana ......
*
Akia ...........................................
*
*
Wherever found ........................
E
asabaliauskas on DSK3SPTVN1PROD with RULES
*
FERNS AND ALLIES
*
*
*
*
Status
*
*
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*
*
*
Asplenium diellaciniatum ...........
*
No common name ....................
*
*
Wherever found ........................
E
*
*
Cyclosorus boydiae ...................
*
Kupukupu makalii .....................
*
*
Wherever found ........................
E
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Scientific name
Common name
Where listed
Deparia kaalaana ......................
No common name ....................
Wherever found ........................
E
*
*
Dryopteris glabra var. pusilla ....
*
Hohiu ........................................
*
*
Wherever found ........................
E
*
*
Huperzia stemmermanniae .......
*
No common name ....................
*
*
Wherever found ........................
E
Hypolepis hawaiiensis var.
mauiensis.
Olua ..........................................
Wherever found ........................
E
*
No common name ....................
*
*
Wherever found ........................
E
*
Microlepia strigosa var.
mauiensis.
*
*
*
*
*
Status
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*
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*
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*
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*
*
Dated: September 12, 2016.
Stephen Guertin,
Acting Director, U.S. Fish and Wildlife
Service.
[FR Doc. 2016–23112 Filed 9–29–16; 8:45 am]
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BILLING CODE 4333–15–P
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Agencies
[Federal Register Volume 81, Number 190 (Friday, September 30, 2016)]
[Rules and Regulations]
[Pages 67786-67860]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2016-23112]
[[Page 67785]]
Vol. 81
Friday,
No. 190
September 30, 2016
Part V
Department of the Interior
-----------------------------------------------------------------------
Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Endangered Status for 49
Species From the Hawaiian Islands; Final Rule
Federal Register / Vol. 81 , No. 190 / Friday, September 30, 2016 /
Rules and Regulations
[[Page 67786]]
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R1-ES-2015-0125; 4500030113]
RIN 1018-BB07
Endangered and Threatened Wildlife and Plants; Endangered Status
for 49 Species From the Hawaiian Islands
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Final rule.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), determine
endangered status under the Endangered Species Act of 1973 (Act), as
amended, for 10 animal species, including the Hawaii DPS of the band-
rumped storm-petrel (Oceanodroma castro), the orangeblack Hawaiian
damselfly (Megalagrion xanthomelas), the anchialine pool shrimp
(Procaris hawaiana), and seven yellow-faced bees (Hylaeus anthracinus,
H. assimulans, H. facilis, H. hilaris, H. kuakea, H. longiceps, and H.
mana), and for 39 plant species from the Hawaiian Islands. This rule
adds these species to the Federal Lists of Endangered and Threatened
Wildlife and Plants.
DATES: This rule is effective October 31, 2016.
ADDRESSES: This final rule is available on the Internet at https://www.regulations.gov and at https://www.fws.gov/pacificislands. Comments
and materials we received, as well as supporting documentation we used
in preparing this rule, are available for public inspection at https://www.regulations.gov, or, by appointment, during normal business hours
at: U.S. Fish and Wildlife Service, Pacific Islands Fish and Wildlife
Office, 300 Ala Moana Boulevard, Honolulu, HI 96850; telephone 808-792-
9400; or facsimile 808-792-9581.
FOR FURTHER INFORMATION CONTACT: Mary M. Abrams, Ph.D., Field
Supervisor, Pacific Islands Fish and Wildlife Office, 300 Ala Moana
Boulevard, Honolulu, HI 96850; telephone 808-792-9400; or facsimile
808-792-9581. Persons who use a telecommunications device for the deaf
(TDD) may call the Federal Information Relay Service (FIRS) at 800-877-
8339.
SUPPLEMENTARY INFORMATION:
Executive Summary
Why we need to publish a rule. Under the Endangered Species Act
(Act), a species may warrant protection through listing if it is
endangered or threatened throughout all or a significant portion of its
range. Listing a species as an endangered or threatened species can
only be completed by issuing a rule. Critical habitat is to be
designated, to the maximum extent prudent and determinable, for any
species determined to be an endangered or threatened species under the
Act.
This rule makes final the listing of 10 animal species (the Hawaii
DPS of the band-rumped storm-petrel (Oceanodroma castro), the
orangeblack Hawaiian damselfly (Megalagrion xanthomelas), the
anchialine pool shrimp (Procaris hawaiana), and seven yellow-faced bees
(Hylaeus anthracinus, H. assimulans, H. facilis, H. hilaris, H. kuakea,
H. longiceps, and H. mana)), and 39 plant species (Asplenium
diellaciniatum (no common name, NCN), Calamagrostis expansa (Maui
reedgrass), Cyanea kauaulaensis (NCN), Cyclosorus boydiae (kupukupu
makalii), Cyperus neokunthianus (NCN), Cyrtandra hematos (haiwale),
Deparia kaalaana (NCN), Dryopteris glabra var. pusilla (hohiu),
Exocarpos menziesii (heau), Festuca hawaiiensis (NCN), Gardenia remyi
(nanu), Huperzia stemmermanniae (NCN), Hypolepis hawaiiensis var.
mauiensis (olua), Joinvillea ascendens ssp. ascendens (ohe), Kadua
fluviatilis (kamapuaa), Kadua haupuensis (NCN), Labordia lorenciana
(NCN), Lepidium orbiculare (anaunau), Microlepia strigosa var.
mauiensis (NCN), Myrsine fosbergii (kolea), Nothocestrum latifolium
(aiea), Ochrosia haleakalae (holei), Phyllostegia brevidens (NCN),
Phyllostegia helleri (NCN), Phyllostegia stachyoides (NCN), Portulaca
villosa (ihi), Pritchardia bakeri (Baker's loulu), Pseudognaphalium
sandwicensium var. molokaiense (enaena), Ranunculus hawaiensis (makou),
Ranunculus mauiensis (makou), Sanicula sandwicensis (NCN), Santalum
involutum (iliahi), Schiedea diffusa ssp. diffusa (NCN), Schiedea
pubescens (maolioli), Sicyos lanceoloideus (anunu), Sicyos macrophyllus
(anunu), Solanum nelsonii (popolo), Stenogyne kaalae ssp. sherffii
(NCN), and Wikstroemia skottsbergiana (akia), as endangered species.
Delineation of critical habitat requires identification of the
physical or biological features essential to the species' conservation.
A careful assessment of the biological needs of the species and the
areas that may have the physical or biological features essential for
the conservation of the species and that may require special management
considerations or protections, and thus qualify for designation as
critical habitat, is required. We require additional time to analyze
the best available scientific data in order to identify specific areas
appropriate for critical habitat designation and to analyze the impacts
of designating such areas as critical habitat. Accordingly, we find
designation of critical habitat to be ``not determinable'' at this
time.
The basis for our action. Under the Act, we can determine that a
species is an endangered or threatened species based on any of five
factors: (A) The present or threatened destruction, modification, or
curtailment of its habitat or range; (B) overutilization for
commercial, recreational, scientific, or educational purposes; (C)
disease or predation; (D) the inadequacy of existing regulatory
mechanisms; or (E) other natural or manmade factors affecting its
continued existence. We have determined that these 49 species are
experiencing population-level impacts as the result of the following
current and ongoing threats:
Habitat loss and degradation due to urbanization;
nonnative feral ungulates (hoofed mammals, e.g., pigs, goats, axis
deer, black-tailed deer, mouflon, and cattle); nonnative plants;
wildfire; and water extraction.
Predation or herbivory by nonnative feral ungulates, rats,
slugs, bullfrogs, Jackson's chameleons, ants, and wasps.
Stochastic events such as landslides, flooding, drought,
tsunami, and hurricanes.
Human activities such as recreational use of anchialine
pools, dumping of nonnative fish and trash into anchialine pools, and
manmade structures and artificial lighting.
Vulnerability to extinction due to small numbers of
individuals and occurrences and lack of regeneration.
Competition with nonnative plants and nonnative
invertebrates.
Existing regulatory mechanisms and conservation efforts are not
adequate to ameliorate the impacts of these threats on any of the 49
species such that listing is not warranted. Environmental effects from
climate change are likely to exacerbate the impacts of these threats.
Peer review and public comment. We sought comments from independent
specialists to ensure that our designation is based on scientifically
sound data, assumptions, and analyses. We invited these peer reviewers
to comment on our listing proposal. We also considered all comments and
information we received during two comment periods, including at one
public hearing.
[[Page 67787]]
Previous Federal Actions
Please refer to the proposed listing rule for the 49 species from
the Hawaiian Islands (80 FR 58820; September 30, 2015) for a detailed
description of previous Federal actions concerning these species.
Summary of Comments and Recommendations
On September 30, 2015, we published a proposed rule to list 49
species (39 plants and 9 animals) from the Hawaiian Islands as
endangered throughout their ranges and the Hawaii population (distinct
population segment (DPS)) of the band-rumped storm-petrel as endangered
(80 FR 58820). The comment period for the proposed rule lasted 60 days,
ending November 30, 2015 We published a public notice of the proposed
rule in the local Honolulu Star Advertiser, West Hawaii Today, Hawaii
Tribune-Herald, Molokai Dispatch, The Maui News, and The Garden Island
newspapers at the beginning of the comment period. We received two
requests for a public hearing. On January 22, 2016 (81 FR 3767), we
reopened the comment period for an additional 30 days, ending on
February 22, 2016, and we announced a public meeting and public hearing
for the proposed rule. We again published a public notice in local
newspapers and provided the public notice to local media. For both
comment periods, we requested that all interested parties submit
comments or information concerning the proposed listing of the 49
species. We contacted all appropriate State and Federal agencies,
county governments, elected officials, scientific organizations, and
other interested parties and invited them to comment. The public
meeting and hearing were held in Hilo, Hawaii, on February 9, 2016.
During the comment periods, we received a total of 41 unique public
comment letters (including comments received at the public hearing) on
the proposed listing of the 49 species. Of the 41 commenters, 21 were
peer reviewers, 3 were Federal agencies (Hawaii Volcanoes National
Park, Haleakala National Park, and Kaloko-Honokohau and Puuhonua o
Honaunau National Historical Parks (NHPs)), 4 were State of Hawaii
agencies (Hawaii Department of Health, Hawaii Department of Land and
Natural Resources Division of Aquatic Resources, Hawaii Division of
Forestry and Wildlife, and Hawaii Department of Hawaiian Home Lands),
and 13 were nongovernmental organizations or individuals (including
those who provided comments or testimony at the public hearing). The
National Park Service (NPS) provided new information about the numbers
and range of species in this rule that occur on NPS lands, and about
graduate research on the orangeblack Hawaiian damselfly. We appreciate
the time and effort taken by all commenters to submit their views and
information, and we have incorporated all substantive new information,
e.g., from the National Park Service, into this final rule. However, we
received some comments from the public on the possible future
designation of critical habitat and on a variety of other topics. To
the extent that comments do not pertain to the proposed listing rule,
we do not address them in this final rule. In this final rule, we
address only those comments relevant to the listing of the 49 species
from the Hawaiian Islands.
All substantive information related to the listing action provided
during the comment periods has either been incorporated directly into
this final rule, or is addressed below. For readers' convenience, we
have combined similar comments into a single comment and response.
Peer Review
In accordance with our peer review policy published in the Federal
Register on July 1, 1994 (59 FR 34270), we solicited expert opinions
from 29 knowledgeable individuals with scientific expertise on one or
more of the 49 Hawaiian Islands species, which include 39 plants, a
seabird, a damselfly, an anchialine pool shrimp, and seven yellow-faced
bees, and their habitats. This expertise also included familiarity with
the geographic region in which these species occur and conservation
biology principles. We received responses from 21 of these individuals.
We reviewed all comments we received from the peer reviewers for
substantive issues and new information regarding the 49 species. Of
these 21 peer reviewers, 18 provided comments or new information on one
or more of the 49 species. Ten peer reviewers stated support for the
proposed listing, and 11 were neutral regarding the proposed listing.
These peer reviewers generally supported our methodology and
conclusions. Peer reviewer comments are either addressed below or are
incorporated into this final rule as appropriate.
(1) Comment: One peer reviewer stated that sea-level rise and
coastal inundation collectively are also potential future threats to
the welfare of Procaris hawaiana, because they may cause further loss
of anchialine pool habitat.
Our Response: We have added sea-level rise and coastal inundation
as threats to P. hawaiana and its habitat under the discussion in this
rule titled ``Climate Change'' (Factor E. Other Natural or Manmade
Factors Affecting Their Continued Existence).
(2) Comment: One peer reviewer stated that because sea-level rise
could increase surface connectivity between currently isolated
anchialine pools, invasion by nonnative fish would be exacerbated.
Our Response: In this rule, we have added surface connectivity to
our summary description of the status and stressors to P. hawaiana as a
factor likely to exacerbate the threat posed by nonnative fish to this
species and its anchialine pool habitat (see Anchialine pool shrimp
(Procaris hawaiana), under Summary of Biological Status of the 49
Hawaiian Islands Species).
(3) Comment: One peer reviewer recommended that the island of
Lanai, and coastal habitat, be included as habitat for the band-rumped
storm-petrel, as birds were observed during the breeding season
transiting this habitat, which is conducive to nesting where crevices
and ledges are numerous and can provide some protection from feral cats
(Felis catus), goats (Capra hircus), and mouflon (Ovis gmelini
musimon).
Our Response: We have added coastal habitat on Lanai in our
description of habitat for the band-rumped storm-petrel in this final
listing rule.
(4) Comment: One peer reviewer recommended that coastal habitat on
leeward east Maui be included for the band-rumped storm-petrel, as
remains of a chick were found there in 1999.
Our Response: We understand that coastal habitat on east Maui may
be part of the species' historical range, but we have not added coastal
areas on leeward east Maui as currently occupied habitat for the band-
rumped storm-petrel in this final rule. Unlike coastal Lanai, in
coastal areas on leeward east Maui, no indication of the species'
presence or use of this habitat has been observed for 17 years.
(5) Comment: One peer reviewer stated that predation by bullfrogs
(Lithobates catesbeianus) should be included as a threat to the
orangeblack Hawaiian damselfly, and that impacts of backswimmers
(Notonectidae family) and caddisflies (Trichoptera order) on the
damselfly are speculative.
Our Response: We have included in this final rule that bullfrogs
are a threat to the orangeblack Hawaiian damselfly, and clarified that
the effects of
[[Page 67788]]
predation by backswimmers and caddisflies are not well understood.
Comments From State Agencies
(6) Comment: The Hawaii Department of Land and Natural Resources'
Division of Forestry and Wildlife did not comment in support of, or in
opposition to, the proposed listing of the 49 species from the Hawaiian
Islands. District botanists from Kauai, Oahu, Maui, and Hawaii Island
provided plant species occurrence updates by island.
Our Response: We appreciate the information provided regarding the
49 plant species from the Hawaiian Islands, and have incorporated it
into the Summary of Biological Status of the 49 Hawaiian Islands
Species for the appropriate species in this final rule.
(7) Comment: The Hawaii Department of Health acknowledged that
protecting wildlife and plants can often be important for human and
environmental health. They further commented that managing and
controlling wild ungulates is necessary for 95 percent of these
proposed plant species, the orangeblack Hawaiian damselfly (Megalagrion
xanthomelas), and the yellow-faced bees (Hylaeus spp.), but that it is
also essential to preventing erosion, and, therefore, protecting water
quality. Fire is a natural process that is now unnaturally frequent,
intense, and destructive to the Hawaiian Islands, in part due to
invasive grasses. Mitigating wildfires is essential to caring for 38
percent of the plant species, the damselfly, and yellow-faced bees, but
it also limits the release of air pollutants that are known to be
harmful to human health. Protection of coastal and wetland habitat such
as that populated by the anchialine pool shrimp (Procaris hawaiana)
limits further human pressures on our sensitive coastlines and aquatic
environments.
Our Response: We agree that managing and controlling ungulates
would provide significant conservation benefits to listed plant and
animal species, and would also prevent erosion and protect water
quality of the islands and near shore reefs. We also acknowledge that
nonnative grasses contribute to the increase in numbers and intensity
of wildfires in Hawaii. Protection of coastal habitat (through
nonnative plant and ungulate control, and prevention of wildfires)
would provide a conservation benefit to the anchialine pool shrimp, and
to other species that depend on coastal habitat.
(8) Comment: The Hawaii Department of Land and Natural Resources
Division of Aquatic Resources concurred that the information in the
proposed rule for the anchialine pool shrimp, Procaris hawaiana, is the
most accurate and up-to-date information available, and supported
listing the species as endangered under the Act (16 U.S.C. 1531 et
seq.).
Our Response: We appreciate this support for the proposed listing
of the anchialine pool shrimp, Procaris hawaiana.
(9) Comment: The Department of Hawaiian Home Lands (DHHL) asked
that the Secretary of the Interior consider the effects of designation
of endangered species that may potentially have critical habitat on
Hawaiian Home Lands in a similar manner to the effects such designation
has on tribal lands, including the impact on tribal sovereignty. DHHL
is aware that Secretarial Order 3206, issued in June 1997, establishes
guidelines for the Service when dealing with Indian tribes relating to
endangered species. Secretarial Order 3206 recognizes that, in order to
respect the cultural and social aspects of Indian tribes, some
environmental restrictions on Indian tribal lands are not appropriate,
and it calls on the Service to preserve endangered species while
respecting tribal authority over their own lands. While native
Hawaiians are not an ``Indian tribe'' under the Order, DHHL's mission,
to place native Hawaiians on its lands for residential, agricultural,
and pastoral homesteading purposes, is analogous to the circumstances
of Indian tribes. The Department also recommends that the Secretaries
of the Interior and Commerce, in determining endangered species and
critical habitat designations, consult directly with the Hawaiian Homes
Commission, DHHL, Office of Native Hawaiian Relations, and
beneficiaries of the Hawaiian Homes Commission Act to include native
intelligence and knowledge on species, habitat, and place-based
management and protection.
Our Response: In accordance with the President's memorandum of
April 29, 1994 (Government-to-Government Relations With Native American
Tribal Governments; 59 FR 22951), Executive Order 13175 (Consultation
and Coordination With Indian Tribal Governments), and the Department of
the Interior's manual at 512 DM 2, we readily acknowledge our
responsibility to communicate meaningfully with recognized Federal
Tribes on a government-to-government basis. In accordance with
Secretarial Order 3206 of June 5, 1997 (American Indian Tribal Rights,
Federal-Tribal Trust Responsibilities, and the Endangered Species Act),
we readily acknowledge our responsibilities to work directly with
tribes in developing programs for healthy ecosystems; to incorporate
native intelligence and knowledge of species, habitat, and place-based
management and protection; to acknowledge that tribal lands are not
subject to the same controls as Federal public lands; to remain
sensitive to Indian culture; and to make information available to
tribes. In addition, a 2004 consolidated appropriations bill (Pub. L.
108-199, see section 148) established the Office of Native Hawaiian
Relations within the Secretary of the Interior's Office, and its duties
include effectuating and implementing the special legal relationship
between the Native Hawaiian people and the United States, and fully
integrating the principle and practice of meaningful, regular, and
appropriate consultation with the Native Hawaiian people by assuring
timely notification of and prior consultation with the Native Hawaiian
people before any Federal agency takes any actions that may have the
potential to significantly affect Native Hawaiian resources, rights, or
lands. A 2011 memorandum of understanding (MOU) signed by the
Department of the Interior states that ``Federal agencies are required
to consult with Native Hawaiian organizations before taking any action
that may have the potential to significantly affect Native Hawaiian
resources, rights, or lands.'' Although native Hawaiians are not
technically a ``recognized Federal tribe'' as referenced in the above
Executive and Secretarial Orders, we endeavor to fully engage and work
directly with native Hawaiians as much as possible. At the time we
published our proposed rule (80 FR 58820; September 30, 2015), we
notified several Hawaiian organizations including the DHHL, Kamehameha
Schools, the Office of Hawaiian Affairs, the Kahoolawe Island Reserve
Commission (KIRC), and Kahea-The Hawaiian-Environmental Alliance. We
contacted the Department of the Interior's Office of Native Hawaiian
Relations on September 28, 2015, to inform them of our proposed listing
action. We also conducted in-person meetings with staff of the
Department of Hawaiian Home Lands, Kamehameha Schools, and KIRC. We
considered all comments and recommendations provided by these
organizations in developing this final listing rule. At the time we
prepare a proposed critical habitat rule for these species, we will
notify these groups and organizations, and carefully consider any
comments and new information they provide regarding habitat for these
species.
[[Page 67789]]
Public Comments
Seven public commenters supported listing of all 49 Hawaiian
Islands species. Seven public commenters opposed the listing of the 49
Hawaiian Islands species, and one of these commenters supported the
intent of listing but opposed designation of critical habitat on their
lands.
(10) Comment: One commenter supported this rule because of the
facts and analysis stated in the proposed rule. Two commenters stated
that humans need to be a voice for plants and animals, and that this
listing will positively impact the conservation of many animals and
positively lead other conversations in the right direction.
Our Response: We appreciate the comments and believe that listing
status will help provide conservation benefits to the species and their
habitats.
(11) Comment: One commenter stated that the 49 species also play a
pivotal role in promoting tourism and building the economy of Hawaii
and that they deserve to be put onto the Lists of Endangered and
Threatened Wildlife and Plants. Two commenters stated that listing
these species will attract wildlife enthusiasts and nature lovers from
all around the world, and their spending and tourism helps to build and
maintain sources of revenue in Hawaii; most markets within the islands
depend on the tourism dollars that wildlife attracts.
Our Response: We do not consider economic consequences in our
decisions to list or not list species as endangered or threatened under
the Act. Section 4(b)(1)(a) of the Act specifies that listing
determinations be made ``solely on the basis of the best scientific and
commercial data available.''
(12) Comment: One commenter stated that the potential negative
impacts of listing to landowners is very small, as the vast majority of
the habitat for these rare species occurs on State and Federal lands,
or in private lands devoted to conservation.
Our Response: We agree that many of the 49 species occur or were
known from State and Federal lands, or in undeveloped areas already
dedicated to conservation. However, listing a species as endangered or
threatened is based on the species' biological status; the development
of a proposed rule for critical habitat for these species will be
completed in a separate rule, and the effects of critical habitat on
landowners will be analyzed upon preparation of that proposed rule.
(13) Comment: One commenter stated that island residents have
entirely lost historical and cultural opportunities and rights as a
result of species protection enforcement and that those in the field of
endangered species protection have a single focus, with little or no
concern for cultural and historical values. Another commenter stated
that this listing would cause a further loss for the public of
cultural, historical, and economic resources. A third commenter stated
that native Hawaiian society believes they should be able to manage
their people, land, and resources autonomously.
Our Response: Listing a species as endangered or threatened does
not cause loss of historical and cultural opportunities; in fact, it
highlights the need to protect the characteristics that are unique to
the Hawaiian Islands. We acknowledge that some economic impacts are a
possible consequence of listing a species under the Act; for example,
there may be costs to the landowner associated with the development of
a habitat conservation plan (HCP). In other cases, if the landowner
does not acquire a permit for incidental take (for animals), the
landowner may choose to forego certain activities on their property to
avoid violating the Act, resulting in potential lost income. However,
the Act does not provide for the consideration of such impacts when
making a listing decision. Section 4(b)(1)(a) of the Act specifies that
listing determinations be made ``solely on the basis of the best
scientific and commercial data available.'' The language provided by
Congress in the Act thus precludes such costs from consideration in
association with a listing determination. We work collaboratively with
private landowners, and strongly encourage those with listed species on
their property to work with us to develop incentive-based measures such
as strategic habitat areas (SHAs) and HCPs, which have the potential to
provide conservation measures that effect positive results for the
species and their habitat while providing regulatory relief for
landowners. The conservation and recovery of endangered and threatened
species, especially of those in Hawaii that occur nowhere else in the
world, and the ecosystems upon which they depend, is the ultimate
objective of the Act, and the Service recognizes the vital importance
of voluntary, nonregulatory conservation measures that provide
incentives for landowners in achieving that objective. In regards to
land management by native Hawaiians, see our response to Comment (9),
above. The Act does provide for the consideration of potential economic
impacts in the course of designating critical habitat (limited to
activities that are funded, authorized, or carried out by a Federal
agency), and that analysis will be conducted as we prepare a rule
proposing critical habitat for the multi-island species.
(14) Comment: Four commenters were concerned that listing a species
would entail removal of nonnative species with cultural significance,
or removal of those used for food and sport hunting, and that control
of nonnative ungulates would not be conducted humanely.
Our Response: Habitat destruction and modification by ungulates is
a threat to 37 of the 39 plants, and to 9 of the 10 animals proposed
for listing. Herbivory by ungulates is a threat to 27 of the 39 plants
proposed for listing. Hawaii was inhabited as early as the 2nd century;
therefore, hunting of game mammals is a relatively recent activity
(Tomich 1986, p. 1). The first Polynesian settlers brought domestic
pigs of southeast Asia (Sus scrofa or a species derived from Sus scrofa
vittatus) with them that were small in size, domesticated, and allowed
to run freely around habitations (Tomich 1986, p. 120). Cook brought
English pigs on his first voyage to Hawaii and landed a boar and sow on
Niihau in 1778 (Tomich 1986, p. 121). Goats and European boars were
introduced and released (on Niihau in 1778) by ship captains with the
intent of establishing feral populations of these animals to be an
available food source in future visits to the islands. Cattle (Bos
taurus) and domestic sheep (Ovis aries) were released in 1794, by
Vancouver. Deer were released later; first, axis deer in 1867, and then
mule deer (black-tailed deer) in 1961 (Tomich 1986, pp. 127, 133, 141,
150, 158). These ungulates multiplied rapidly, with immense negative
impacts to native vegetation (Loope 1988, pp. 274-276). The need for
control of feral cattle was recognized as early as 1918, by C.S. Judd
(Tomich 1986, p. 146). The commenter may be referring to the Federal
court order mandating the removal of sheep and goats for protection of
the palila (Loxioides bailleui), an endangered bird endemic to Hawaii.
Aerial hunting is an efficient control method and was chosen by the
State to comply with this order. Carcasses taken during hunts (in both
2014 and 2015) were available to the permitted public for salvage (DLNR
2014, in litt.; DLNR 2015, in litt.). Aerial hunting is not conducted
by the Service in Hawaii.
(15) Comment: One commenter stated that once species are listed for
protection under the Act, there is no public recourse.
Our Response: There is public recourse after a rulemaking is
published in the Federal Register. Under the Act,
[[Page 67790]]
an interested person may petition to add a species to, or to remove a
species from, either of the Lists of Endangered and Threatened Wildlife
and Plants. Within 12 months of the petition, the Secretary will make a
finding as to whether the petition presents substantial scientific or
commercial information indicating that the petitioned action may be
warranted. Persons may also petition to designate or revise a critical
habitat designation. Our petition regulations are set forth at 50 CFR
424.14.
(16) Comment: Two commenters expressed concern that the magnitude
of the proposed listing rule and the subsequent designation of critical
habitat will have negative effects on Hawaii's economy, property
values, and land use.
Our Response: We understand there is confusion and concern about
the effects of listing the 49 multi-island species. Listing provides
certain protections to the species under the Act. Section 7 of the Act
states that each Federal agency (through consultation) shall insure
that any action authorized, funded, or carried out by the agency is not
likely to jeopardize the continued existence of any endangered or
threatened species. For endangered species of fish or wildlife, section
9 of the Act prohibits any person subject to the jurisdiction of the
United States to import or export; ``take'' (defined as harass, harm,
pursue, hunt, shoot, wound, kill, trap, capture, collect, or attempt
any of these actions) within the United States or the territorial sea
of the United States; take upon the high seas; deliver, receive, carry,
transport, or ship in interstate or foreign commerce in the course of a
commercial activity; or sell or offer for sale in interstate or foreign
commerce. For endangered plants, section 9 of the Act prohibits any
person subject to the jurisdiction of the United States to import or
export; deliver, receive, carry, transport, or ship in interstate or
foreign commerce in the course of a commercial activity; sell or offer
for sale in interstate or foreign commerce; remove and reduce the
species to possession from areas under Federal jurisdiction;
maliciously damage or destroy any such species on areas under Federal
jurisdiction; or remove, cut, dig up, or damage or destroy any species
species in knowing violation of any State law or regulations or in the
course of any violation of a State criminal trespass law. Section 10 of
the Act provides for permitting of actions that may enhance the
propagation or survival of the species, or that may ``take'' a species.
We acknowledge that some economic impacts are a possible consequence of
listing a species under the Act; for example, there may be costs to the
landowner associated with the development of an HCP. In other cases, if
the landowner does not acquire a permit for incidental take, the
landowner may choose to forego certain activities on their property to
avoid violating the Act, resulting in potential lost income. However,
the statute does not provide for the consideration of such impacts when
making a listing decision. Listing determinations are made ``solely on
the basis of the best scientific and commercial data available.'' This
rule only lists the 49 species from the Hawaiian Islands; it does not
designate critical habitat.
(17) Comment: Two commenters stated that listing species and
designating critical habitat on private property in Hawaii will
alienate ranchers, a group that can help with species and habitat
conservation. The commenters state that conservation can best be
achieved by cooperation and coordination with private landowners.
Our Response: This rule only addresses the listing of 49 species
from the Hawaiian Islands and does not designate critical habitat. We
agree that partnerships can provide benefits for listed species and
their habitat through development of conservation plans and
implementation of management actions.
(18) Comment: One commenter stated that the Service should include
the public now, not after designating critical habitat, with outreach,
public forums, presentations, and meetings on every island for
community groups, industry and business groups, the Soil and Water
Conservation Districts, the Farm Bureau, Hawaii Cattlemen's Council,
and schools.
Our Response: As described above, the publication of the proposed
listing rule did not include a critical habitat proposal. We opened a
60-day comment period on the proposed listing rule, obtained extensive
peer review, published notices in numerous local newspapers, reopened
the comment period, and held a public hearing and information meeting.
We considered all comments we received in preparing this final listing
rule, and this rule incorporates new, substantive information provided
to us by commenters.
Summary of Changes From the Proposed Rule
In preparing this final rule, we reviewed and fully considered
comments from the public and peer reviewers on the proposed rule, and
incorporated the following substantive changes into this final rule.
None of the new information we received changed our evaluation of the
threats to these species or our determinations in this final rule that
they are endangered.
(1) We made revisions to the demographic status or distribution of
31 species of plants, based on comments from peer reviewers, by
correcting current locations or numbers of individuals for: Asplenium
diellaciniatum, Calamagrostis expansa, Cyanea kauaulaensis, Cyclosorus
boydiae, Cyrtandra hematos, Dryopteris glabra var. pusilla, Exocarpos
menziesii, Gardenia remyi, Huperzia stemmermanniae, Joinvillea
ascendens ssp. ascendens, Kadua fluviatilis, Microlepia strigosa var.
mauiensis, Myrsine fosbergii, Nothocestrum latifolium, Ochrosia
haleakalae, Phyllostegia brevidens, P. helleri, P. stachyoides,
Portulaca villosa, Pritchardia bakeri, Pseudognaphalium sandwicensium
var. molokaiense, Ranunculus hawaiensis, R. mauiensis, Sanicula
sandwicensis, Santalum involutum, Schiedea diffusa ssp. diffusa, S.
pubescens, Sicyos lanceoloideus, S. macrophyllus, Stenogyne kaalae ssp.
sherffii, and Wikstroemia skottsbergiana.
(2) We made revisions to specific threats to 31 plant species,
based on comments from peer reviewers, including: Asplenium
diellaciniatum, Calamagrostis expansa, Cyanea kauaulaensis, Cyclosorus
boydiae, Cyperus neokunthianus, Cyrtandra hematos, Deparia kaalaana,
Dryopteris glabra var. pusilla, Exocarpos menziesii, Huperzia
stemmermanniae, Hypolepis hawaiiensis var. mauiensis, Joinvillea
ascendens ssp. ascendens, Kadua fluviatilis, K. haupuensis, Labordia
lorenciana, Lepidium orbiculare, Microlepia strigosa var. mauiensis,
Myrsine fosbergii, Nothocestrum latifolium, Ochrosia haleakalae,
Phyllostegia brevidens, P. helleri, P. stachyoides, Portulaca villosa,
Sanicula sandwicensis, Santalum involutum, Schiedea diffusa ssp.
diffusa, S. pubescens, Sicyos lanceoloideus, Solanum nelsonii, and
Wikstroemia skottsbergiana.
(3) We corrected the taxonomy for the nonnative plant, California
grass, from Brachiaria mutica to Urochloa mutica.
(4) We added further references concerning genetic research that
supports differences in populations of the band-rumped storm-petrel
breeding in different oceans and archipelagos.
(5) We added additional information on current nesting sites of the
band-rumped storm-petrel on Lehua Island, Kauai, Molokai (coastal),
Lanai (coastal), Hawaii Island (Hawaii Volcanoes National Park), and
subalpine habitat
[[Page 67791]]
(Hawaii Island), based on comments regarding audio detections.
(6) We added information regarding additional populations of the
orangeblack Hawaiian damselfly on Hawaii Island.
(7) We added information on predation of the orangeblack Hawaiian
damselfly by Jackson's chameleons, backswimmers, and bullfrogs as a
threat, and predation by the black twig borer as a threat to Labordia
lorenciana and Nothocestrum latifolium.
(8) We added competition with caddisflies for resources, prey, and
space as a potential threat to the orangeblack Hawaiian damselfly.
(9) We made revisions to the demographic status or distribution of
the yellow-faced bees Hylaeus anthracinus, H. facilis, and H.
longiceps.
(10) We added tsunami as a threat to the yellow-faced bees that
occur in coastal areas (Hylaeus anthracinus, H. assimulans, H. facilis,
H. hilaris, and H. longiceps), and to Solanum nelsonii, also in coastal
areas.
(11) We changed ``Australian colletid'' to ``alien Hylaeus'' bees,
and included competition with sweat bees (Lasioglossum spp.) as a
threat to the yellow-faced bees.
(12) We noted that transmission of diseases carried by nonnative
insects through shared food sources could be a threat to the yellow-
faced bees, but we have no specific evidence of this type of disease
transmission.
(13) We added drought as a potential threat to all seven yellow-
faced bees.
(14) We added infiltration of waste water, fertilizers, or
pesticides resulting from development activities as a potential threat
to the anchialine pool shrimp.
(15) We added sea-level rise and coastal inundation as a potential
threat to Solanum nelsonii, as occurrences in low-lying coastal areas
are at risk, and to the anchialine pool shrimp, as these events could
increase connectivity of anchialine pools leading to further incursion
by nonnative fish from one pool to another.
Background
Please refer to the proposed listing rule for the 49 species from
the Hawaiian Islands (80 FR 58820; September 30, 2015), available at
https://www.regulations.gov (see ADDRESSES), for the following
information:
For background information on the Hawaii Islands, see
``The Hawaiian Islands'' under Background;
For ecosystem descriptions, see An Ecosystem-Based
Approach To Assessing the Conservation Status of the 49 Species in the
Hawaiian Islands;
For detailed descriptions of the species and their
taxonomy, see Description of the 49 Hawaiian Islands Species.
Hawaiian Islands Species Addressed in This Final Rule
Table 1A (plants) and Table 1B (animals), below, provide the common
name, scientific name, and range (by Hawaiian Island) for the 49
species addressed in this final rule.
Table 1A--Plant Species Listed as Endangered
------------------------------------------------------------------------
Scientific name Common name Hawaiian Island
------------------------------------------------------------------------
Plants
Asplenium diellaciniatum...... No common name Kauai.
(NCN).
Calamagrostis expansa......... Maui reedgrass... Hawaii, Maui.
Cyanea kauaulaensis........... NCN.............. Maui.
Cyclosorus boydiae............ kupukupu makalii. Hawaii (H), Maui,
Oahu.
Cyperus neokunthianus......... NCN.............. Maui (H).
Cyrtandra hematos............. haiwale.......... Molokai.
Deparia kaalaana.............. NCN.............. Hawaii (H), Maui,
Kauai (H).
Dryopteris glabra var. pusilla hohiu............ Kauai.
Exocarpos menziesii........... heau............. Hawaii, Lanai (H).
Festuca hawaiiensis........... NCN.............. Hawaii, Maui (H).
Gardenia remyi................ nanu............. Hawaii, Maui,
Molokai, Kauai.
Huperzia stemmermanniae....... NCN.............. Hawaii, Maui (H).
Hypolepis hawaiiensis var. olua............. Maui.
mauiensis.
Joinvillea ascendens ssp. ohe.............. Hawaii, Maui,
ascendens. Molokai, Oahu,
Kauai.
Kadua fluviatilis............. kamapuaa......... Oahu, Kauai.
Kadua haupuensis.............. NCN.............. Kauai (H).
Labordia lorenciana........... NCN.............. Kauai.
Lepidium orbiculare........... anaunau.......... Kauai.
Microlepia strigosa var. NCN.............. Hawaii, Maui, Oahu.
mauiensis.
Myrsine fosbergii............. kolea............ Oahu, Kauai.
Nothocestrum latifolium....... aiea............. Maui, Lanai (H),
Molokai, Oahu, Kauai
(H).
Ochrosia haleakalae........... holei............ Hawaii, Maui.
Phyllostegia brevidens........ NCN.............. Hawaii, Maui.
Phyllostegia helleri.......... NCN.............. Kauai.
Phyllostegia stachyoides...... NCN.............. Hawaii (H), Maui,
Molokai.
Portulaca villosa............. ihi.............. Hawaii, Maui,
Kahoolawe, Lanai
(H), Molokai, Oahu
(H), Kaula (H),
Lehua (H), Nihoa
(H).
Pritchardia bakeri............ Baker's loulu.... Oahu.
Pseudognaphalium sandwicensium enaena........... Maui, Lanai (H),
var. molokaiense. Molokai, Oahu (H).
Ranunculus hawaiensis......... makou............ Hawaii, Maui (H).
Ranunculus mauiensis.......... makou............ Hawaii (H), Maui,
Molokai (H), Oahu
(H), Kauai.
Sanicula sandwicensis......... NCN.............. Hawaii, Maui.
Santalum involutum............ iliahi........... Kauai.
Schiedea diffusa ssp. diffusa. NCN.............. Maui, Molokai (H).
Schiedea pubescens............ maolioli......... Maui, Lanai (H),
Molokai.
Sicyos lanceoloideus.......... anunu............ Oahu, Kauai.
Sicyos macrophyllus........... anunu............ Hawaii, Maui (H).
Solanum nelsonii.............. popolo........... Hawaii, Maui (H),
Molokai, Niihau (H),
Pearl & Hermes,
Kure, Midway,
Laysan, Nihoa.
Stenogyne kaalae ssp. sherffii NCN.............. Oahu (H).
[[Page 67792]]
Wikstroemia skottsbergiana.... akia............. Kauai.
------------------------------------------------------------------------
(H) = historically known from island, but not observed in the past 20
years.
Table 1B--Animal Species Listed as Endangered
------------------------------------------------------------------------
Common name Scientific name Hawaiian Island
------------------------------------------------------------------------
Animals
Band-rumped storm-petrel...... Oceanodroma Hawaii, Maui,
castro. Kahoolawe, Lanai,
Molokai (H), Oahu
(H), Kauai, Lehua.
Yellow-faced bee.............. Hylaeus Hawaii, Maui,
anthracinus. Kahoolawe, Lanai
(H), Molokai, Oahu.
Yellow-faced bee.............. Hylaeus Maui, Kahoolawe,
assimulans. Lanai, Oahu (H).
Yellow-faced bee.............. Hylaeus facilis.. Maui (H), Lanai (H),
Molokai, Oahu.
Yellow-faced bee.............. Hylaeus hilaris.. Maui (H), Lanai (H),
Molokai.
Yellow-faced bee.............. Hylaeus kuakea... Oahu.
Yellow-faced bee.............. Hylaeus longiceps Maui, Lanai, Molokai,
Oahu.
Yellow-faced bee.............. Hylaeus mana..... Oahu.
Orangeblack Hawaiian damselfly Megalagrion Hawaii, Maui, Lanai,
xanthomelas. Molokai, Oahu, Kauai
(H).
Anchialine pool shrimp........ Procaris hawaiana Hawaii, Maui.
------------------------------------------------------------------------
(H) = Historically known from the island, but not observed in the last
20 years.
Summary of Biological Status of the 49 Hawaiian Islands Species
The Act directs us to determine whether any species is an
endangered species or a threatened species because of any one of the
factors listed in section 4(a)(1). We summarize, below, the biological
condition of, and factors affecting, each of the 49 species and
determine whether each species is endangered or threatened. The
summaries below include only brief lists of factors affecting each
species. Each of these factors is fully considered, in detail, in the
subsequent section, Summary of Factors Affecting the 49 Species From
the Hawaiian Islands.
Climate Change Vulnerability Assessment for the Hawaiian Plants
Twenty-seven of the plant species described below were evaluated
for their vulnerability to climate change as part of a comprehensive
vulnerability analysis of native Hawaiian plants, as indicated in Table
2 (Fortini et al. 2013, 134 pp.). This analysis used ``climate
envelopes'' (geographic ranges encompassing suitable climate for each
species, as defined by temperature and moisture (Fortini et al. 2013,
p. 17)) developed from field records by Price et al. (2012) to project
each species' potential range in the year 2100. The location and
spatial extent of these future ranges, and their overlap with current
ranges, allows calculation of a vulnerability score. Estimates of
vulnerability based on climate-envelope modeling are conservative in
that they do not take into account potential changes in interspecific
interactions such as predation, disease, pollination, or competition.
This study provides a landscape- or island-scale picture of potential
climate-change vulnerability of Hawaiian plants; the results are less
clear at finer spatial scales (Fortini et al. 2013, p. 42). However,
all 27 of these plant species scored moderately or extremely vulnerable
in the analysis because of their relative inability to exhibit the
possible responses necessary for persistence under projected climate
change (Fortini et al. 2013, 134 pp.). These responses include the
migration response (dispersal and establishment in new areas beyond
their current distribution), the microrefugia response (persistence in
topographically complex areas that are less exposed), evolutionary
adaptation response (morphological changes in response to the changing
environment), and toleration response (adaptation to environmental
changes through phenotypic plasticity). In the study, response
probabilities ranged from 0 (not vulnerable at all) to 1.0 (extremely
vulnerable; species likely to disappear or ``wink out'' by the year
2100) (Fortini et al. 2013, pp. 6-7). Many species found to be
moderately vulnerable in this study, with scores of 0.5 or greater,
already are listed as endangered; some already are extinct (Fortini et
al. 2013, pp. 24, 93). Therefore, because the species in this rule were
found by the Fortini et al. (2013) study to be moderately (0.5) to
extremely (1.0) vulnerable, we deem the likelihood of their persistence
to be low with the impacts of climate change in addition to other
threats these species face. The environmental changes associated with
climate change are likely to exacerbate these ongoing threats and
further reduce the likelihood that these species will persist in the
future.
Plants
Asplenium diellaciniatum (no common name, NCN), a terrestrial or
epipetric (growing on rocks) fern in the spleenwort family
(Aspleniaceae), is endemic to Kauai (Palmer 2003, p. 117). Little is
known of the historical distribution of this species. It was described
from a collection from ``Halemanu,'' the Knudsen homestead area on
western Kauai. Currently, this fern is found in montane mesic forest at
Kawaiiki and Kaluahaulu Ridge (Palmer 2003, p. 117; HBMP 2010; Lorence
et al. 2013, p. 167) in 3 occurrences, totaling approximately 100
individuals, 30 of which are in an ungulate exclosure (TNCH 2007; HBMP
2010; Lorence et al. 2013, p. 167; Wood 2013, in litt.; Plant
Extinction Prevention Program (PEPP) 2014, pp. 33, 59; Kishida 2015, in
litt.; Williams 2015, in litt.).
Feral pigs (Sus scrofa), goats (Capra hircus), and black-tailed
deer (Odocoileus hemionus columbianus) modify and destroy the habitat
of Asplenium diellaciniatum on Kauai, with evidence of the activities
of these animals reported in the areas where A. diellaciniatum occurs
(Service 1999, p. 72; HBMP 2010). Feral pigs, goats, and black-tailed
deer also forage on A. diellaciniatum. Ungulates are managed in Hawaii
as game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt; Hawaii
Administrative
[[Page 67793]]
Rule-Hawaii Department of Land and Natural Resources (HAR-DLNR) 2010,
in litt.). Nonnative plants, such as Adiantum hispidulum (rough
maidenhair fern), Blechnum appendiculatum (no common name), Erigeron
karvinskianus (daisy fleabane), and Rubus argutus (prickly Florida
blackberry), compete with A. diellaciniatum, modify and destroy native
habitat, and displace native plant species by competing for water,
nutrients, light, and space; they may also produce chemicals that
inhibit growth of other plants (Smith 1985, pp. 180-250; Vitousek et
al. 1987 in Cuddihy and Stone 1990, p. 74; Williams 2015, in litt.).
Additionally, the small number of individuals of A. diellaciniatum
limits this species' ability to adapt to environmental change.
The remaining occurrences of Asplenium diellaciniatum are at risk;
A. diellaciniatum numbers are decreasing on Kauai, and both the species
and its habitat continue to be negatively affected by destruction and
modification by ungulates and by direct competition by nonnative
plants, combined with herbivory by nonnative ungulates. Because of the
threats described above, we find that this species is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Calamagrostis expansa (Maui reedgrass), a perennial in the grass
family (Poaceae), is known from the islands of Maui and Hawaii
(O'Connor 1999, p. 1509; Wagner and Herbst 2003, p. 59). Historically,
C. expansa was known from wet forest, open bogs, and bog margins on
Maui at 17 locations on east Maui, and in a large occurrence covering
nearly the entire summit on west Maui, and was discovered in 7
occurrences totaling approximately 750 individuals on the island of
Hawaii in 1995 (O'Connor 1999, p. 1509; HBMP 2010; Smithsonian National
Museum of Natural History (NMNH) Botany Collections 2014, in litt.;
Vetter 2015, in litt.). Currently, this species is known from 13 to 33
occurrences totaling fewer than 750 individuals. This species is
rhizomatous (growing from underground stems), making it difficult to
determine exact numbers of distinct individuals and populations, and
botanists' estimations vary. On the island of Maui, there are 2
occurrences in the west Maui Mountains (approximately 100 individuals)
and from 7 to as many as 40 occurrences in the east Maui Mountains
(totaling at least 200 individuals), often along ridges above 6,000
feet (ft) (1,830 meters (m)), or on raised hummocks in wet forest and
bogs, in the montane wet ecosystem (Wood 2005a, in litt.; TNCH 2007;
Welton 2008 and 2010, in litt.; Fay 2010, in litt.; HBMP 2010;
Oppenheimer 2010, in litt.; Agorastos 2011, in litt.; Vetter 2015, in
litt.). Most of the east Maui occurrences are in exclosures (Duvall
2015, in litt.). On the island of Hawaii, there are 3 occurrences in
the Kohala Mountains (totaling several hundred individuals) and 1
occurrence of 6 individuals last observed in 2004 in Upper Waiakea
Forest Reserve, in the montane wet ecosystem (Perry 2006, in litt; TNCH
2007; HBMP 2010; Perry 2015, in litt.).
Feral pigs modify and destroy the habitat of Calamagrostis expansa
on Maui and Hawaii, with evidence of the activities of feral pigs
reported in the areas where C. expansa occurs on east Maui, and on
Hawaii Island in the Kohala Mountains and in the Waiakea Forest Reserve
(Hobdy 1996, in litt.; Perlman 1996, in litt.; Wood 1996, in litt.;
Perry 2006, in litt.; HBMP 2010). Some occurrences on east and west
Maui are currently fenced; however, ungulate and weed control
activities must be maintained to provide continued protection (Duvall
2015, in litt.). Ungulates are managed in Hawaii as game animals, but
public hunting does not adequately control the numbers of ungulates to
eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.). Rats
have been noted by biologists as a threat to C. expansa at Laupahoehoe
Natural Area Reserve (NAR) on Hawaii Island, by consuming seeds (HBMP
2010). Nonnative plants compete with this species and modify and
destroy native habitat, negatively affecting C. expansa on east and
west Maui and Hawaii Island. Additionally, the small number of
individuals limits this species' ability to adapt to environmental
change. Fortini et al. (2013, p. 68) found that, as environmental
conditions are altered by climate change, C. expansa is unlikely to
tolerate or adapt to projected changes in temperature and moisture, and
is unlikely to be able to move to areas with more suitable climatic
conditions. Although we cannot predict the timing, extent, or magnitude
of specific impacts, we do expect the effects of climate change to
exacerbate the threats to C. expansa described above (see ``Climate
Change'' under Factor E. Other Natural or Manmade Factors Affecting
Their Continued Existence, below).
The remaining occurrences of Calamagrostis expansa are at risk; C.
expansa populations are decreasing on Maui and Hawaii Island, and this
species continues to be negatively affected by habitat modification and
destruction by feral pigs, and by direct competition from nonnative
plants, combined with herbivory by feral pigs and rats. This species is
vulnerable to the effects of climate change, and the likelihood of its
persistence with the impacts of climate change, exacerbated by the
ongoing threats, is low. We find that this species is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Cyanea kauaulaensis (NCN), a shrub in the bellflower family
(Campanulaceae), is endemic to Maui (Oppenheimer and Lorence 2012, p.
15). Cyanea kauaulaensis occurs on leeward west Maui, on talus or
basalt boulder-strewn slopes along perennial streams from 2,400 to
3,000 ft (730 to 900 m), in the lowland wet ecosystem (TNCH 2007; HBMP
2010; Oppenheimer and Lorence 2012, pp. 17-18). This species was first
collected during a botanical survey in 1989. Further surveys (in 2008,
2009, and 2011) revealed more individuals, and study of the collections
indicated that it was a new species of Cyanea. Currently, C.
kauaulaensis is known from Kauaula Valley (approximately 100
individuals) (Oppenheimer and Lorence 2012, pp. 15-16, 20; Duvall 2015,
in litt.; Oppenheimer 2015, in litt.).
The greatest threats to this species currently are the low numbers
of occurrences and individuals, its limited range, poor seedling
recruitment, and loss of pollinators and dispersal agents (Oppenheimer
and Lorence 2012, pp. 20-21; Duvall 2015, in litt.). Rats and slugs are
noted as a threat to Cyanea kauaulaensis because of their herbivory and
seed predation. Additionally, nonnative plants modify and destroy
native habitat and outcompete native species, negatively affecting C.
kauaulaensis and its habitat. Although feral ungulates are present on
west Maui, the known occurrences of C. kauaulaensis may be less at risk
from this particular threat because of their location in extremely
steep and rugged terrain; however, erosion, landslides, flooding, and
drying due to climate change affect this species because of the terrain
where it occurs (Oppenheimer and Lorence 2012, pp. 20-21; Duvall 2015,
in litt.). The remaining occurrence of Cyanea kauaulaensis is at risk.
Because of the threats described above, we find that this species is
endangered throughout all of its range, and,
[[Page 67794]]
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Cyclosorus boydiae (previously Christella boydiae) (kupukupu
makalii) is a small to medium-sized member of the thelypteroid fern
family (Thelypteridaceae) (Pukui and Elbert 1986, p. 186; Palmer 2003,
pp. 87-88). Typical habitat for C. boydiae is exposed, rocky, or moss-
covered banks of stream courses in dense-wet Metrosideros-Acacia (ohia-
koa) forest, from 2,300 to 4,400 ft (700 to 1,350 m), with other native
ferns, grasses, and dwarfed woody species, in the lowland wet and
montane wet ecosystems (Hillebrand 1888, p. 572; Medeiros et al. 1993,
p. 87; Wagner (W.H.) et al. 1999, p. 156; TNCH 2007; HBMP 2010; Gates
2015, in litt.). Historically, this fern was known from near sea level
to 4,400 ft (1,350 m) on Oahu, Maui, and Hawaii Island (Hillebrand
1888, p. 572; Medeiros et al. 1993, pp. 86-87; Palmer 2003, pp. 87-88).
Currently, C. boydiae is found on Oahu and east Maui, in 13 occurrences
totaling approximately 400 individuals (Palmer 2003, pp. 87-88;
Oppenheimer 2008, in litt.; Fay 2010, in litt.; HBMP 2010; Welton 2010,
in litt.). On east Maui, there are at least 11 occurrences (over 1,000
individuals) in the lowland wet and montane wet ecosystems, and on Oahu
there are 2 occurrences in the Koolau Mountains in the montane wet
ecosystem, totaling 40 individuals, and one historic occurrence in
Kaluanui Drainage, but the status of the species at this location is
currently unknown (Palmer 2003, pp. 87-88; Wood 2007a, in litt.; Kam
2008, in litt.; Oppenheimer 2008 and 2010, in litt.; HBMP 2010; Welton
2010, in litt.; Ching 2011, in litt.; Ching Harbin 2015, in litt.;
Oppenheimer 2015, in litt.). The historical occurrence of C. boydiae on
the island of Hawaii was found in the lowland wet ecosystem (HBMP
2010).
Feral pigs modify and destroy the habitat of Cyclosorus boydiae on
Maui and Oahu, with evidence of their activities reported at three
occurrences of C. boydiae on east Maui and at two occurrences on Oahu.
However, on east Maui, two of the five occurrences are provided
protection in Haleakala National Park (Wood 2007a, in litt.; HBMP 2010;
Kawelo 2011, in litt.). Ungulates are managed in Hawaii as game
animals, but public hunting does not adequately control the numbers of
ungulates to eliminate habitat modification and destruction or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Historical occurrences of C. boydiae on Oahu have
dramatically declined in numbers or disappeared as a result of habitat
modification and destruction, landslides and flooding, invasion of
lower elevation stream courses by nonnative plants, and manmade stream
diversions (Medeiros et al. 1993, p. 88; Palmer 2003, p. 88). Nonnative
plants, such as Tibouchina herbacea (glorybush), modify and destroy
native habitat of. C. boydiae and outcompete this and other native
species for water, nutrients, light, and space (Smith 1985, pp. 180-
250; Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74). Herbivory
by feral pigs negatively impacts this species (HBMP 2010). This species
occurs on stream banks at or just above water level, and flash floods
or drought can damage and destroy it (Ching Harbin 2015, in litt.).
Fortini et al. (2013, p. 72) found that, as environmental conditions
are altered by climate change, C. boydiae is unlikely to tolerate or
adapt to projected changes in temperature and moisture, and is unlikely
to be able to move to areas with more suitable climatic conditions.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to C. boydiae described above.
The remaining occurrences of Cyclosorus boydiae are at risk; C.
boydiae populations are decreasing on Oahu and Maui, and the species
continues to be negatively affected by habitat loss and destruction by
ungulates, direct competition with nonnative plants, and herbivory by
ungulates. Flash floods and drought can damage and destroy this
species. The effects of climate change are likely to further exacerbate
these threats. Because of the threats describe above, we find that this
species is endangered throughout all of its range, and, therefore, find
that it is unnecessary to analyze whether it is endangered or
threatened in a significant portion of its range.
Cyperus neokunthianus (NCN) is a perennial plant in the sedge
family (Cyperaceae) (Koyama 1999, p. 1420). Cyperus neokunthianus
occurs in riparian areas of the lowland wet ecosystem on west Maui
(Koyama 1999, p. 1420; TNCH 2007; HBMP 2010). Historically, this
species was known from Honokohau Falls and Waihee Valley (HBMP 2010;
Global Biodiversity Information Facility (GBIF) database 2014, in
litt.). This species was last observed in 1996. Currently, there are no
known individuals in the wild; however, Waihee Valley and Maui County
lands have been suggested as potential habitat for further surveys
(PEPP 2013, p. 32; PEPP 2014, p. 59; Duvall 2015, in litt.).
Feral pigs modify and destroy the habitat of Cyperus neokunthianus
on west Maui, with evidence of the activities of feral pigs reported in
the area where this species was last observed (HBMP 2010). Habitat
modifications resulting from activities of feral pigs that affect C.
neokunthianus include direct destruction of this species and other
native plants, disruption of topsoil leading to erosion, and
establishment and spread of nonnative plants. Ungulates are managed in
Hawaii as game animals, but public hunting does not adequately control
the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Additionally, nonnative plants modify
and destroy native habitat and outcompete native species, also
negatively affecting habitat of C. neokunthianus on west Maui.
Currently, there are no known extant individuals, and low numbers makes
this species more vulnerable to extinction because of the higher risks
from genetic bottlenecks, random demographic fluctuations, and
localized catastrophes.
Cyperus neokunthianus is at risk and continues to be negatively
affected by modification and destruction by nonnative animals and
plants (Duvall 2015, in litt.). Because of the threats described above,
we find that this species is endangered throughout all of its range,
and, therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Cyrtandra hematos (haiwale), a shrub in the African violet family
(Gesneriaceae), is endemic to Molokai (Wagner et al. 1999, pp. 760,
762). Cyrtandra hematos occurs in wet forest from 3,400 to 3,800 ft
(1,030 to 1,150 m) on eastern Molokai, in the montane wet ecosystem
(Wagner et al. 1999, pp. 760, 762; HBMP 2010; TNCH 2007). Historically,
this species was known from four locations on Molokai (Wagner et al.
1999, pp. 760, 762). Currently, there are fewer than 100 individuals at
two locations on Molokai (Duvall 2015, in litt.; Oppenheimer 2015, in
litt.).
Feral pigs and goats modify and destroy the habitat of Cyrtandra
hematos on Molokai, with evidence of the activities of these animals
reported in the areas where this species occurs (Service 2015, in
litt.). Ungulates are managed in Hawaii as game animals, but public
hunting does not adequately control the numbers of ungulates to
eliminate habitat modification and
[[Page 67795]]
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Additionally, nonnative plants modify
and destroy native habitat and outcompete this and other native species
for water, nutrients, light, and space, or a nonnative plant may
produce chemicals that inhibit growth of other plants (Smith 1985, pp.
180-250; Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74; Service
2015, in litt.). This species experiences reduced reproductive vigor
due to low numbers and lack of regeneration, leading to diminished
capacity to adapt to environmental changes, and thereby lessening the
probability of long-term persistence (Barrett and Kohn 1991, p. 4;
Newman and Pilson 1997, p. 361). This species hybridizes with C.
grayana (Oppenheimer 2015, in litt.). Fortini et al. (2013, p. 72)
found that, as environmental conditions are altered by climate change,
C. hematos is unlikely to tolerate or adapt to projected changes in
temperature and moisture, and is unlikely to be able to move to areas
with more suitable climatic conditions. Although we cannot predict the
timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to C. hematos
described above.
The remaining occurrences of Cyrtandra hematos are at risk. The
known individuals are restricted to a small area on Molokai and
continue to be negatively affected by habitat modification and
destruction by ungulates and nonnative plants, and by direct
competition with nonnative plants. The low number of remaining
individuals limits this species' ability to adapt to environmental
changes. Hybridization results in a reduction of the numbers of C.
hematos. The effects of climate change are likely to further exacerbate
these threats. We find that this species is endangered throughout all
of its range, and, therefore, find that it is unnecessary to analyze
whether it is endangered or threatened in a significant portion of its
range.
Deparia kaalaana (NCN), a small, terrestrial fern in the ladyfern
family (Athyriaceae), is recognized as a distinct taxon by Palmer
(2003, pp. 109-111) and Christenhusz et al. (2012, p. 16). This fern is
historically known from the islands of Kauai, Maui, and Hawaii, on
rocky stream banks and in wet forest, in the lowland mesic and lowland
wet ecosystems (Palmer 2003, pp. 109-111; TNCH 2007; HBMP 2010;
Oppenheimer and Bustamente 2014, p. 103; PEPP 2014, p. 95). Deparia
kaalaana was presumed extinct on all three islands where it previously
occurred until one individual was discovered on east Maui, growing
along a perennial stream on the western side of a small pool with other
native ferns and herbaceous plants (Oppenheimer and Bustamente 2014,
pp. 103-107; PEPP 2014, p. 95).
Feral pigs modify and destroy habitat of Deparia kaalaana by
facilitating the spread of nonnative plants, which converts vegetation
communities from native to nonnative (Cuddihy and Stone 1990, p. 63;
Oppenheimer and Bustamente 2014, p. 106). Ungulates are managed in
Hawaii as game animals, but public hunting does not adequately control
the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt; HAR-DLNR 2010, in litt.). Nonnative plants, such as Blechnum
appendiculatum (NCN), Clidemia hirta (Koster's curse), Hedychium
gardnerianum (kahili ginger), Prunella vulgaris (selfheal), and Rubus
argutus, are capable of displacing all of the riparian habitat
elements, including native plants, in the area where D. kaalaana
occurs. Nonnative slugs such as Derocerus laeve and Limax maximus are
common in the area and can consume young plants (Joe and Daehler 2008,
pp. 252-253). Flash floods and drought can damage and destroy this
species at its only known location. A single catastrophic event may
result in extirpation of the remaining individual.
The remaining occurrence of Deparia kaalaana is at risk, and both
the species and its habitat on Hawaii, Maui, and Kauai continues to be
negatively affected by modification and destruction by nonnative
ungulates, and by direct competition with nonnative plants, combined
with herbivory by nonnative ungulates and slugs. Although we cannot
predict the timing, extent, or magnitude of specific impacts, we do
expect the effects of climate change to exacerbate the threats to D.
kaalaana described above. We find that this species is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Dryopteris glabra var. pusilla (hohiu) is a small, terrestrial fern
in the wood fern family (Dryopteridaceae) (Palmer 2003, p. 144).
Habitat for D. glabra var. pusilla is deep shade on rocky, mossy
streambanks in wet forest at about 4,000 ft (1,200 m), in the montane
wet ecosystem on Kauai (Palmer 2003, p. 144; TNCH 2007; HBMP 2010).
Historically, D. glabra var. pusilla was known from the Kawaikoi stream
area (HBMP 2010). Currently, this variety is known from fewer than 250
individuals in the Alakai Wilderness Preserve on Kauai (National
Tropical Botanical Garden (NTBG) Herbarium Database 1995, in litt.;
HBMP 2010; Wood 2015, in litt.).
Dryopteris glabra var. pusilla is at risk from habitat modification
and destruction by nonnative plants, feral pigs, and black-tailed deer
(Wood 2015, in litt.). Most individuals occur in the Alakai Wilderness
Preserve; however, only portions of the Preserve are fenced to prevent
ungulate incursion. Ungulates are managed in Hawaii as game animals,
but public hunting does not adequately control the numbers of ungulates
to eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.).
Nonnative plants modify and destroy native habitat and outcompete this
and other native species for water, nutrients, light, and space, or a
nonnative plant may produce chemicals that inhibit growth of other
plants, also negatively affecting habitat of D. glabra var. pusilla
(Smith 1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy and Stone
1990, p. 74). Herbivory by rats and slugs is a threat to D. glabra var.
pusilla (Wood 2015, in litt.). In addition, the limited number of
occurrences and few individuals lead to a diminished capacity to adapt
to environmental changes, thereby lessening the probability of long-
term persistence, and a single catastrophic event may result in
extirpation of remaining occurrences. Landslides along streambanks have
been known to destroy populations of this fern (Wood 2015, in litt.).
Fortini et al. (2013, p. 74) found that, as environmental
conditions are altered by climate change, D. glabra var. pusilla is
unlikely to tolerate or adapt to projected changes in temperature and
moisture, and is unlikely to be able to move to areas with more
suitable climatic conditions. Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to D. glabra var. pusilla
described above. Because of these threats, we find that this variety is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Exocarpos menziesii (heau) is a shrub in the sandalwood family
(Santalaceae) (Wagner et al. 1999, p. 1218). This species occurs in
Metrosideros shrubland or drier forest areas, and on lava flows with
sparse vegetation, from 4,600 to 6,900 ft (1,400 to 2,100 m), in the
montane dry ecosystem on the island of Hawaii (Wagner et al. 1999, p.
[[Page 67796]]
1218; TNCH 2007), and historically occurred in the lowland mesic (Lanai
and Hawaii Island) and montane mesic ecosystems (Hawaii Island) (TNCH
2007; Bishop Museum 2014). Exocarpos menziesii is historically known
from the island of Lanai and was wide-spread on the island of Hawaii
(Wagner et al. 1999, p. 1218; TNCH 2007; Bishop Museum 2014).
Currently, there are seven scattered occurrences on Hawaii Island, six
of which consist of only a few individuals, the seventh totals an
estimated 1,800 individuals (PEPP 2013, pp. 10, 33; Thomas 2014, in
litt.; Evans 2015a, in litt.; Orlando 2015, in litt.; Perry 2015, in
litt.). There are no currently known occurrences of this species on
Lanai.
Feral goats, mouflon, and sheep modify and destroy the habitat of
Exocarpos menziesii on Hawaii Island, and may forage on this species,
with evidence of the activities of these animals reported in the areas
where this species occurs (Service 2015, in litt.). Ungulates are
managed in Hawaii as game animals, but public hunting does not
adequately control the numbers of ungulates to eliminate habitat
modification and destruction or herbivory by these animals (Anderson et
al. 2007, in litt; HAR-DLNR 2010, in litt.). Feral ungulate management
is incorporated into the U.S. Army's Pohakuloa Training Area (PTA)
management plan, and plants at PTA may be provided some protection
within fenced management units in the training area (Evans 2015a, in
litt.); however, it is reported that feral goats are still being
removed from within the fenced area (Nadig 2015, in litt.). Any
individuals of E. menziesii outside of fenced exclosures or outside of
the managed area are at risk. Additionally, nonnative plants modify and
destroy native habitat and outcompete this and other native species for
water, nutrients, light, and space, or a nonnative plant may produce
chemicals that inhibit growth of other plants, also negatively
affecting habitat of E. menziesii (Smith 1985, pp. 180-250; Vitousek et
al. 1987 in Cuddihy and Stone 1990, p. 74). Occurrences and numbers of
individuals have declined on the island of Hawaii (HBMP 2010; Thomas
2014, in litt.), where E. menziesii was once widely distributed from
the south to the west sides of the island, and are now restricted to
seven locations. Consequently, E. menziesii experiences reduced
reproductive vigor due to reduced levels of genetic variability,
leading to diminished capacity to adapt to environmental changes,
thereby reducing the probability of long-term persistence (Barrett and
Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). Fire is a likely
threat to this species; although the U.S. Army has constructed
firebreaks and has standard operating procedures (SOPs) in place for
prevention and suppression of wildfires at the PTA, wildfires may
encroach from other areas (U.S. Army Garrison 2013, in litt.). The
small number of individuals outside the larger occurrence at the PTA
limits this species' ability to adapt to environmental changes. Fortini
et al. (2013, p. 76) found that, as environmental conditions are
altered by climate change, E. menziesii is unlikely to tolerate or
adapt to projected changes in temperature and moisture, and is unlikely
to be able to move to areas with more suitable climatic conditions.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to E. menziesii described above.
The remaining occurrences of Exocarpos menziesii are at risk from
modification and destruction by feral goats, mouflon, and sheep; from
herbivory by these ungulates; and by the small number of remaining
occurrences. Fire is a likely threat to this species. The effects of
climate change are likely to exacerbate these threats. Because of these
threats, we find that this species is endangered throughout all of its
range, and, therefore, find that it is unnecessary to analyze whether
it is endangered or threatened in a significant portion of its range.
Festuca hawaiiensis (NCN) is a cespitose (growing in tufts or
clumps) annual in the grass family (Poaceae) (O'Connor 1999, p. 1547).
Typical habitat for this species is dry forest at 6,500 ft (2,000 m),
in the montane dry ecosystem (O'Connor 1999, p. 1547). Historically, F.
hawaiiensis occurred at Hualalai and Puu Huluhulu on the island of
Hawaii, and possibly at Ulupalakua on Maui; however, it is no longer
found at these sites (O'Connor 1999, p. 1547). Currently, F.
hawaiiensis is only known from the U.S. Army's PTA on the island of
Hawaii (HBMP 2010). These remaining four occurrences are within an area
of less than 10 square miles (mi) (26 square kilometers (km)) and total
approximately 1,500 individuals (U.S. Army Garrison 2013, in litt.;
Evans 2015a, in litt.).
Habitat destruction and modification by feral goats and sheep is a
threat to Festuca hawaiiensis. These ungulates also browse on native
plants such as grasses, including F. hawaiiensis. Ungulates are managed
in Hawaii as game animals, but public hunting does not adequately
control the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Feral ungulate management is
incorporated into the U.S. Army's PTA management plan, and these plants
are provided some protection within fenced management units in the
training area (Evans 2015a, in litt.); however, feral goats are still
being removed from inside the fenced area (Nadig 2015, in litt.). In
addition, any individuals of F. hawaiiensis outside of fenced
exclosures or outside of the managed area are at risk. Nonnative
plants, such as Cenchrus setaceus (Pennisetum setaceum; fountain
grass), are naturalized in the area and outcompete F. hawaiiensis and
other native plants. Occurrences and numbers of individuals are
declining on the island of Hawaii, and F. hawaiiensis experiences
reduced reproductive vigor due to reduced levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, thereby reducing the probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361; HBMP
2010). Fire is a likely threat to this species, especially because of
the ingress of nonnative grass species. Although the U.S. Army has
constructed firebreaks and has SOPs in place for prevention and
suppression of wildfires at the PTA, fires may encroach from other
areas, exacerbated by fuel loads provided by nonnative grasses (U.S.
Army Garrison 2013, in litt.). Fortini et al. (2013, p. 76) found that,
as environmental conditions are altered by climate change, F.
hawaiiensis is unlikely to tolerate or adapt to projected changes in
temperature and moisture, and is unlikely to be able to move to areas
with more suitable climatic conditions. Although we cannot predict the
timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to F. hawaiiensis
described above.
The remaining occurrences of Festuca hawaiiensis are at risk; F.
hawaiiensis occurrences have decreased on Hawaii Island, as it no
longer occurs at Hualalai and Puu Huluhulu, and the species may be
extirpated from Maui. This species continues to be negatively affected
by habitat modification and destruction by ungulates and by direct
competition with nonnative plants, combined with herbivory by
ungulates. Fire is a likely threat to the species and its habitat. The
effects of climate change are likely to further exacerbate these
threats. Because
[[Page 67797]]
of the threats described above, we find that this species is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Gardenia remyi (nanu) is a tree in the coffee family (Rubiaceae)
(Wagner et al. 1999, p. 1133). Typical habitat for G. remyi is mesic to
wet forest from 190 to 3,000 ft (60 to 760 m), in the lowland mesic
(Kauai, Molokai, and Hawaii Island) and lowland wet ecosystems (Kauai,
Molokai, Maui, and Hawaii Island) (Wagner et al. 1999, p. 1133; TNCH
2007; HBMP 2010; Oppenheimer 2015, in litt.). Historically, this
species was found on the island of Hawaii at Wao Kele O Puna NAR,
Waiakea Forest Reserve, Pahoa, and Hakalau Nui. On Maui, this species
was known from Wailuaiki and Waikamoi in the Koolau Forest Reserve, and
from Papaaea and Kipahulu. On Molokai, this species was known from
Keopukaloa, Pukoo, Honomuni, Halawa, and Kaluaaha (HBMP 2010). On
Kauai, this species ranged across the island, and was known from
Halelea, Kealia, Moloaa, and Lihue-Koloa Forest Reserves, including
Hanakapiai Valley, Mahaulepu, and east Wahiawa Bog. Currently, G. remyi
is known from 16 occurrences totaling approximately 90 individuals on
the islands of Hawaii, Maui, Molokai, and Kauai (Wood 2005b, in litt.;
Oppenheimer 2006, in litt; Perry 2006, in litt.; Welton 2008, in litt.;
Agorastos 2010, in litt.; HBMP 2010; Perlman 2010, in litt.). An
occurrence on east Maui has been observed to decline from 14
individuals in 1992, to only 1 individual by 2015 (Duvall 2015, in
litt.).
Habitat modification and destruction by feral pigs, goats, and axis
deer negatively affects Gardenia remyi and areas suitable for its
reintroduction (Perry, in litt. 2006; PEPP 2008, p. 102; HBMP 2010).
Feral pigs and signs of their activities have been reported at
occurrences of G. remyi on the island of Hawaii, on Kauai, on east and
west Maui, and on Molokai. Goats and signs of their activities are
reported at the occurrences G. remyi on Kauai and Molokai. Axis deer
and signs of their activities are reported at the occurrences of G.
remyi on Molokai (HBMP 2010). Herbivory by these ungulates is a threat
to G. remyi, as they browse on leaves and other parts of almost any
woody or fleshy plant species. Ungulates are managed in Hawaii as game
animals, but public hunting does not adequately control the numbers of
ungulates to eliminate habitat modification and destruction or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants, such as Clidemia hirta, Hedychium
gardnerianum, Psidium cattleianum (strawberry guava), and Tibouchina
herbacea on Hawaii Island (Perry 2006, in litt.); Lantana camara
(lantana), Psidium guajava, and Rubus argutus on Kauai (Wood 2004, in
litt.); Ageratina adenophora (Maui pamakani), Rubus rosifolius
(thimbleberry), and T. herbacea on Maui (HBMP 2010); and C. hirta and
P. cattleianum on Molokai (HBMP 2010), modify and destroy native
habitat of G. remyi and outcompete this and other native plants for
water, nutrients, light, and space in areas where G. remyi occurs on
these islands. Landslides are a threat to occurrences and habitat of G.
remyi on Hawaii Island (Perry 2006, in litt.). Lack of pollination was
suggested as the cause for abortion of immature fruits that were seen
among plants on Hawaii Island (PEPP 2010, p. 73). Similarly, Agorastos
(2011, in litt.) reported no viable seed production in the wild or
within ex situ collections and no recruitment in the wild among the 14
individuals observed on the island of Hawaii, Maui, and Molokai, for
unknown reasons (Duvall 2015, in litt.; Oppenheimer 2015, in litt.).
Some species of Gardenia are dioecious (male and female flowers on
separate plants) and although the breeding system of G. remyi is
currently unknown, this may be a cause of failure to produce viable
seed in isolated individuals (Lorence 2015, in litt.). Predation of
seeds by rats is reported as a threat to individuals on Kauai (NTBG
2008, in litt.). Fortini et al. (2013, p. 76) found that, as
environmental conditions are altered by climate change, G. remyi is
unlikely to tolerate or adapt to projected changes in temperature and
moisture, and is unlikely to be able to move to areas with more
suitable climatic conditions. Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to G. remyi described above.
The remaining occurrences of Gardenia remyi are at risk. Gardenia
remyi continues to be negatively affected by habitat modification and
destruction by ungulates, and by direct competition from nonnative
plants, combined with herbivory by ungulates and seed predation by
rats. Natural events such as landslides are a threat to occurrences on
the island of Hawaii. Pollination and seed production are observed to
be limited. Low numbers of individuals (90 total individuals
distributed across 4 islands) makes this species more vulnerable to
extinction because of the higher risks from genetic bottlenecks, random
demographic fluctuations, and localized catastrophes. The effects of
climate change are likely to exacerbate these threats. Because of the
threats, we find that this species is endangered throughout all of its
range, and, therefore, find that it is unnecessary to analyze whether
it is endangered or threatened in a significant portion of its range.
Huperzia stemmermanniae (NCN) is an epiphytic hanging fir-moss (a
fern ally) in the club moss family (Lycopodiaceae) (Palmer 2003, pp.
257-259). This species is epiphytic on rough bark of living trees or
fallen logs in Metrosideros polymorpha-Acacia koa forest on the island
of Hawaii, from 3,200 to 3,800 ft (975 to 1,160 m), in the montane wet
ecosystem (Medeiros et al. 1996b, p. 93; Palmer 2003, pp. 257, 259;
TNCH 2007; HBMP 2010). There is little information available on the
historical range of this species. Huperzia stemmermanniae was first
collected in 1981, from two occurrences totaling 10 individuals in
Laupahoehoe NAR on the island of Hawaii, and was mistakenly identified
as H. mannii (Medeiros et al. 1996b, p. 93; HBMP 2010). One individual
occurred in Kaapahu Valley on east Maui, but has not been relocated
since 1995 (Perry 2006, in litt.; Welton 2008, in litt.; HBMP 2010;
Conry 2012, in litt.; Perry 2015, in litt.). In 2006, there were
estimated to be as many as 20 individuals in Laupahoehoe (Perry 2006,
in litt.). Currently, there are only a few individuals remaining due to
prolonged drought conditions (Perry 2015, in litt.).
Feral pigs, goats, axis deer, and cattle modify and destroy the
habitat of Huperzia stemmermanniae on Maui, and feral pigs modify and
destroy the habitat of this species on Hawaii Island (Medeiros et al.
1996b, p. 96; Wood 2003, in litt.; HBMP 2010). Herbivory by these
ungulates is a threat to H. stemmermanniae. Ungulates are managed in
Hawaii as game animals, but public hunting does not adequately control
the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Nonnative plants, such as Clidemia
hirta, Miconia calvescens, Psidium cattleianum, and Cyathea cooperi
(Australian tree fern), modify and destroy the forest habitat that
supports the native species upon which this epiphytic plant grows, and
drought also negatively affects this species and its habitat (Medeiros
et al. 1996b, p. 96; Perry 2006, in litt.; HBMP 2010). Huperzia
stemmermanniae
[[Page 67798]]
experiences reduced reproductive vigor due to reduced levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, thereby lessening the probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361; HBMP
2010). Fortini et al. (2013, p. 77) found that, as environmental
conditions are altered by climate change, H. stemmermanniae is unlikely
to tolerate or adapt to projected changes in temperature and moisture,
and is unlikely to be able to move to areas with more suitable climatic
conditions. Although we cannot predict the timing, extent, or magnitude
of specific impacts, we do expect the effects of climate change to
exacerbate the threats to H. stemmermanniae described above.
The remaining occurrences of Huperzia stemmermanniae are at risk.
The known individuals are restricted to a small area on Hawaii Island,
and this species continues to be negatively affected by habitat
modification and destruction by ungulates. The low numbers of
individuals H. stemmermanniae reduces the probability of its long-term
persistence. The effects of climate change are likely to further
exacerbate these threats. Because of the threats described above, we
find that this species is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Hypolepis hawaiiensis var. mauiensis (olua) is a small terrestrial
member of the bracken fern family (Dennstaedtiaceae), and is recognized
as a distinct taxon by Palmer (2003, pp. 168-169). Hypolepis
hawaiiensis var. mauiensis occurs in wet forest, predominately in the
montane wet ecosystem (Palmer 2003, pp. 168-170; Oppenheimer 2015, in
litt.). This variety is historically known from west Maui (Palmer 2003,
pp. 168-170). Currently, 5 to 10 individuals are known from openings
between bogs on west Maui, and a few individuals are known from east
Maui (Maui Nui Task Force (MNTF) 2010, in litt.).
Nonnative plants, such as Tibouchina herbacea, modify and destroy
the habitat of Hypolepis hawaiiensis var. mauiensis on east and west
Maui (HBMP 2010; MNTF 2010, in litt.). Nonnative plants also displace
this and other native plant species by competing for water, nutrients,
light, and space, or they may produce chemicals that inhibit growth of
other plants (Smith 1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy
and Stones 1990, p. 74; MNTF 2010, in litt.). Herbivory by slugs is a
threat (Oppenheimer 2015, in litt.). This fern experiences reduced
reproductive vigor due to low numbers of individuals, leading to
diminished capacity to adapt to environmental changes, and thereby
lessening the probability of long-term persistence (Barrett and Kohn
1991, p. 4; Newman and Pilson 1997, p. 361). Fortini et al. (2013, p.
78) found that, as environmental conditions are altered by climate
change, H. hawaiiensis var. mauiensis is unlikely to tolerate or adapt
to projected changes in temperature and moisture, and is unlikely to be
able to move to areas with more suitable climatic conditions. Although
we cannot predict the timing, extent, or magnitude of specific impacts,
we do expect the effects of climate change to exacerbate the threats to
H. hawaiiensis var. mauiensis described above.
The remaining occurrences of Hypolepis hawaiiensis var. mauiensis
are at risk. Nonnative plants modify and destroy native habitat, and
also outcompete native plants, and this plant is threatened by
herbivory by slugs. This fern is also vulnerable to the impacts of
climate change, and the small number of remaining individuals limits
its ability to adapt to environmental change. Because of these threats,
we find that this variety is endangered throughout all of its range,
and, therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Joinvillea ascendens ssp. ascendens (ohe) is an erect perennial
herb in the joinvillea family (Joinvilleaceae) (Wagner et al. 1999, p.
1450). Joinvillea ascendens ssp. ascendens occurs in wet to mesic
Metrosideros polymorpha-Acacia koa lowland and montane forest, and
along intermittent streams, from 1,000 to 4,300 ft (305 to 1,300 m); in
the lowland mesic (Kauai and Oahu), lowland wet (Oahu, Molokai, Maui,
and Hawaii Island), montane wet (Kauai, Oahu, Molokai, Maui, and Hawaii
Island), and montane mesic ecosystems (Kauai) (TNCH 2007; HBMP 2010).
Historically, this subspecies was found in widely distributed
occurrences on the islands of Kauai, Oahu, Molokai, Maui, and Hawaii
Island (HBMP 2010). On Kauai, this subspecies was wide-ranging across
the mountains and into coastal areas (HBMP 2010). On Oahu, this
subspecies was known from the summit area of the Waianae Mountains, and
ranged along the entire length of the Koolau Mountain range. On
Molokai, this subspecies was known from the eastern half of the island
ranging from Pelekunu Preserve and east to Halawa Valley. On west Maui,
this subspecies occurred in the summit area, and on the northeastern
side of east Maui it ranged from the Koolau FR to Kaapahu (Gates 2015,
in litt.). On Hawaii Island, it occurred almost island-wide. Currently,
J. ascendens ssp. ascendens is still found on the same islands, in only
56 occurrences totaling approximately 200 individuals (HBMP 2010; Conry
2012, in litt.).
Nonnative ungulates modify and destroy habitat on all of the
islands where Joinvillea ascendens ssp. ascendens occurs (Moses 2006,
in litt.; Oppenheimer 2006, in litt.; Welton and Haus 2008, p. 16; HBMP
2010; Perlman 2010, in litt.). Herbivory by feral pigs, goats, axis
deer, black-tailed deer, and rats is a threat to this subspecies (HBMP
2010; Williams 2015, in litt.). Ungulates are managed in Hawaii as game
animals, but public hunting does not adequately control the numbers of
ungulates to eliminate habitat modification and destruction or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Many nonnative plant species, such as Passiflora
tarminiana (banana poka), Rubus ellipticus (yellow Himalayan
raspberry), and Setaria palmifolia (palmgrass) on Hawaii Island;
Clidemia hirta, Psidium cattleianum, and P. guajava on Kauai; C. hirta
and Tibouchina herbacea on Maui; Juncus effusus (Japanese mat rush) on
Molokai; and C. hirta and P. cattleianum on Oahu, modify and destroy
habitat and outcompete this subspecies (HBMP 2010). Randomly occurring
natural events, such as landslides, are a threat to the occurrences of
J. ascendens ssp. ascendens on Kauai and Molokai (HBMP 2010). Fire is
likely to be a threat to this subspecies in the drier areas of the
Waianae Mountains of Oahu (HBMP 2010). This subspecies is usually found
as widely separated individuals. Seedlings have rarely been observed in
the wild, and, although mature seeds germinate in cultivation, these
seedlings also rarely survive to maturity. It is uncertain if this
rarity of reproduction is typical, or if it is related to habitat
disturbance, or possibly a lack of soil mycorrhizae (symbiotic
relationship between fungi and plants) required for successful
establishment (Wagner et al. 1999, p. 1451; Oppenheimer 2015, in
litt.). Fortini et al. (2013, p. 76) found that, as environmental
conditions are altered by climate change, J. ascendens ssp. ascendens
is unlikely to tolerate or adapt to projected changes in temperature
and moisture, and is unlikely to be able to move to areas with
[[Page 67799]]
more suitable climatic conditions. Although we cannot predict the
timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to J. ascendens
ssp. ascendens described above.
The remaining occurrences of Joinvillea ascendens ssp. ascendens
are at risk. The known individuals continue to be negatively affected
by habitat modification and destruction by ungulates, compounded with
herbivory by ungulates and rats. The small number of remaining
individuals, smaller distribution, and poor recruitment in the wild
limits this subspecies' ability to adapt to environmental changes.
Destruction by fire, landslides, rockfalls, and floods can occur at any
time. The effects of climate change are likely to further exacerbate
these threats. Because of these threats, we find that this subspecies
is endangered throughout all of its range, and, therefore, find that it
is unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Kadua fluviatilis (previously Hedyotis fluviatilis) (kamapuaa) is a
climbing shrub in the coffee family (Rubiaceae) family (Wagner et al.
1999, pp. 1142-1144). Typical habitat for this species on Kauai is
mixed native shrubland and Metrosideros forest from 750 to 2,200 ft
(230 to 680 m), in the lowland mesic ecosystem (TNCH 2007; HBMP 2010);
and in open shrubland with sparse tree cover in the lowland mesic
ecosystem (Wood 1998, in litt.; TNCH 2007). On Oahu, K. fluviatilis
occurs along rocky streambanks in wet Metrosideros forest from 820 to
1,990 ft (250 to 607 m) in the lowland wet ecosystem (TNCH 2007; HBMP
2010).
Historically, Kadua fluviatilis was known from the island of Kauai
in at least 5 occurrences ranging from the north coast across the
central plateau to the south coast, and from the island of Oahu in at
least 11 occurrences in the northern Koolau Mountains (HBMP 2010;
Williams 2015, in litt.). Currently, during surveys on Oahu in 2013,
only 20 to 25 individuals were observed in one occurrence (Wood 2005b,
in litt., NTBG 2009, in litt.; HBMP 2010; Ching Harbin 2015, in litt.).
On Kauai, K. fluviatilis is known from two occurrences totaling
approximately 500 individuals (HBMP 2010).
Feral pigs and goats modify and destroy habitat of Kadua
fluviatilis (HBMP 2010). Evidence of the activities of feral pigs has
been reported at the occurrences on Kauai and Oahu (Wood 1998, in
litt.; HBMP 2010). Feral goats and evidence of their activities have
been observed at one location on Kauai (HBMP 2010). Herbivory by feral
pigs and goats is a threat to K. fluviatilis. Ungulates are managed in
Hawaii as game animals, but public hunting does not adequately control
the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Nonnative plant species, such as
Lantana camara, Paspalum conjugatum (Hilo grass), Psidium cattleianum,
P. guajava, Rubus rosifolius, and Schinus terebinthifolius (Christmas
berry), degrade habitat and outcompete this and other native species
for water, nutrients, light, and space, or may produce chemicals that
inhibit growth of other plants (Smith 1985, pp. 180-250; Vitousek et
al. 1987 in Cuddihy and Stone 1990, p. 74; Wood 1998, in litt.; HBMP
2010). Kadua fluviatilis is negatively affected by landslides on Kauai
(HBMP 2010). Fortini et al. (2013, p. 78) found that, as environmental
conditions are altered by climate change, K. fluviatilis is unlikely to
tolerate or adapt to projected changes in temperature and moisture, and
is unlikely to be able to move to areas with more suitable climatic
conditions. Although we cannot predict the timing, extent, or magnitude
of specific impacts, we do expect the effects of climate change to
exacerbate the threats to K. fluviatilis described above.
The remaining occurrences of Kadua fluviatilis are at risk. Numbers
of occurrences and individuals are decreasing on Oahu and Kauai, from
16 occurrences to 3, and from over 1,000 individuals to about 500
individuals (HBMP 2010; OTFM 2014, in litt.). This species continues to
be negatively affected by habitat modification and destruction by feral
pigs and goats, stochastic events such as landslides, and direct
competition from nonnative plants, combined with herbivory by nonnative
ungulates. Climate change is likely to further exacerbate these
threats. Because of these threats, we find that this species is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Kadua haupuensis (NCN) is a shrub in the coffee family (Rubiaceae)
(Lorence et al. 2010, p. 137). There is no historical information for
this species as it was recently discovered and described from one
occurrence just below and along cliffs in an isolated area on southern
Kauai, from 980 to 1,640 ft (300 to 500 m), in the lowland mesic
ecosystem (TNCH 2007; Lorence et al. 2010, pp. 137-144). Currently,
however, there are no known extant individuals of K. haupuensis; the
single natural occurrence is thought to be extirpated. Ten individuals
were propagated from seed collected in 1999, with cuttings from these
currently under cultivation. Seeds are in storage at NTBG's seed bank
(Lorence 2015, in litt.).
Feral pigs modify and destroy the habitat of Kadua haupuensis on
Kauai (Lorence et al. 2010, p. 140). Ungulates are managed in Hawaii as
game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Predation of fruits and seeds by rats is a threat.
Landslides are an additional threat to this species at its last known
occurrence. Nonnative plants, such as Caesalpinia decapetala (wait-a-
bit), Passiflora laurifolia (yellow granadilla), and various nonnative
grasses, modify and destroy native habitat, outcompete native plants,
and are found at the last known location of K. haupuensis. The small
number of remaining individuals limits this species' ability to adapt
to environmental change. Because of these threats, we find that K.
haupuensis is endangered throughout all of its range, and, therefore,
find that it is unnecessary to analyze whether it is endangered or
threatened in a significant portion of its range.
Labordia lorenciana (NCN) is a small tree in the Logania family
(Loganiaceae) (Wood et al. 2007, pp. 195-197). This species occurs on
the island of Kauai at 3,800 ft (1,160 m), in forest in the montane
mesic ecosystem (Wood et al. 2007, pp. 197-198). Currently, there are
four known individuals. Additional surveys for L. lorenciana have not
been successful; however, experts believe this species may occur in
other areas (Wood et al. 2007, p. 198).
Labordia lorenciana is at risk from habitat modification and
destruction and herbivory by nonnative mammals, displacement of
individuals through competition with nonnative plants, stochastic
events, and problems associated with small populations. Feral pigs,
goats, and black-tailed deer modify and destroy the habitat of L.
lorenciana (Wood et al. 2007, p. 198; Kishida 2015, in litt.).
Ungulates are managed in Hawaii as game animals, but public hunting
does not adequately control the numbers of ungulates to eliminate
habitat modification and destruction by these animals (Anderson et al.
2007, in litt; HAR-DLNR 2010, in litt.). Predation of seeds by rats is
a threat to this species (Wood et al. 2007, p. 198). Habitat
[[Page 67800]]
destruction and modification by nonnative plants, and competition with
nonnative plants including Lantana camara, Passiflora tarminiana,
Psidium cattleianum, and Rubus argutus, are a threat to Labordia
lorenciana, as these nonnative plants have the ability to spread
rapidly and cover large areas in the forest understory (Smith 1985, pp.
180-250; Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74; Wood et
al. 2007, p. 198). Randomly occurring natural events, such as
landslides, flash floods, fallen tree limbs, and fire, are threats to
L. lorenciana where it occurs on Kauai (Wood et al. 2007, p. 198). This
species experiences reduced reproductive vigor as there is no in situ
seedling recruitment and a very small number of individuals remain
(Wood et al. 2007, p. 198). Infestation by the black twig borer
(Xylosandrus compactus) is a threat to this species (Kishida 2015, in
litt.). Because of these threats, we find that L. lorenciana is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Lepidium orbiculare (anaunau) is a small, many-branched shrub in
the mustard family (Brassicaceae) (St. John 1981, pp. 371-373; Wagner
et al. 1999, p. 409). This species occurs in mesic forest on the island
of Kauai, in the lowland mesic ecosystem (Wagner et al. 1999, p. 409;
TNCH 2007; HBMP 2010; PEPP 2014, p. 34). Historically, this species was
known from widely scattered occurrences on Kauai (Wagner et al. 1999,
p. 409). Currently, there is one occurrence of fewer than 50
individuals (Wagner et al. 2012, p. 19; PEPP 2014, p. 34; Smithsonian
Institution 2015, in litt.).
Feral pigs and goats have been documented to modify and destroy
habitat of other rare and endangered native plant species at the same
location on Kauai (Lorence et al. 2010, p. 140; Kishida 2015, in
litt.); therefore, we consider that activities of feral pigs and goats
also pose a threat to Lepidium orbiculare. Ungulates are managed in
Hawaii as game animals, but public hunting does not adequately control
the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Nonnative plants, such as Melinis
minutiflora (molasses grass) and Stachytarpheta jamaicensis (Jamaica
vervain), degrade native habitat, outcompete native plants, and are
found at the last known location of L. orbiculare (HBMP 2010).
Landslides are an additional threat to this species. Because there are
fewer than 50 individuals, L. orbiculare experiences reduced
reproductive vigor due to reduced levels of genetic variability,
leading to diminished capacity to adapt to environmental changes, and
thereby lessening the probability of long-term persistence (Barrett and
Kohn 1991, p. 4; Newman and Pilson 1997, p. 361; PEPP 2014, p. 34).
The remaining occurrence of Lepidium orbiculare is at risk and the
species continues to be negatively affected by the threats described
above. Because of these threats, we find that this species is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Microlepia strigosa var. mauiensis (NCN) is a terrestrial, medium-
sized fern in the bracken fern family (Dennstaedtiaceae) (Palmer 2003,
p. 186). Typical habitat for M. strigosa var. mauiensis is mesic to wet
forest from 1,400 to 6,000 ft (425 to 1,830 m), in the lowland mesic
(Oahu), montane mesic (Hawaii Island), and montane wet (Maui and Hawaii
Island) ecosystems (Palmer 2003, p. 186; TNCH 2007: HBMP 2010). Little
is known of the historical locations of M. strigosa var. mauiensis;
however, it was wide-ranging on the islands of Hawaii, Maui, and Oahu
(HBMP 2010). Currently, M. strigosa var. mauiensis is known from nine
occurrences totaling fewer than 100 individuals on the islands of Oahu
(about 40 individuals), Maui (fewer than 20 individuals on east and
west Maui), and Hawaii (35 individuals last observed in 2004) (Palmer
2003, p. 186; Lau 2007, pers.comm.; Oppenheimer 2007 and 2008, in
litt.; Welton 2008, in litt.; Ching 2011, in litt.; Ching Harbin 2015,
in litt.; Oppenheimer 2015, in litt.).
Habitat modification and destruction by feral pigs and goats is a
threat to Microlepia strigosa var. mauiensis (Oppenheimer 2007, in
litt.; Bily 2009, in litt.; HBMP 2010). Herbivory by feral pigs is a
threat to M. strigosa var. mauiensis (Oppenheimer 2007, in litt.; Bily
2009, in litt.; HBMP 2010). Ungulates are managed in Hawaii as game
animals, but public hunting does not adequately control the numbers of
ungulates to eliminate habitat modification and destruction or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants, such as Ageratina adenophora, Juncus
acuminatus (rush), Plantago major (broad-leaved plantain), and
Tibouchina herbacea, degrade habitat and outcompete this variety on
Maui (Oppenheimer 2007, in litt.). Hybridization with other species and
varieties of Microlepia is a threat to this plant on Oahu and is
compounded by the low number of individuals (Kawelo 2010, in litt.).
Because of these threats, we find that this variety is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Myrsine fosbergii (kolea) is a branched shrub or small tree in the
myrsine family (Myrsinaceae) (Wagner et al. 1999, p. 940). Typical
habitat for this species on Oahu is Metrosideros-mixed native
shrubland, from 2,200 to 2,800 ft (670 to 850 m) (Wagner et al. 1999,
p. 940; TNCH 2007; HBMP 2010). Typical habitat for this species on
Kauai is Metrosideros-Diospyros (ohia-lama) lowland mesic forest and
Metrosideros-Cheirodendron (ohia-olapa) montane wet forest, often on
watercourses or stream banks, from 900 to 4,300 ft (270 to 1,300 m), in
the lowland mesic, lowland wet, and montane wet ecosystems (TNCH 2007;
HBMP 2010; Wagner et al. 2012, p. 53). Myrsine fosbergii was
historically known from the summit ridges of the Koolau Mountains of
Oahu (HBMP 2010). This species was first collected on Kauai in 1987.
Currently, on Oahu, there are fewer than 30 individuals in the Koolau
Mountains (lowland mesic and lowland wet ecosystems) (HBMP 2010; OTFM
2014, in litt.; Reynolds 2015, in litt.; Sailer 2015, in litt.).
Propagation attempts of the Oahu plants have been unsuccessful (Ching
Harbin 2015, in litt.). On Kauai, this species was once widely
scattered in the northwest and central areas, but is currently known
from only 55 remaining individuals (Wood 2005e and 2007c, in litt.;
HBMP 2010).
Myrsine fosbergii is at risk from habitat modification and
destruction by nonnative ungulates and plants. On Oahu, evidence of the
activities of feral pigs has been reported at all summit occurrences
(HBMP 2010). On Kauai, evidence of the activities of feral pigs has
been reported at the remaining occurrence (Wood 2005e and 2007c, in
litt.; HBMP 2010), and evidence of the activities of feral goats has
also been reported (HBMP 2010). Herbivory by feral pigs and goats is a
threat to M. fosbergii (Wood 2005e and 2007c, in litt.; HBMP 2010).
Ungulates are managed in Hawaii as game animals, but public hunting
does not adequately control the numbers of ungulates to eliminate
habitat modification and destruction or herbivory by these animals
(Anderson et al. 2007, in litt.;
[[Page 67801]]
HAR-DLNR 2010, in litt.). Nonnative plants, such as Axonopus
fissifolius (narrow-leaved carpetgrass), Clidemia hirta, Erigeron
karvinskianus, Psidium cattleianum, P. guajava, and Rubus rosifolius,
compete with M. fosbergii and modify and destroy its native habitat on
Oahu and Kauai (HBMP 2010). Hybridization is a threat to this species,
as M. fosbergii hybridizes with other Myrsine species, and the number
of non-hybrid individuals may actually be lower than estimated (Ching
Harbin 2015, in litt.). Fortini et al. (2013, p. 82) found that, as
environmental conditions are altered by climate change, M. fosbergii is
unlikely to tolerate or adapt to projected changes in temperature and
moisture, and is unlikely to be able to move to areas with more
suitable climatic conditions. Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to M. fosbergii described
above.
The remaining occurrences of Myrsine fosbergii are at risk from the
threats described above. The effects of climate change are likely to
exacerbate the threats described above. Because of these threats, we
find that M. fosbergii is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Nothocestrum latifolium (aiea) is a small tree in the nightshade
family (Solanaceae) (Symon 1999, p. 1263). Typical habitat for this
species is dry to mesic forest in the dry cliff (Kauai, Oahu, Lanai,
and Maui), lowland dry (Oahu, Lanai, and Maui), and lowland mesic
(Oahu, Molokai, Lanai, and Maui) ecosystems (TNCH 2007; HBMP 2010).
Historically, N. latifolium was known from the Waianae Mountains of
Oahu, Molokai, Lanai, and Maui (HBMP 2010; Sailer 2015, in litt.). This
species was collected once on Kauai in 1986, but has not been observed
there before or after that time (Symon 1999, p. 1263; BISH 504035-
Montgomery; Williams 2015, in litt.). Currently, on the island of Oahu,
there is one individual remaining, with only one of the other
previously extant individuals represented in an ex situ collection
(Moses 2006, in litt.; Starr 2006, in litt.; HBMP 2010; Kawakami 2010,
in litt.; Kawelo 2010, in litt.; Welton 2010, in litt.; Ching 2011, in
litt.; Ching Harbin 2015, in litt.; Sailer 2015, in litt.). On Molokai,
there a few individuals on the central south slope (Oppenheimer 2015,
in litt.). There are 18 occurrences totaling approximately 1,600
individuals on east and west Maui (Ching 2011, in litt.; HBMP 2010;
Oppenheimer 2015, in litt.). On Lanai, no individuals were found during
surveys in 2012, and this species may be extirpated from this island,
although there are plans to continue surveying suitable habitat (PEPP
2012, p. 129; Oppenheimer 2015, in litt.). In summary, the species'
range on each island has decreased dramatically since 2001 (Kawelo 2005
and 2010, in litt.; HBMP 2010; Oppenheimer 2011, in litt.).
Feral pigs (Oahu, Maui, Kauai), goats (Maui, Kauai), mouflon
(Lanai), feral cattle (Maui), axis deer (Lanai, Maui), and black-tailed
deer (Kauai) modify and destroy habitat of Nothocestrum latifolium
(HBMP 2010; Oppenheimer 2015, in litt.). Herbivory by these animals
also poses a threat to this species. Ungulates are managed in Hawaii as
game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants, such as Fraxinus uhdei (tropical
ash), Grevillea robusta (silk oak), Lantana camara, Leucaena
leucocephala (koa haole), Melinis minutiflora, Passiflora suberosa
(huehue haole), Schinus terebinthifolius, and Toona ciliata (Australian
red cedar), outcompete N. latifolium and modify and destroy habitat at
all known occurrences. Wildfire, and fire caused by military training
activities, is a threat to this species and its habitat (Sailer 2015,
in litt.). Low numbers of individuals limits this species' ability to
adapt to environmental change. Infestation by the black twig borer is a
threat to N. latifolium (Ching Harbin 2015, in litt.). This species
continues to decline, and, for unknown reasons, there is an observed
lack of regeneration in N. latifolium in the wild (HBMP 2010; Duvall
2015, in litt.). Fortini et al. (2013, p. 83) found that, as
environmental conditions are altered by climate change, N. latifolium
is unlikely to tolerate or adapt to projected changes in temperature
and moisture, and is unlikely to be able to move to areas with more
suitable climatic conditions. Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to N. latifolium described
above.
The remaining occurrences of Nothocestrum latifolium are at risk
from the threats described above. Because of these threats, we find
that this species is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Ochrosia haleakalae (holei) is a tree in the dogbane family
(Apocynaceae) (Wagner et al. 1999, p. 218). Typical habitat for this
species is dry to mesic forest, sometimes wet forest, and often lava,
from 2,300 to 4,000 ft (700 to 1,200 m), in the dry cliff (Maui),
lowland mesic (Maui and Hawaii Island), and montane mesic (Maui)
ecosystems (Medeiros et al. 1986, pp. 27-28; Wagner et al. 1999, p.
218; TNCH 2007; HBMP 2010). Historically, this species was known from
east Maui and Hawaii Island (HBMP 2010). Currently, O. haleakalae is
known from 4 occurrences totaling about 15 individuals on the island of
Maui (Medeiros 2007, in litt.; Oppenheimer 2008, in litt.; HBMP 2010;
Oppenheimer 2015, in litt.). On Hawaii Island, there are two
occurrences totaling at least 150 individuals in Hawaii Volcanoes
National Park, with 150 outplanted in nearby kipuka (vegetated areas
surrounded by lava flows), and one individual in the Laupahoehoe
section of Hilo Forest Reserve (Pratt 2005, in litt.; Bio 2008a, in
litt.; HBMP 2010; Pratt 2011, in litt.; Conry 2012, in litt.; Orlando
2015, in litt.; Perry 2015, in litt.).
Feral pigs and goats modify and destroy the habitat of Ochrosia
haleakalae on Maui and Hawaii Island; in addition, cattle modify and
destroy the habitat of this species on Maui (Medeiros 1995, in litt.;
Pratt 2005, in litt.; Oppenheimer 2015, in litt.). Herbivory by these
animals also poses a threat to this species. Ungulates are managed in
Hawaii as game animals, but public hunting does not adequately control
the numbers of ungulates to eliminate habitat modification and
destruction or herbivory by these animals (Anderson et al. 2007, in
litt.; HAR-DLNR 2010, in litt.). Nonnative plant species, such as
Cestrum diurnum (day cestrum), Fraxinus uhdei, Psidium cattleianum, P.
guajava, Rubus argutus, Setaria palmifolia (palmgrass), and Toona
ciliata, modify and destroy habitat and outcompete native plants,
including O. haleakalae (HBMP 2010). In dry areas, wildfires affecting
the habitat of this species are exacerbated by the presence of
introduced grass species such as Pennisetum clandestinum (kikuyu grass)
(HBMP 2010; Oppenheimer 2015, in litt.). Herbivory and seed predation
by slugs and rats is a threat to this species (Oppenheimer 2015, in
litt.). There is low to no reproduction observed in the wild, and this
reduced reproductive vigor is due to reduced levels of genetic
variability resulting from low numbers of individuals. This decreases
the
[[Page 67802]]
species' capacity to adapt to environmental changes, and thereby
lessens the probability of its long-term persistence (Barrett and Kohn
1991, p. 4; Newman and Pilson 1997, p. 361; Duvall 2015, in litt.).
Fortini et al. (2013, p. 83) found that, as environmental conditions
are altered by climate change, O. haleakalae is unlikely to tolerate or
adapt to projected changes in temperature and moisture, and is unlikely
to be able to move to areas with more suitable climatic conditions.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to O. haleakalae described above.
Ochrosia haleakalae is at risk from habitat degradation and loss by
feral pigs, goats, cattle, and nonnative plants; the displacement of
individuals due to competition with nonnative plants for space,
nutrients, water, air, and light; herbivory by feral pigs, goats, and
cattle; seed predation by slugs and rats; and by the small number of
remaining individuals. The effects of climate change are likely to
further exacerbate these threats. Because of these threats, we find
that this species is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Phyllostegia brevidens (NCN) is a scandent (climbing) subshrub in
the mint family (Lamiaceae) (Wagner et al. 1999, pp. 814-815). This
species occurs in wet forest on the islands of Maui and Hawaii from
2,900 to 3,200 ft (880 to 975 m), in the lowland wet (Maui), montane
wet (Hawaii Island), and wet cliff (Maui) ecosystems (Wagner et al.
1999, pp. 814-815; TNCH 2007; HBMP 2010). Phyllostegia brevidens is
historically known from Hilo Forest Reserve, Mauna Kea, and Kulani on
Hawaii Island; and from Kipahulu Valley on Maui (Haleakala National
Park) (Wagner et al. 1999, p. 815; HBMP 2010; Smithsonian Institution
2015, in litt.). Currently, there is one individual on the island of
Maui and two individuals on Hawaii Island (PEPP 2009, p. 90; Wagner et
al. 2012, p. 46; PEPP 2014, p. 136; Gates 2015, in litt.; Oppenheimer
2015, in litt.; Perry 2015, in litt.).
Feral pigs modify and destroy habitat of this species on Maui (PEPP
2014, p. 136). The two remaining individuals on Hawaii Island are
currently fenced (Perry 2015, in litt.); however, owing to the
potential for accidental damage or vandalism (irrespective of
maintenance), fences do not guarantee protection from ungulate ingress.
Herbivory by feral pigs also poses a threat to this species on Maui.
Ungulates are managed in Hawaii as game animals, but public hunting
does not adequately control the numbers of ungulates to eliminate
habitat modification and destruction or herbivory by these animals
(Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.). Nonnative
plants, such as Clidemia hirta and Hedychium gardnerianum, modify and
destroy habitat and outcompete P. brevidens on Maui (PEPP 2009, p. 90).
Herbivory by slugs is a threat to the remaining individual on Maui
(PEPP 2014, p. 136). In addition, natural events such as landslides and
erosion are threats to the occurrence on Maui (PEPP 2014, p. 136). The
small number of remaining individuals limits this species' ability to
adapt to environmental change. Fortini et al. (2013, p. 84) found that,
as environmental conditions are altered by climate change, P. brevidens
is unlikely to tolerate or adapt to projected changes in temperature
and moisture, and is unlikely to be able to move to areas with more
suitable climatic conditions. Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to P. brevidens described
above.
The remaining occurrences of Phyllostegia brevidens are at risk.
The species continues to be negatively affected by habitat modification
and destruction by ungulates and nonnative plants, and by direct
competition from nonnative plants, combined with herbivory by ungulates
and slugs. The effects of climate change are likely to further
exacerbate these threats. We find that P. brevidens is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Phyllostegia helleri (NCN) is a weakly erect to climbing shrub in
the mint family (Lamiaceae) (Wagner et al. 1999, pp. 816-817). This
species occurs on ridges or spurs from 2,800 to 4,000 ft (860 to 1,200
m) in diverse forest on Kauai in the lowland wet, montane wet, and wet
cliff ecosystems (Wagner et al. 1999, p. 817; TNCH 2007; HBMP 2010).
Historically, P. helleri was wide-ranging on the island of Kauai, from
the north and east sides throughout the central plateau (Wagner et al.
1999, p. 817; HBMP 2010). Currently, this species is limited to one
occurrence of four individuals (PEPP 2014, p. 35; Kishida 2015, in
litt.).
Feral pigs and goats modify and destroy the habitat of Phyllostegia
helleri on Kauai (HBMP 2010). Ungulates are managed in Hawaii as game
animals, but public hunting does not adequately control the numbers of
ungulates to eliminate habitat modification and destruction or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Herbivory on fruits and seeds by rats negatively
affects the remaining individuals (HBMP 2010). The only known
occurrence of this species is located at the base of cliffs, and
landslides are an additional threat (HBMP 2010). Nonnative plants, such
as Erigeron karvinskianus, Kalanchoe pinnata (air plant), Psidium
guajava, Rubus rosifolius, and various grasses, modify and destroy
native habitat, outcompete native plants, and are found at the last
known occurrence of Phyllostegia helleri (HBMP 2010). This species
experiences reduced reproductive vigor due to reduced levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, and thereby lessening the probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). Fortini
et al. (2013, p. 84) found that, as environmental conditions are
altered by climate change, P. helleri is unlikely to tolerate or adapt
to projected changes in temperature and moisture, and is unlikely to be
able to move to areas with more suitable climatic conditions. Although
we cannot predict the timing, extent, or magnitude of specific impacts,
we do expect the effects of climate change to exacerbate the threats to
P. helleri described above.
The remaining occurrence of Phyllostegia helleri is at risk. The
numbers of individuals are decreasing on Kauai, as this species was
wide-ranging on the island, extending from the north and east sides
throughout the central plateau, and is now known from only one
occurrence of four individuals. These four individuals continue to be
negatively affected by habitat modification and destruction by
ungulates and nonnative plants, direct competition by nonnative plants,
and by seed predation by rats. Natural events such as landslides may
damage or destroy the remaining four individuals. The small number of
remaining individuals limits this species' ability to adapt to
environmental changes. The effects of climate change are likely to
further exacerbate these threats. Because of these threats, we find
that P. helleri is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
[[Page 67803]]
endangered or threatened in a significant portion of its range.
Phyllostegia stachyoides (NCN) is a weakly erect to climbing
subshrub in the mint family (Lamiaceae) (Wagner et al. 1999, p. 823).
This species occurs in mesic to wet forest from 3,600 to 4,600 ft
(1,000 to 1,400 m), in the montane wet (Hawaii Island, Maui, and
Molokai) and montane mesic (Hawaii Island and Maui) ecosystems (Wagner
et al. 1999, p. 823; TNCH 2007; HBMP 2010). Phyllostegia stachyoides is
historically known from the eastern and central Molokai, west Maui, and
wide-ranging occurrences on Hawaii Island (Wagner et al. 1999, p. 823;
HBMP 2010; VanDeMark 2016, in litt.). Currently, occurrences on west
Maui total about 15 individuals (Oppenheimer 2015, in litt.). Those on
Molokai occur at 5 locations and total fewer than 30 individuals
(Orlando 2015, in litt.; PEPP 2012, p. 156). Plants on Hawaii Island
are now considered to be P. ambigua (VanDeMark 2016, in litt.).
Feral pigs, goats, and axis deer modify and destroy the habitat of
Phyllostegia stachyoides on Maui, with evidence of the activities of
these animals reported in areas where this species occurs (HBMP 2010;
PEPP 2014, p. 141). Ungulates are managed in Hawaii as game animals,
but public hunting does not adequately control the numbers of ungulates
to eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.).
Nonnative plants, such as Ageratina adenophora, Erigeron karvinskianus,
and Tibouchina herbacea, compete with P. stachyoides, modify and
destroy its native habitat, and displace other native plant species
(Smith 1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy and Stone
1990, p. 74; PEPP 2014, pp. 141-142). Herbivory by slugs and rats on
leaves and nutlets of P. stachyoides poses a threat to this species at
known locations on Maui and Molokai (PEPP 2014, pp. 140-142). On Maui,
stochastic events such as floods and drought (with ensuing erosion)
pose a threat to small, isolated occurrences of P. stachyoides;
rockfalls and landslides are a threat to occurrences on Molokai (PEPP
2014, pp. 140-142). This species experiences reduced reproductive vigor
due to reduced levels of genetic variability, leading to diminished
capacity to adapt to environmental changes, and thereby lessening the
probability of long-term persistence (Barrett and Kohn 1991, p. 4;
Newman and Pilson 1997, p. 361). Fortini et al. (2013, p. 84) found
that, as environmental conditions are altered by climate change, P.
stachyoides is unlikely to tolerate or adapt to projected changes in
temperature and moisture, and is unlikely to be able to move to areas
with more suitable climatic conditions. Although we cannot predict the
timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to P. stachyoides
described above.
The remaining occurrences of Phyllostegia stachyoides are at risk.
The known individuals are restricted to small areas on west Maui and
Molokai, and continue to be negatively affected by habitat modification
and destruction by ungulates and by direct competition with nonnative
plants, combined with herbivory by slugs and rats. The small number of
remaining individuals limits this species' ability to adapt to
environmental changes. Flooding, drought, and the effects of climate
change are likely to further exacerbate these threats. Because of these
threats, we find that this species is endangered throughout all of its
range, and, therefore, find that it is unnecessary to analyze whether
it is endangered or threatened in a significant portion of its range.
Portulaca villosa (ihi) is a perennial herb in the purslane family
(Portulacaceae) (Wagner et al. 1999, p. 1074). Portulaca villosa occurs
on dry, rocky, clay, lava, or coralline reef sites, from sea level to
1,600 ft (490 m), in the coastal (Lehua, Kaula, Oahu, Kahoolawe, Maui,
and Hawaii Island) and lowland dry (Oahu, Molokai, Lanai, Kahoolawe,
Maui, and Hawaii Island) ecosystems, and one reported occurrence in the
montane dry (Hawaii Island) ecosystem (Wagner et al. 1999, p. 1074;
TNCH 2007; HBMP 2010). Portulaca villosa is historically known from all
the main Hawaiian Islands except Niihau and Kauai (Wagner et al. 1999,
p. 1074). Portulaca villosa has been observed on the small islets of
Kaula and Lehua (west of Kauai and Niihau), and on Nihoa (NWHI);
however, the current status of these occurrences is unknown. This
species has not been observed on Oahu since the 1960s, when it was
locally abundant at Kaohikaipu Island (HBMP 2010). Historically, on the
island of Hawaii, this species occurred in the coastal area of Hawaii
Volcanoes National Park west of Kamoamoa, but was extirpated in 1993 by
lava flows (Orlando 2015, in litt.). On the island of Lanai, two
individuals were last observed in 1996 (HBMP 2010). Currently, P.
villosa is known from a few individuals on Molokai, 2 individuals on
east Maui and 24 individuals on west Maui, fewer than 15 individuals on
Kahoolawe, and five occurrences totaling 10 individuals on Hawaii
Island (MNTF 2010, in litt.; Evans 2015a, in litt.).
Axis deer (Maui and Lanai), goats (Maui), mouflon (Lanai), and
cattle (Hawaii Island) modify and destroy the habitat of Portulaca
villosa (HBMP 2010; Oppenheimer 2015, in litt.). These ungulates also
forage directly on this species. Ungulates are managed in Hawaii as
game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants, such as Lantana camara, Nicotiana
glauca (tree tobacco), Pennisetum ciliare (buffelgrass), and Prosopis
pallida (kiawe, mesquite), compete with Portulaca villosa, modify and
destroy its native habitat, displace other native plant species, and
pose a threat to the known occurrences on Hawaii Island, Maui,
Kahoolawe, and Molokai (Smith 1985, pp. 180-250; Vitousek et al. 1987
in Cuddihy and Stone 1990, p. 74). P. villosa occurs in drier coastal
and lowland habitats, all of which are affected by wildfires. Some
coastal habitat includes exposed cliffs, which erode and cause
landslides and rockfalls in areas where P. villosa occurs (Kahoolawe),
posing a threat to this species (HBMP 2010). This species experiences
reduced reproductive vigor due to low levels of genetic variability,
leading to diminished capacity to adapt to environmental changes, and
thereby lessening the probability of long-term persistence (Barrett and
Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). Fortini et al. (2013,
p. 86) found that, as environmental conditions are altered by climate
change, P. villosa is unlikely to tolerate or adapt to projected
changes in temperature and moisture, and is unlikely to be able to move
to areas with more suitable climatic conditions. Although we cannot
predict the timing, extent, or magnitude of specific impacts, we do
expect the effects of climate change to exacerbate the threats to P.
villosa described above.
The remaining occurrences of Portulaca villosa are at risk; the
numbers of individuals are decreasing on Maui, Molokai, and Hawaii
Island, and the species continues to be negatively affected by
continued habitat modification and destruction by feral ungulates and
nonnative plants, and by competition with nonnative plants. Because of
its small and isolated remaining occurrences, natural events such as
rockfalls, landslides, and wildfires may pose a threat to this species.
The small number of remaining individuals limits this species' ability
to
[[Page 67804]]
adapt to environmental changes. The effects of climate change are
likely to further exacerbate these threats. Because of these threats,
we find that this species is endangered throughout all of its range,
and, therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Pritchardia bakeri (Baker's loulu) is a small to medium-sized tree
in the palm family (Arecaceae) (Hodel 2009, pp. 173-179; Hodel 2012,
pp. 70-73). This species occurs in the lowland mesic ecosystem in the
Koolau Mountains on Oahu, from 1,500 to 2,100 ft (457 to 640 m), in
disturbed, windswept, and mostly exposed shrubby or grassy areas, and
sometimes on steep slopes in these areas (Bacon et al. 2012, pp. 1-17;
Hodel 2012, pp. 71-73). Currently, occurrences total fewer than 100
individuals (Ching Harbin 2015, in litt.).
Habitat modification and destruction by feral pigs impact the range
and abundance of Pritchardia bakeri. Ungulates are managed in Hawaii as
game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants compete with and degrade and destroy
native habitat of P. bakeri, and displace native plant species by
competing for water, nutrients, light, and space, or they may produce
chemicals that inhibit growth of other plants (Smith 1985, pp. 180-250;
Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74). Stochastic
events such as hurricanes modify and destroy the habitat of P. bakeri,
and can directly damage or kill plants. Rats eat the fruit before they
mature, leading to minimal or no recruitment (Hodel 2012, pp. 42, 73).
This species experiences reduced reproductive vigor due to low levels
of genetic variability caused by seed predation by rats and widely
separated occurrences, leading to diminished capacity to adapt to
environmental changes, and thereby lessening the probability of long-
term persistence (Barrett and Kohn 1991, p. 4; Newman and Pilson 1997,
p. 361; Hodel 2012, p. 73).
The remaining occurrences of Pritchardia bakeri are at risk; the
known individuals are restricted to small areas on Oahu, and continue
to be negatively affected by habitat degradation and loss by feral pigs
and nonnative plants, fruit predation by rats, and the small number and
reduced range of remaining individuals. Although we cannot predict the
timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to P. bakeri
described above. Based on these threats, we find that this species is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Pseudognaphalium sandwicensium var. molokaiense (enaena) is a
perennial herb in the sunflower family (Asteraceae) (Wagner et al.
1999, p. 321). Typical habitat for this variety is strand vegetation in
dry consolidated dunes, in the coastal ecosystem (Wagner et al. 1999,
p. 321; TNCH 2007; HBMP 2010). Historically, this variety was known
from Molokai, Oahu, Maui, and Lanai (HBMP 2010; MNTF 2010, in litt.).
Currently, P. sandwicensium var. molokaiense is known only from two
locations on Molokai (as many as 20,000 individuals, depending on
rainfall), and from fewer than 25 individuals on the northwest coast of
Maui (Moses 2006, in litt.; Starr 2006, in litt.; Kallstrom 2008, in
litt.; Oppenheimer 2015, in litt.). This variety was last observed on
Lanai in 1960, and on Oahu (5 individuals) in the 1980s (HBMP 2010).
Goats and axis deer modify and destroy the habitat of
Pseudognaphalium sandwicensium var. molokaiense, with evidence of the
activities of these animals reported in the areas where this plant
occurs (Moses 2006, in litt.; Starr 2006, in litt.; Kallstrom 2008, in
litt; HBMP 2010). Ungulates are managed in Hawaii as game animals, but
public hunting does not adequately control the numbers of ungulates to
eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.).
Additionally, nonnative plants, such as Atriplex semibaccata
(Australian saltbush), Chenopodium murale (aheahea, goosefoot),
Pennisetum ciliare, Prosopis pallida, and Setaria parviflora (foxtail),
compete with and displace native plant species by competing for water,
nutrients, light, and space, or they may produce chemicals that inhibit
growth of other plants (Smith 1985, pp. 180-250; Vitousek et al. 1987
in Cuddihy and Stone 1990, p. 74; Moses 2006, in litt.). This variety
experiences reduced reproductive vigor due to low levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, and thereby lessening the probability of long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361).
Rockfalls and landslides are a threat to the occurrence of this variety
on a sea cliff on west Maui (HBMP 2010). Fortini et al. (2013, p. 86)
found that, as environmental conditions are altered by climate change,
P. sandwicensium var. molokaiense is unlikely to tolerate or adapt to
projected changes in temperature and moisture, and is unlikely to be
able to move to areas with more suitable climatic conditions. Although
we cannot predict the timing, extent, or magnitude of specific impacts,
we do expect the effects of climate change to exacerbate the threats to
P. sandwicensium var. molokaiense described above.
The remaining occurrences of Pseudognaphalium sandwicensium var.
molokaiense on Molokai and Maui are at risk; individuals no longer
occur on Oahu and Lanai. Occurrences on Maui and Molokai continue to be
negatively affected by habitat modification and destruction by
ungulates, and by direct competition with nonnative plants. The small
number of remaining occurrences limits this plant's ability to adapt to
environmental changes. The effects of climate change are likely to
further exacerbate these threats. Because of these threats, we find
that this variety is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Ranunculus hawaiensis (makou) is an erect or ascending perennial
herb in the buttercup family (Ranunculaceae) (Duncan 1999, p. 1088).
Typical habitat is mesic forest on grassy slopes and scree, and in open
pastures, from 6,000 to 6,700 ft (1,800 to 2,000 m), in the montane
mesic (Hawaii Island), montane dry (Hawaii Island), and subalpine
(Hawaii Island and Maui) ecosystems (Bio 2008a, in litt; Pratt 2007, in
litt.; Duncan 1999, p. 1088; TNCH 2007; HBMP 2010). Historically, R.
hawaiensis was wide-ranging on the island of Hawaii. On Maui, this
species was known from Haleakala National Park (HBMP 2010). In the
1980s and 1990s, this species numbered several hundred individuals on
both islands. Currently, there are six occurrences totaling 14
individuals on Hawaii Island (Bio 2008a, in litt.; PEPP 2008, p. 108;
Pratt 2008, in litt.; HBMP 2010; Agorastos 2011, in litt.; Imoto 2013,
in litt.; Orlando 2015, in litt.). On Maui, a few individuals were
observed on a cliff in 1994; however, this occurrence was not relocated
in further surveys (PEPP 2013, p. 177). Additionally, no individuals
were re-observed in Haleakala National Park (DLNR 2006, p. 61).
Feral pigs, mouflon, and cattle modify and destroy the habitat of
Ranunculus hawaiensis on Hawaii Island, with
[[Page 67805]]
evidence of the activities of these animals reported in the areas where
this species occurs (HBMP 2010). These ungulates also forage on R.
hawaiensis. Ungulates are managed in Hawaii as game animals, but public
hunting does not adequately control the numbers of ungulates to
eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.).
Nonnative plants, such as Ehrharta stipoides (meadow ricegrass), Holcus
lanatus (common velvetgrass), and various grasses, modify and destroy
native habitat, outcompete native plants, and have been reported in
areas where R. hawaiensis occurs (HBMP 2010). Drought and erosion pose
a threat in the areas of the last known occurrences of R. hawaiensis on
Maui (PEPP 2013, p. 177). This species experiences reduced reproductive
vigor due to low levels of genetic variability, leading to diminished
capacity to adapt to environmental changes, and thereby lessening the
probability of long-term persistence (Barrett and Kohn 1991, p. 4;
Newman and Pilson 1997, p. 361). Fortini et al. (2013, p. 86) found
that, as environmental conditions are altered by climate change, R.
hawaiensis is unlikely to tolerate or adapt to projected changes in
temperature and moisture, and is unlikely to be able to move to areas
with more suitable climatic conditions. Although we cannot predict the
timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to R. hawaiensis
described above.
The remaining occurrences of Ranunculus hawaiensis are at risk; the
known individuals are restricted to small areas on Hawaii Island and
continue to be negatively affected by habitat modification and
destruction by feral ungulates, and by direct competition with
nonnative plants, combined with predation by ungulates. Drought and
erosion pose a threat in the areas of the last known occurrences on
Maui. The small number of remaining individuals limits this species'
ability to adapt to environmental changes. The effects of climate
change are likely to further exacerbate these threats. Because of these
threats, we find that this species is endangered throughout all of its
range, and, therefore, find that it is unnecessary to analyze whether
it is endangered or threatened in a significant portion of its range.
Ranunculus mauiensis (makou) is an erect to weakly ascending
perennial herb in the buttercup family (Ranunculaceae) (Duncan 1999, p.
1089). Typical habitat for R. mauiensis is open sites in mesic to wet
forest and along streams, from 3,500 to 5,600 ft (1,060 to 1,700 m), in
the montane wet (Kauai, Oahu, Molokai, and Maui), montane mesic (Kauai,
Molokai, Maui, and Hawaii Island), and wet cliff (Molokai and Maui)
ecosystems (Duncan 1999, p. 1089; TNCH 2007; HBMP 2010). Historically,
R. mauiensis was known from Kauai, Oahu, Molokai, Maui, and Hawaii
(HBMP 2010). Oahu occurrences have not been observed since the 1800s,
and Hawaii Island occurrences have not been observed since 1980 (HBMP
2010). Currently, R. mauiensis is known from Kauai (53 individuals) and
east Maui (112 individuals). Two individuals formerly known from
Molokai have not been observed on recent surveys (Bily 2007, in litt.;
Perlman 2007a, in litt.; Wood 2007b, in litt.; HBMP 2010; PEPP 2010, p.
105; Bakutis 2011, in litt.; PEPP 2011, p. 161; PEPP 2013, p. 177;
Oppenheimer 2015, in litt.).
Feral pigs, goats, axis deer, black-tailed deer, and cattle modify
and destroy the habitat of Ranunculus mauiensis on Kauai, Molokai, and
Maui, with evidence of the activities of these animals reported in the
areas where this species occurs (HBMP 2010; PEPP 2014, pp. 155-156).
Ungulates are managed in Hawaii as game animals (except for cattle),
but public hunting does not adequately control the numbers of ungulates
to eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.).
Nonnative plants, such as Buddleja asiatica (dog tail), Clidemia hirta,
Erigeron karvinskianus, Hedychium gardnerianum, Lantana camara,
Passiflora edulis (passion fruit), P. tarminiana, Psidium cattleianum,
Rubus argutus, R. rosifolius, and Tibouchina herbacea, modify and
destroy the native habitat of Ranunculus mauiensis and displace native
plant species by competing for water, nutrients, light, and space; they
may also produce chemicals that inhibit the growth of other plants
(Smith 1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy and Stone
1990, p. 74; HBMP 2010; PEPP 2014, p. 155). Herbivory by slugs (Maui)
and seed predation by rats (Maui, Kauai) are both reported as threats
to R. mauiensis (HBMP 2010; PEPP 2014, pp. 154-155). Stochastic events
such as drought (Maui), landslides (Kauai), and fire (Maui) are also
reported as threats to R. mauiensis (HBMP 2010). Erosion is a threat to
occurrences on Maui and Kauai (PEPP 2014, pp. 155-156). This species
experiences reduced reproductive vigor due to low levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, thereby lessening the probability of its long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). Fortini
et al. (2013, p. 86) found that, as environmental conditions are
altered by climate change, R. mauiensis is unlikely to tolerate or
adapt to projected changes in temperature and moisture, and is unlikely
to be able to move to areas with more suitable climatic conditions.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to R. mauiensis described above.
The remaining occurrences of Ranunculus mauiensis are at risk, the
known individuals are restricted to small areas on Kauai and Maui, and
continue to be negatively affected by habitat modification and
destruction by ungulates, direct competition with nonnative plants, and
herbivory and predation by slugs and rats. Because of its small,
isolated occurrences, landslides, drought, and erosion also negatively
affect this species. The small number of remaining individuals limits
this species' ability to adapt to environmental changes. The effects of
climate change are likely to further exacerbate these threats. Because
of these threats, we find that this species is endangered throughout
all of its range, and, therefore, find that it is unnecessary to
analyze whether it is endangered or threatened in a significant portion
of its range.
Sanicula sandwicensis (NCN) is a stout, erect, perennial herb in
the parsley family (Apiaceae) (Constance and Affolter 1999, p. 210).
This species occurs from 6,500 to 8,500 ft (2,000 to 2,600 m) in
shrubland and woodland on the islands of Maui and Hawaii Island, in the
montane mesic (Hawaii Island and Maui), montane dry (Hawaii Island),
and subalpine (Hawaii Island and Maui) ecosystems (Constance and
Affolter 1999, p. 210; TNCH 2007; NTBG Database 2014, in litt.).
Sanicula sandwicensis is historically known from Haleakala on Maui and
from Mauna Kea, Mauna Loa, and Hualalai on Hawaii Island (Constance and
Affolter 1999, p. 210). Currently, there are more than 50 individuals
of S. sandwicensis on east and west Maui (MNTF 2010, in litt.; PEPP
2011, pp. 162-164; Oppenheimer 2015, in litt.). In 2008, an occurrence
of fewer than 20 individuals was found in Hawaii Volcanoes National
Park (Benitez et al. 2008, p. 59). Following ungulate removal, this
occurrence increased to as many as 45 individuals, with many juvenile
plants
[[Page 67806]]
(Orlando 2015, in litt.). A single individual was found farther east at
about 7,400 ft (Orlando 2015, in litt.).
Feral pigs and goats modify and destroy the habitat of Sanicula
sandwicensis on Maui, with evidence of the activities of these animals
reported in the areas where this species occurs (PEPP 2011, pp. 162-
164; Oppenheimer 2015, in litt.). Ungulates are managed in Hawaii as
game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants modify and destroy the habitat of S.
sandwicensis and displace native plant species by competing for water,
nutrients, light, and space; they may also produce chemicals that
inhibit the growth of other plants (Smith 1985, pp. 180-250; Vitousek
et al. 1987 in Cuddihy and Stone 1990, p. 74; PEPP 2011, pp. 162-164).
Those nonnative plants observed to directly affect S. sandwicensis and
its habitat are Ageratina adenophora, Anthoxanthum odoratum (sweet
vernalgrass), Epilobium ciliatum (willow herb), Holcus lanatus (common
velvetgrass), Pinus spp., Prunella vulgaris, and Rubus argutus (PEPP
2011, pp. 162-164). Stochastic events such as drought, flooding, and
fires are all reported to pose threats to this species (PEPP 2011, pp.
162-164). Erosion is a threat to occurrences on Maui (PEPP 2011, pp.
162-163). Herbivory by rats also is a threat because they eat the
taproot, killing the plant (Oppenheimer 2015, in litt.). This species
experiences reduced reproductive vigor due to low levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, thereby lessening the probability of its long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). Fortini
et al. (2013, p. 88) found that, as environmental conditions are
altered by climate change, S. sandwicensis is unlikely to tolerate or
adapt to projected changes in temperature and moisture, and is unlikely
to be able to move to areas with more suitable climatic conditions.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to S. sandwicensis described above.
The remaining occurrences of Sanicula sandwicensis are at risk; the
known individuals are restricted to small areas on Maui and Hawaii
Island and continue to be negatively affected by habitat modification
and destruction by feral pigs and goats and by direct competition with
nonnative plants. Stochastic events such as drought, flooding, erosion,
and fires are threats to this species. The small number of remaining
individuals limits this species' ability to adapt to environmental
changes. The effects of climate change are likely to further exacerbate
these threats. Because of these threats, we find that this species is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Santalum involutum (iliahi) is a shrub or small tree in the
sandalwood family (Santalaceae) (Harbaugh et al. 2010, pp. 827-838).
Habitat for S. involutum is mesic and wet forest on Kauai, from 400 to
2,500 ft (120 to 750 m), in the lowland mesic and lowland wet
ecosystems (TNCH 2007; Harbaugh et al. 2010, pp. 827-838).
Historically, this species was known from northern Kauai at Kee,
Hanakapiai, and Wainiha, and from southern Kauai at Wahiawa, but has
not been observed in these areas for 30 years (Harbaugh et al. 2010, p.
835). Currently, approximately 50 to 100 individuals occur in isolated
forest pockets on Kauai (Harbaugh et al. 2010, p. 835; Wood 2015, in
litt.).
Feral pigs and goats modify and destroy the habitat of Santalum
involutum on Kauai, with evidence of the activities of these animals
reported in the areas where this species occurs (Harbaugh et al. 2010,
pp. 835-836; Wood 2015, in litt.). Ungulates are managed in Hawaii as
game animals, but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants modify and destroy the native habitat
of S. involutum and displace native plant species by competing for
water, nutrients, light, and space; they may also produce chemicals
that inhibit the growth of other plants (Smith 1985, pp. 180-250;
Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74; HBMP 2010).
Nonnative plants reported to modify and destroy habitat of S. involutum
include Clidemia hirta, Hedychium gardnerianum, Lantana camara, Melinis
minutiflora, Psidium cattleianum, P. guajava, and Rubus argutus
(Harbaugh et al. 2010, p. 836). Herbivory and seed predation by rats is
a threat to this species (Harbaugh et al. 2010, p. 836; Wood 2015, in
litt.). Wildfire is a threat to this species in mesic areas (Harbaugh
et al. 2010, p. 836). This species experiences reduced reproductive
vigor due to low levels of genetic variability, leading to diminished
capacity to adapt to environmental changes, thereby lessening the
probability of its long-term persistence (Barrett and Kohn 1991, p. 4;
Newman and Pilson 1997, p. 361).
The remaining occurrences of Santalum involutum are at risk; the
known individuals are restricted to a small area on Kauai and continue
to be negatively affected by habitat modification and destruction by
ungulates, direct competition with nonnative plants, and by herbivory
and fruit predation by rats. The small number of remaining individuals
limits this species' ability to adapt to environmental changes.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to S. involutum described above. Because of these threats, we
find that this species is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Schiedea diffusa ssp. diffusa (NCN) is a reclining or weakly
climbing vine in the pink family (Caryophyllaceae) (Wagner et al. 1999,
pp. 511-512; Wagner et al. 2005, pp. 103-106). Schiedea diffusa ssp.
diffusa occurs in wet forest from 3,000 to 5,300 ft (915 to 1,600 m) on
Molokai, and to 6,700 ft (2,050 m) on Maui, in the lowland wet (Maui)
and montane wet (Maui and Molokai) ecosystems (Wagner et al. 1999, p.
512; TNCH 2007; HBMP 2010). Historically, on Molokai, this subspecies
was known from Kawela to Waikolu valleys, and on Maui it was wide-
ranging on both the east and west mountains (Wagner et al. 2005, p.
106). Currently, S. diffusa ssp. diffusa is known only from east Maui
in scattered occurrences (fewer than 50 individuals total), in a much
smaller range, with some remaining in Haleakala National Park (HBMP
2010; Gates 2015, in litt.). Two occurrences were observed within
Hanawi NAR in 2005; however, their current status is unknown (Vetter
2015, in litt.). On Molokai, there were two occurrences totaling fewer
than 10 individuals; however, these have not been seen since the 1990s
(HBMP 2010; Oppenheimer 2015, in litt.).
Feral pigs modify and destroy the habitat of Schiedea diffusa ssp.
diffusa on Maui and Molokai, with evidence of the activities of these
animals reported in the areas where this subspecies occurs (HBMP 2010;
PEPP 2014, p. 159). Ungulates are managed in Hawaii as game animals,
but public hunting does not adequately control the numbers of
[[Page 67807]]
ungulates to eliminate habitat modification and destruction or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants, such as Adiantum raddianum (NCN),
Ageratina adenophora, Hypochaeris radicata (hairy cat's ear), Juncus
planifolius (rush), Passiflora tarminiana, Prunella vulgaris, Rubus
argutus, and R. rosifolius, modify and destroy the native habitat of S.
diffusa ssp. diffusa and displace native plant species by competing for
water, nutrients, light, and space; they may also produce chemicals
that inhibit the growth of other plants (Smith 1985, pp. 180-250;
Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74; HBMP 2010; PEPP
2014, p. 159). Herbivory by slugs and seed predation by rats are both
reported as threats to this subspecies (HBMP 2010; PEPP 2014, p. 159;
Duvall 2015, in litt.). This subspecies experiences reduced
reproductive vigor due to low levels of genetic variability, leading to
diminished capacity to adapt to environmental changes, thereby
lessening the probability of its long-term persistence (Barrett and
Kohn 1991, p. 4; Newman and Pilson 1997, p. 361).
The remaining occurrences of Schiedea diffusa ssp. diffusa are at
risk. The known individuals are restricted to small areas on Maui and
continue to be negatively affected by habitat modification and
destruction by ungulates, direct competition with nonnative plants, and
herbivory and predation by slugs and rats. The small number of
remaining individuals limits this subspecies' ability to adapt to
environmental changes. Although we cannot predict the timing, extent,
or magnitude of specific impacts, we do expect the effects of climate
change to exacerbate the threats to S. diffusa ssp. diffusa described
above. Because of these threats, we find that this subspecies is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Schiedea pubescens (maolioli) is a reclining or weakly climbing
vine in the pink family (Caryophyllaceae) (Wagner et al. 1999, p. 519;
Wagner et al. 2005, pp. 99-102). This species occurs in diverse mesic
to wet Metrosideros forest from 2,000 to 4,000 ft (640 to 1,220 m) in
the lowland wet, montane wet, montane mesic, and wet cliff ecosystems
(Wagner et al. 1999, 519; Wagner et al. 2005, p 100; TNCH 2007; HBMP
2010). Historically, on Molokai, this species was known from Kalae to
Pukoo ridge; on Lanai, it was known from the Lanaihale summit area but
has not been observed since 1922; on Maui, it was known from the
western mountains at Olowalu, Kaanapali, and Waihee, with a possible
occurrence the eastern mountains at Makawao (HBMP 2010). Currently,
this species is known from one occurrence on Molokai totaling fewer
than 30 individuals. The occurrence on east Maui has not been re-
observed, but this species is found at seven locations on west Maui
(Wood 2001, in litt.; Oppenheimer 2006, in litt.; Bakutis 2010, in
litt.; HBMP 2010; MNTF 2010, in litt.; Oppenheimer 2010, in litt.; PEPP
2014, pp. 162-163; Oppenheimer 2015, in litt.). It was determined that
the report of 4 to 6 individuals of S. pubescens at the PTA on Hawaii
Island was a misidentification of individuals from the species Schiedea
hawaiiensis (Wagner et al. 2005, pp. 93, 95).
Feral pigs, goats, axis deer, and cattle modify and destroy the
habitat of Schiedea pubescens on Maui and Molokai, with evidence of the
activities of these animals reported in the areas where this species
occurs (HBMP 2010; PEPP 2014, p. 162). Ungulates are managed in Hawaii
as game animals (except for cattle), but public hunting does not
adequately control the numbers of ungulates to eliminate habitat
modification and destruction or herbivory by these animals (Anderson et
al. 2007, in litt.; HAR-DLNR 2010, in litt.). Nonnative plants, such as
Buddleja asiatica, Cestrum nocturnum (night cestrum), Clidemia hirta,
Erigeron karvinskianus, Psidium cattleianum, Rubus rosifolius, and
Tibouchina herbacea, modify and destroy the native habitat of S.
pubescens and displace native plant species by competing for water,
nutrients, light, and space; they may also produce chemicals that
inhibit the growth of other plants (Smith 1985, pp. 180-250; Vitousek
et al. 1987 in Cuddihy and Stone 1990, p. 74; HBMP 2010; PEPP 2014, pp.
162-163). Herbivory by slugs and seed predation by rats are both
reported to be threats to S. pubescens on Maui (HBMP 2010; PEPP 2014,
p. 162; Duvall 2015, in litt.). Stochastic events such as drought,
erosion, fire, and flooding are also reported as threats to S.
pubescens (HBMP 2010; PEPP 2014, p. 162; Oppenheimer 2015, in litt.).
This species is outcrossing; however, very low population sizes may
have reduced its genetic variation (Weller 2015, in litt.). Fortini et
al. (2013, p. 88) found that, as environmental conditions are altered
by climate change, S. pubescens is unlikely to tolerate or adapt to
projected changes in temperature and moisture, and is unlikely to be
able to move to areas with more suitable climatic conditions. Although
we cannot predict the timing, extent, or magnitude of specific impacts,
we do expect the effects of climate change to exacerbate the threats to
S. pubescens described above.
The remaining occurrences of Schiedea pubescens are at risk. The
known individuals are restricted to small areas on Molokai and Maui,
and continue to be negatively affected by habitat modification and
destruction by ungulates, direct competition with nonnative plants, and
herbivory and predation by slugs and rats. Landslides, flooding, fire,
and drought impact this species. The small number of remaining
individuals limits this species' ability to adapt to environmental
changes. The effects of climate change are likely to further exacerbate
these threats. Because of these threats, we find that this species is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Sicyos lanceoloideus (anunu) is a perennial vine in the gourd
family (Cucurbitaceae) (Telford 1999, p. 581; Wagner and Shannon 1999,
p. 444). Sicyos lanceoloideus occurs on ridges or spurs in mesic forest
from 1,800 to 2,700 ft (550 to 800 m), in the dry cliff (Oahu), lowland
mesic (Oahu, Kauai), and montane mesic (Kauai) ecosystems (Telford
1999, p. 581; TNCH 2007; HBMP 2010). Sicyos lanceoloideus was
historically found at Kalalau Valley and Waimea Canyon on Kauai and in
the Waianae Mountains on Oahu (Telford 1999, p. 581). Currently, on
Kauai, there are four individuals in three locations (Kishida 2015, in
litt.). On Oahu, this species occurs in 5 locations in the Waianae
Mountains totaling fewer than 35 individuals (HBMP 2010; U.S. Army 2014
database). Because this species is a vine, determining exact numbers is
difficult (PEPP 2013, p. 189). In addition, occurrences and numbers
vary widely as individuals have been observed to persist for fewer than
7 years (Sailer 2015, in litt.).
Feral pigs, goats, and black-tailed deer modify and destroy the
habitat of Sicyos lanceoloideus on Kauai and Oahu, with evidence of the
activities of these animals reported in the areas where this species
occurs (HBMP 2010; PEPP 2013, p. 189; PEPP 2014, p. 166; Williams 2015,
in litt.). Ungulates are managed in Hawaii as game animals, but public
hunting does not adequately control the numbers of ungulates to
eliminate habitat modification and destruction or herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in
[[Page 67808]]
litt.). Nonnative plants, such as Clidemia hirta, Lantana camara, Melia
azedarach (chinaberry), Paspalum urvillei (vasey grass), Passiflora
edulis, Pluchea carolinensis (sourbush), Psidium cattleianum, P.
guajava, Ricinus communis (castor bean), Rubus argutus, Schinus
terebinthifolius, and Stachytarpheta jamaicensis, modify and destroy
the native habitat of Sicyos lanceoloideus, and displace native plant
species by competing for water, nutrients, light, and space; they may
also produce chemicals that inhibit the growth of other plants (Smith
1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy and Stone 1990, p.
74; HBMP 2010; Sailer 2015, in litt.). Drought is also reported as a
threat to S. lanceoloideus (PEPP 2014, p. 166; HBMP 2010; Sailer 2015,
in litt.). Fires are a threat to the occurrence in the Waianae
Mountains of Oahu (Sailer 2015, in litt.). Because of the small
remaining number of individuals, this species experiences reduced
reproductive vigor due to low levels of genetic variability, leading to
diminished capacity to adapt to environmental changes, thereby
lessening the probability of its long-term persistence (Barrett and
Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). Fortini et al. (2013,
p. 89) found that, as environmental conditions are altered by climate
change, S. lanceoloideus is unlikely to tolerate or adapt to projected
changes in temperature and moisture, and is unlikely to be able to move
to areas with more suitable climatic conditions. Although we cannot
predict the timing, extent, or magnitude of specific impacts, we do
expect the effects of climate change to exacerbate the threats to S.
lanceoloideus described above.
The remaining occurrences of Sicyos lanceoloideus are at risk. The
known individuals are restricted to small areas on Kauai and Oahu and
continue to be negatively affected by habitat modification and
destruction by ungulates, direct competition with nonnative plants, and
stochastic events such as drought and fire. The small number of
remaining individuals limits this species' ability to adapt to
environmental change. The effects of climate change are likely to
further exacerbate these threats. Because of these threats, we find
that this species is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Sicyos macrophyllus (anunu) is a perennial vine in the gourd family
(Cucurbitaceae) (Telford 1999, p. 578; Wagner and Shannon 1999, p.
444). Typical habitat is wet Metrosideros polymorpha forest and Sophora
chrysophylla-Myoporum sandwicense (mamane-naio) forest, from 4,000 to
6,600 ft (1,200 to 2,000 m) in the montane mesic (Hawaii Island),
montane wet (Maui), and montane dry (Hawaii Island) ecosystems (Telford
1999, p. 578; TNCH 2007; HBMP 2010). Historically, S. macrophyllus was
known from Puuwaawaa, Laupahoehoe, Puna, and South Kona on Hawaii
Island, and from Kipahulu Valley on the island of Maui (HBMP 2010).
Currently, S. macrophyllus is known from 10 occurrences totaling
between 24 and 26 individuals on Hawaii Island (Bio 2008, pers. comm.;
Pratt 2008, in litt.; HBMP 2010; Evans 2015b, in litt.; Orlando 2015,
in litt.). This species has been outplanted at several sites in Hawaii
Volcanoes National Park and is persisting (Orlando 2015, in litt.). The
individual on Maui has not been observed since 1987 (HBMP 2010).
Feral pigs, mouflon, and cattle modify and destroy the habitat of
Sicyos macrophyllus on the island of Hawaii, with evidence of the
activities of these animals reported in the areas where this species
occurs (HBMP 2010). Ungulates are managed in Hawaii as game animals
(except for cattle), but public hunting does not adequately control the
numbers of ungulates to eliminate habitat modification and destruction
or herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants, such as Cenchrus setaceus, Delairea
odorata (German ivy), Ehrharta stipoides, and Pennisetum clandestinum,
modify and destroy the native habitat of S. macrophyllus and displace
native plant species by competing for water, nutrients, light, and
space; they may also produce chemicals that inhibit the growth of other
plants (Smith 1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy and
Stone 1990, p. 74; HBMP 2010). Seed predation by rats is reported to
pose a threat to this species (HBMP 2010). Stochastic events such as
fire are also reported as a threat to S. macrophyllus (HBMP 2010). This
species experiences reduced reproductive vigor due to low levels of
genetic variability, leading to diminished capacity to adapt to
environmental changes, thereby lessening the probability of its long-
term persistence (Barrett and Kohn 1991, p. 4; Newman and Pilson 1997,
p. 361). Fortini et al. (2013, p. 89) found that, as environmental
conditions are altered by climate change, S. macrophyllus is unlikely
to tolerate or adapt to projected changes in temperature and moisture,
and is unlikely to be able to move to areas with more suitable climatic
conditions. Although we cannot predict the timing, extent, or magnitude
of specific impacts, we do expect the effects of climate change to
exacerbate the threats to S. macrophyllus described above.
The remaining occurrences of Sicyos macrophyllus are at risk. The
only known individuals are restricted to small areas on Hawaii Island
and continue to be negatively affected habitat modification and
destruction by ungulates, direct competition with nonnative plants, and
seed predation by rats. Fire is also a threat to this species. The
small number of remaining individuals limits this species' ability to
adapt to environmental changes. The effects of climate change are
likely to further exacerbate these threats. Because of these threats,
we find that this species is endangered throughout all of its range,
and, therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Solanum nelsonii (popolo) is a sprawling or trailing shrub up to 3
ft (1 m) tall, in the nightshade family (Solanaceae) (Symon 1999, pp.
1273-1274). Typical habitat for this species is coral rubble or sand in
coastal sites up to 490 ft (150 m), in the coastal ecosystem (Symon
1999, pp. 1273-1274; TNCH 2007; HBMP 2010). Historically, S. nelsonii
was known from Kaalualu, Kamilo, and Kaulana Bay, and South Point (5
individuals total) on Hawaii Island; from Kealea Bay, Kawaewaae, and
Leahi on Niihau; and from the Northwest Hawaiian Islands of Nihoa,
Laysan, Pearl and Hermes, and Kure Atoll (Green Island) (Lamoureux
1963, p. 6; Clapp et al. 1977, p. 36; HBMP 2010). This species was last
collected on Niihau in 1949 (HBMP 2010). The only known individual on
Maui was reported to have disappeared in the mid-1990s after cattle had
been allowed to graze in its last known habitat (HBMP 2010; Duvall
2015, in litt.). Currently, S. nelsonii occurs in the coastal ecosystem
on the islands of Hawaii and Molokai (approximately 50 individuals),
and on the Northwest Hawaiian Islands of Kure (an unknown number of
individuals), Midway (approximately 260 individuals on Sand, Eastern,
and Spit islands), Laysan (approximately 490 individuals), Pearl and
Hermes (30 to 100 individuals), and Nihoa (8,000 to 15,000 individuals)
(Aruch 2006, in litt.; Rehkemper 2006, in litt.; Tangalin 2006, in
litt.; Bio 2008 a and 2008b, in litt.; Vanderlip 2011, in litt.; Conry
2012, in litt.; PEPP 2013, pp. 190-191).
[[Page 67809]]
Axis deer and feral cattle modify and destroy the habitat of
Solanum nelsonii on the main Hawaiian islands of Maui, Molokai, and
Hawaii, with evidence of the activities of these animals reported in
the areas where this species occurs (HBMP 2010). Ungulates are managed
in Hawaii as game animals (except for cattle), but public hunting does
not adequately control the numbers of ungulates to eliminate habitat
modification and destruction, and herbivory by these animals (Anderson
et al. 2007, in litt.; HAR-DLNR 2010, in litt.). Nonnative plants, such
as Lantana camara, Leucaena leucocephala, Pennisetum ciliare, Prosopis
pallida, and Setaria verticillata (bristly foxtail), modify and destroy
the native habitat of S. nelsonii both on the main Hawaiian Islands and
on some of the Northwest Hawaiian Islands (HBMP 2010). Nonnative plants
displace native plant species by competing for water, nutrients, light,
and space, or they may produce chemicals that inhibit the growth of
other plants (Smith 1985, pp. 180-250; Vitousek et al. 1987 in Cuddihy
and Stone 1990, p. 74; PEPP 2008, p. 110; HBMP 2010). Seed predation by
rats has been reported as a threat to S. nelsonii on Molokai (PEPP
2014, p. 167). Stochastic events such as drought, erosion, fire, and
flooding are also reported as threats to S. nelsonii (HBMP 2010; PEPP
2014, p. 167). In 2011, a tsunami swept over Midway Atoll's Eastern
Island and Kure Atoll's Green Island, inundating S. nelsonii plants,
spreading plastic debris, and destroying seabird nesting areas,
reaching about 500 ft (150 m) inland (DOFAW 2011, in litt.; Starr 2011,
in litt.; USFWS 2011, in litt.). Occurrences of this species on the
main Hawaiian Islands and on some of the Northwest Hawaiian Islands
experience reduced reproductive vigor due to low levels of genetic
variability, leading to diminished capacity to adapt to environmental
changes, thereby lessening the probability of its long-term persistence
(Barrett and Kohn 1991, p. 4; Newman and Pilson 1997, p. 361). The
effects of climate change resulting in sea-level rise will alter
environmental conditions and ecosystem that support this species.
Fortini et al. (2013, p. 89) found that, as environmental conditions
are altered by climate change, S. nelsonii is unlikely to tolerate or
adapt to projected changes in temperature and moisture, and is unlikely
to be able to move to areas with more suitable climatic conditions.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to S. nelsonii described above.
The remaining occurrences of Solanum nelsonii on the main Hawaiian
Islands are restricted to small areas of Molokai and Hawaii Island, and
continue to be negatively affected by habitat modification and
destruction by ungulates, direct competition with nonnative plants, and
herbivory and predation by rats. Even though most individuals of S.
nelsonii in the Northwestern Hawaii Islands are found on lands managed
by the Service as part of the Hawaiian Islands National Wildlife
Refuge, the relatively isolated occurrences of S. nelsonii there are
negatively affected by nonnative plants. The small number of remaining
individuals limits this species' ability to adapt to environmental
changes. A tsunami occurred and impacted habitat for this species, and
sea level rise associated with global warming will modify and destroy
habitat for S. nelsonii in the low-lying Northwestern Hawaiian Islands.
Because of these threats, we find that this species is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Stenogyne kaalae ssp. sherffii (NCN) is a climbing vine in the mint
family (Lamiaceae) (Wagner and Weller 1999, pp. 448-449; Weller and
Sakai 1999, p. 838). This species occurs in the Koolau Mountains of
Oahu, in diverse forest from 1,500 to 1,600 ft (450 to 490 m) in the
lowland wet ecosystem (Wagner and Weller 1999, pp. 448-449; TNCH 2007;
HBMP 2010; U.S. Army 2014 database). Stenogyne kaalae ssp. sherffii is
historically known from diverse mesic forest in the Waianae Mountains
of Oahu, and from the lowland wet ecosystem of the Koolau Mountains
(although, as described in the proposed rule, it was thought to be a
different species, S. sherffii, until the mid-1990s). This subspecies
occurred within a very small range in the northern Koolau Mountains,
but now all wild individuals are extirpated. There are propagules from
collections from those plants that have been outplanted in the same
areas (PEPP 2014, p. 169; Ching Harbin 2015, in litt.).
Feral pigs modify and destroy the habitat of Stenogyne kaalae ssp.
sherffii on Oahu, with evidence of the activities of these animals
reported in the areas where this subspecies occurred (HBMP 2010; PEPP
2014, p. 169). Ungulates are managed in Hawaii as game animals, but
public hunting does not adequately control the numbers of ungulates to
eliminate habitat destruction and modification, and herbivory by these
animals (Anderson et al. 2007, in litt.; HAR-DLNR 2010, in litt.).
Nonnative plants, such as Blechnum appendiculatum (NCN), Clidemia
hirta, Cyclosorus parasiticus (NCN), Psidium cattleianum, and Rubus
rosifolius, destroy and modify the native habitat of S. kaalae ssp.
sherffii and displace native plant species by competing for water,
nutrients, light, and space; they may also produce chemicals that
inhibit the growth of other plants (Smith 1985, pp. 180-250; Vitousek
et al. 1987 in Cuddihy and Stone 1990, p. 74; HBMP 2010). This
subspecies experiences reduced reproductive vigor due to low levels of
genetic variability, leading to diminished capacity to adapt to
environmental changes, thereby lessening the probability of its long-
term persistence (Barrett and Kohn 1991, p. 4; Newman and Pilson 1997,
p. 361). Fortini et al. (2013, p. 90) found that, as environmental
conditions are altered by climate change, S. kaalae ssp. sherffii is
unlikely to tolerate or adapt to projected changes in temperature and
moisture, and is unlikely to be able to move to areas with more
suitable climatic conditions. Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to S. kaalae ssp. sherffii
described above.
Any remaining occurrences of Stenogyne kaalae ssp. sherffii are at
risk; the last known wild individuals were restricted to a very small
area on Oahu, and the habitat continues to be negatively affected by
habitat modification and destruction by ungulates and direct
competition with nonnative plants. The small number of remaining
individuals (outplanted only) limits this subspecies' ability to adapt
to environmental changes. The effects of climate change are likely to
further exacerbate these threats. Because of these threats, we find
that this subspecies is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Wikstroemia skottsbergiana (akia) is a shrub or small tree in the
akia family (Thymelaceae) (Peterson 1999, p. 1290). Wikstroemia
skottsbergiana occurs in wet forest on the island of Kauai, in the
lowland wet ecosystem (Peterson 1999, p. 1290; TNCH 2007). Wikstroemia
skottsbergiana is historically known from the Wahiawa Mountains,
Hanalei Valley, and Kauhao Valley, on the island of Kauai (Peterson
1999, p. 1290). Currently, this species is limited to 30
[[Page 67810]]
individuals at one location (PEPP 2012, p. 26; Wood 2015, in litt.).
Feral pigs and goats destroy and modify the habitat of Wikstroemia
skottsbergiana on Kauai, with evidence of the activities of these
animals reported in the areas where this species occurs (DLNR 2005, in
litt.; Wood 2015, in litt.). Ungulates are managed in Hawaii as game
animals, but public hunting does not adequately control the numbers of
ungulates to eliminate habitat destruction and modification or
herbivory by these animals (Anderson et al. 2007, in litt.; HAR-DLNR
2010, in litt.). Nonnative plants destroy and modify the native habitat
of W. skottsbergiana and displace native plant species by competing for
water, nutrients, light, and space; they may also produce chemicals
that inhibit the growth of other plants (Smith 1985, pp. 180-250;
Vitousek et al. 1987 in Cuddihy and Stone 1990, p. 74; HBMP 2010).
Predation of seeds by rats is a threat to this species (DLNR 2005, in
litt.). Landslides are a threat to the only known occurrence of this
species (Wood 2015, in litt.). This species experiences reduced
reproductive vigor due to low levels of genetic variability, leading to
diminished capacity to adapt to environmental changes, thereby
lessening the probability of its long-term persistence (Barrett and
Kohn 1991, p. 4; Newman and Pilson 1997, p. 361; DLNR 2005, in litt.).
The remaining occurrences of Wikstroemia skottsbergiana are at
risk. The known individuals are restricted to a very small area on
Kauai and continue to be negatively affected by habitat modification
and destruction by ungulates, direct competition with nonnative plants,
and seed predation by rats. The small number of remaining individuals
limits this species' ability to adapt to environmental changes.
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to W. skottsbergiana described above. Because of these threats,
we find that this species is endangered throughout all of its range,
and, therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Animals
Band-Rumped Storm-Petrel (Oceanodroma castro)--Hawaii Population
The band-rumped storm-petrel, a small seabird, is a member of the
family Hydrobatidae (order Procellariiformes) and a member of the
Northern Hemisphere subfamily Hyrdrobatinae (Slotterback 2002, p. 2).
This seabird is found in several areas of the subtropical Pacific and
Atlantic Oceans (del Hoyo et al. 1992 in Bird Life International 2015,
in litt.). The Atlantic breeding populations are restricted to islands
in the eastern portions: Cape Verde, Ascension, Madeira, and the Azores
Islands (Allan 1962, p. 274; Harrison 1983, p. 274). Wintering birds
may occur as far west as the mid-Atlantic; however, Atlantic breeding
populations are not within the borders of the United States or in areas
under U.S. jurisdiction. Three widely separated breeding areas occur in
the Pacific: in Japan, Hawaii, and Galapagos (Richardson 1957, p. 19;
Harris 1969, p. 96; Harrison 1983, p. 274). The Japanese population,
which breeds on islets off the east coast of Japan (Hidejima and
Sanganjima in Allan 1962, p. 274; Harris 1969, p. 96), ranges within
860 mi (1,400 km) east and south of the breeding colonies. Populations
in Japan and Galapagos total as many as 23,000 pairs (Boersma and Groom
1993, p. 114); however, a recent survey on Hidejima Island revealed
only 117 burrows, some of which were occupied by Leach's storm petrels
(Oceanodroma leucorhoa) (Biodiversity Center of Japan 2014, p. 1).
Surveyors noted that the nesting area had been affected by extensive
erosion caused by the 2011 earthquake and tsunami (Biodiversity Center
of Japan 2014, p. 1).
When Polynesians arrived about 1,500 years ago, the band-rumped
storm-petrel probably was common on all of the main Hawaiian Islands
(Harrison et al. 1990, pp. 47-48). As evidenced by bones found in
middens on Hawaii Island (Harrison et al. 1990, pp. 47-48) and in
excavation sites on Oahu and Molokai (Olson and James 1982, pp. 30,
33), band-rumped storm-petrels were once numerous enough to be
harvested for food and possibly for their feathers (Harrison et al.
1990, p. 48).
In Hawaii, band-rumped storm-petrels are known to nest in remote
cliff locations on Kauai and Lehua Island, in steep open to vegetated
cliffs, and in little vegetated, high-elevation lava fields on Hawaii
Island (Wood et al. 2002, p. 17-18; VanderWerf et al. 2007, pp. 1, 5;
Joyce and Holmes 2010, p. 3; Banko 2015 in litt.; Raine 2015, in
litt.). Vocalizations were heard in Haleakala Crater on Maui in 1992
(Johnston 1992, in Wood et al. 2002, p. 2), on Lanai (Penniman 2015, in
litt.), and in Hawaii Volcanoes National Park (Orlando 2015, in litt.).
Based on the scarcity of known breeding colonies in Hawaii and their
remote, inaccessible locations today compared to prehistoric population
levels and distribution, the band-rumped storm-petrel appears to be
significantly reduced in numbers and range following human occupation
of the Hawaiian Islands, likely as a result of predation by nonnative
mammals and habitat loss.
Taxonomists have typically combined the Pacific populations of
band-rumped storm-petrel into a single taxon, and currently the
American Ornithologist's Union (AOU) regards the species as monotypic
(2015, in litt.). However, molecular studies are ongoing and indicate
genetic differences between populations in different oceans and
archipelagos (Friesen et al. 2007b, pp. 18590-18952; Smith et al. 2007,
p. 770; Taylor et al., in prep in Raine 2015, in litt.) and between
sympatric populations that breed in different seasons (e.g., in the
Galapagos Islands; Smith and Friesen 2007, pp. 1599-1560; Smith et al.
2007, p. 756).
Band-rumped storm-petrels are regularly observed in coastal waters
around Kauai, Niihau, and Hawaii Island (Harrison et al. 1990, p. 49;
Holmes and Joyce 2009, 4 pp.), and in ``rafts'' (regular
concentrations) of a few birds to as many as 100, possibly awaiting
nightfall before coming ashore to breeding colonies. Kauai likely has
the largest population, with an estimated 221 nesting pairs in cliffs
along the north shore of the island in 2002, and additional
observations on the north and south side of the island in 2010
(Harrison et al. 1990, p. 49; Wood et al. 2002, pp. 2-3; Holmes and
Joyce 2009, 4 pp.; Joyce and Holmes 2010, pp. 1-3). Audio detections
for Kauai indicate this species may be predominantly breeding on the Na
Pali coast and Waimea Canyon, with a very small number in Wainiha
Valley (Raine 2015, in litt.). The band-rumped storm-petrel is also
known from Lehua Island (as detected there by auditory surveys)
(VanderWerf et al. 2007, p.1; Raine 2015, in litt.), Maui (Mitchell et
al. 2005, in litt.), Kahoolawe (Olson 1992, pp. 38, 112), Lanai
(Penniman 2015, in litt.) and Hawaii Island (Mitchell et al. 2005, in
litt.; Orlando 2015, in litt.). Additional surveys have been conducted
on several islands in recent years, including surveys confirming the
presence of band-rumped storm-petrels at the PTA on Hawaii Island, but
further data are not yet available (Swift 2015, in litt.). The species
likely once nested in coastal Maui, where the remains of a chick were
found in 1999, and islands such as Niihau and Kaula, where surveys have
not been conducted, likely have suitable nesting habitat and may harbor
the species (Penniman 2015, in litt.). We do
[[Page 67811]]
not have a current estimate of total numbers in Hawaii at this time.
Nesting sites are in burrows and in crevices, holes, and on
protected ledges along cliff faces, where a single egg is laid (Allan
1962, p. 274-275; Harris 1969, pp. 104-105; Slotterback 2002, p. 11).
Predation by nonnative animals on nests and adults during the breeding
season is the greatest threat to the Hawaiian population of the band-
rumped storm-petrel. These predators include feral cats (Felis catus),
barn owls (Tyto alba), small Indian mongoose (Herpestes auropunctatus),
black rats (Rattus rattus), Norway rats (R. norvegicus), and Polynesian
rats (R. exulans) (Scott et al. 1986, pp. 1, 363-364; Tomich 1986, pp.
37-45; Harrison et al. 1990, pp. 47-48; Slotterback 2002, p. 19; Raine
2015, in litt.). Attraction of fledglings to artificial lights and
collisions with structures, such as communication towers and utility
lines, is also a threat (Reed et al. 1985, p. 377; Telfer et al. 1987,
pp. 412-413; Harrison et al. 1990, p. 49; Banko et al. 1991, p. 651;
Cooper and Day 1998, p. 18; Podolsky et al. 1998, pp. 21, 27-30; Holmes
and Joyce 2009, p. 2). Monitoring of power lines on Kauai has recorded
over 1,000 strikes by seabirds annually (mostly Newell's shearwaters
(Puffinus newelli); Travers et al. 2014, pp. 19, 42) that may result in
injury or death. Recent studies of attraction of seabirds to artificial
lights indicate that 40 percent of those downed by exhaustion (from
circling the lights) are killed by collisions with cars or other
objects (Anderson 2015, p. 4-13). The small numbers of these birds and
their nesting areas on remote cliffs make population-level impacts
difficult to document. However, the band-rumped storm-petrel has
similar behavior, life-history traits, and habitat needs to the
Newell's shearwater, a threatened species that has sustained major
losses as a result of light attraction and collisions with lines or
other objects (Banko et al. 1991, p. 651; Banko 2015, in litt.; Raine
2015, in litt.). Therefore, we conclude that these are threats to the
band-rumped storm-petrel as well. Erosion and landslides at nest sites
caused by the actions of nonnative ungulates is a threat in some
locations on the island of Kauai (Raine 2015, in litt.). Nonnative
plants outcompete native plants and can also affect nesting sites of
the band-rumped storm-petrel by accelerating erosion, leading to
landslides and rockfalls (Wood et al. 2002, pp. 7-19). Regulatory
mechanisms (e.g., the Migratory Bird Treaty Act (MBTA; 16 U.S.C. 703 et
seq.)) contribute minimally to the active recovery and management of
this species (USFWS 2013, in litt.). The small population size and
limited distribution of the band-rumped storm-petrel in Hawaii is a
threat to this population (Soul[eacute] 1987, p. 8; Lande 1988, pp.
1455, 1458-1459; Harrison et al. 1990, p. 50; Furness 2003, p. 33).
During the breeding season, a single hurricane or landslide caused by
erosion could cause reproductive failure and kill a significant number
of adult birds. Commercial fisheries and ocean pollution have negative
impacts to seabirds, and also are likely to have negative impacts to
the band-rumped storm petrel, although the information about the
impacts of fisheries and plastics on storm-petrel species is limited.
In this rule, our listing determination applies only to the Hawaiian
population of the band-rumped storm-petrel (see Distinct Population
Segment (DPS) Analysis, below). Because of the deleterious and
cumulative effects to the band-rumped storm-petrel caused by the
threats described above, we find that the Hawaii population is
endangered throughout its range, and, therefore, we find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Yellow-Faced Bees (Hylaeus spp.)
Bees in the genus Hylaeus (family Colletidae), which includes the
seven species in this final rule, are commonly known as yellow-faced
bees or masked bees for their yellow-to-white facial markings. All
Hylaeus bees roughly resemble small wasps in appearance; however,
Hylaeus bees have plumose (branched) hairs on the body that are longest
on the sides of the thorax, which readily distinguish them from wasps
(Michener 2000, in litt.). Bees in the family Colletidae are also
referred to as plasterer bees because they line their nests with a
self-secreted, cellophane-like material. Eggs hatch and develop into
larvae (immature stage) and as larvae grow, they molt through three
successive stages (instars), then change into pupae (a resting form) in
which they metamorphose and emerge as adults (Michener 2000, in litt.).
The diet of the larval stages is unknown, although it is presumed the
larvae feed on stores of pollen and nectar collected and deposited in
the nest by the adult female.
Yellow-Faced Bee (Hylaeus anthracinus)
Hylaeus anthracinus was historically known from numerous coastal
and lowland dry forest habitats up to 2,000 ft (610 m) in elevation on
the islands of Hawaii, Maui, Lanai, Molokai, and Oahu, and in some
areas was ``locally abundant.'' Between 1997 and 1998, surveys for
Hawaiian Hylaeus were conducted at 43 sites that were either historical
collecting localities or potential suitable habitat. Hylaeus
anthracinus was observed at 13 of the 43 survey sites, but was not
found at any of the 9 historically occupied sites (Daly and Magnacca
2003, p. 217). Several of the historical collection sites have been
urbanized or are dominated by nonnative vegetation (Liebherr and
Polhemus 1997, pp. 346-347; Daly and Magnacca 2003, p. 55; Magnacca
2007, pp. 186-188). There has been a dramatic decline in abundance or
presence of H. anthracinus since surveys conducted in 1999 through
2002, noted on surveys conducted between 2011 and 2013 (Magnacca 2015,
in litt.). Currently, H. anthracinus is known from 15 small patches of
coastal and lowland dry forest habitat (Magnacca 2005a, in litt., p.
2); 5 locations on the island of Hawaii in the coastal ecosystem; 2
locations on Maui in the coastal and lowland dry ecosystems; 1 location
on Kahoolawe in the lowland dry ecosystem; 3 locations on Molokai in
the coastal ecosystem, and 4 locations on Oahu in the coastal ecosystem
(Daly and Magnacca 2003, p. 217; Magnacca 2005a, in litt., p. 2;
Magnacca and King 2013, pp. 13-14; Graham 2015, in litt.). These 15
locations supported small populations of H. anthracinus, but the number
of individual bees is unknown. In 2004, a single individual was
collected in montane dry forest on the island of Hawaii (possibly a
vagrant); however, the presence of additional individuals has not been
confirmed at this site (Magnacca 2005a, in litt., p. 2). Although this
species was previously unknown from the island of Kahoolawe, it was
observed at one location on the island in 2002 (Daly and Magnacca 2003,
p. 55). Additionally, during surveys between 1997 and 2008, H.
anthracinus was absent from 17 other sites with potentially suitable
habitat from which other species of Hylaeus were collected (Daly and
Magnacca 2003, pp. 4, 55) on Hawaii Island, Maui, Lanai, Molokai, and
Oahu.
Habitat destruction and modification by urbanization and land use
conversion lead to the direct fragmentation of foraging and nesting
areas used by Hylaeus anthracinus. Habitat destruction and modification
by nonnative plants adversely impacts native plant species by modifying
the availability of light, altering soil-water regimes, modifying
nutrient cycling, altering the fire characteristics
[[Page 67812]]
(increasing the fire cycle), and ultimately converting native dominated
plant communities to nonnative plant communities, and results in
removal of food sources and nesting sites for H. anthracinus (Graham
2015, in litt.). Habitat modification and destruction by nonnative
animals such as feral pigs, goats, axis deer, and cattle, is considered
one of the primary factors underlying degradation of native vegetation
in the Hawaiian Islands, and these habitat changes also remove food
sources and nesting sites for H. anthracinus (Stone 1985, pp. 262-263;
Cuddihy and Stone 1990, pp. 60-66, 73). Fire is a threat to H.
anthracinus, as it destroys native coastal and lowland plant
communities on which the species depends, and opens habitat for
increased invasion by nonnative plants. Because of the greater
frequency, intensity, and duration of fires that have resulted from the
human alteration of landscapes and the introduction of nonnative
plants, especially grasses, fires are now more destructive to the
coastal and lowland dry ecosystems (Brown and Smith 2000, p. 172). A
single grass-fueled fire often kills most native trees and shrubs in
the area (D'Antonio and Vitousek 1992, p. 74) and could destroy food
and nesting resources for H. anthracinus. The number and size of
wildfires are increasing in the main Hawaiian Islands; however, their
occurrences and locations are unpredictable, and could affect habitat
for yellow-faced bees at any time (Gima 1998, in litt.; County of Maui
2009, Ch. 3, p. 3; Hamilton 2009, in litt.; Honolulu Advertiser 2010,
in litt.; Pacific Disaster Center 2011, in litt.). Random, naturally
occurring events such as hurricanes, tsunami, and drought can also
modify and destroy habitat of H. anthracinus by creating disturbed
areas conducive to invasion by nonnative plants and by eliminating food
and nesting resources (Kitayama and Mueller-Dombois 1995, p. 671;
Businger 1998, pp. 1-2; Magnacca 2015, in litt.). Predation by
nonnative ants including the big-headed ant (Pheidole megacephala), the
yellow crazy ant (Anoplolepis gracilipes), Solenopsis papuana (NCN),
and S. geminata (NCN) on Hylaeus egg, larvae, and pupal stages is a
threat to H. anthracinus, and ants also compete with H. anthracinus for
their nectar food and nesting resources (Howarth 1985, p. 155; Hopper
et al. 1996, p. 9; Holway et al. 2002, pp. 188, 209; Daly and Magnacca
2003, p. 9; Lach 2008, p. 155; Graham 2015, in litt.). Predation by
nonnative western yellow jacket wasps (Vespula pensylvanica) is a
threat to H. anthracinus because the wasp is an aggressive, generalist
predator, and occurs in great numbers in many habitat types, from sea
level to over 8,000 ft (2,450 m), including areas where H. anthracinus
and other yellow-faced bees occur (Gambino et al. 1987, p. 169; Graham
2015, in litt.). Existing regulatory mechanisms and agency policies do
not address the primary threats to the yellow-faced bees and their
habitat from nonnative ungulates. Competition with nonnative bees
(honeybees, carpenter bees, sweat bees (Lasioglossum spp.), and alien
Hylaeus bees) for nectar and pollen, and by exclusion from foraging, is
a potential threat to H. anthracinus (Magnacca 2007, p. 188; Graham
2015, in litt.; Magnacca 2015, in litt.). The small number of
populations and individuals of H. anthracinus makes this species more
vulnerable to extinction because of the higher risks from genetic
bottlenecks, random demographic fluctuations, and localized
catastrophes such as hurricanes, tsunami, and drought (Daly and
Magnacca 2003, p. 3; Magnacca 2007, p. 173; Magnacca 2015, in litt.).
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to H. anthracinus described above. In addition, disease has
been suggested as a threat, as pathogens carried by nonnative bees,
wasps, and ants could be transmitted to H. anthracinus through shared
food sources (Graham 2015, in litt.); however, we have no reports of
this type of disease transmission at this time.
The remaining populations of Hylaeus anthracinus and its habitat
are at risk. The known individuals are restricted to 15 locations on
Hawaii, Maui, Kahoolawe, Molokai, and Oahu and continue to be
negatively affected by habitat destruction and modification by
urbanization and land-use conversion, and by habitat destruction and
removal of food and nesting sites by nonnative ungulates and nonnative
plants. Habitat destruction by fire is a threat. Randomly occurring
events such as hurricanes and drought modify habitat and remove food
and nesting sources for H. anthracinus. Predation by nonnative ants and
wasps is a threat. Existing regulatory mechanisms and agency policies
do not address the primary threats to the yellow-faced bees and their
habitat from nonnative ungulates. Competition with nonnative bees for
food and nesting sites is a threat. The small number of remaining
populations limits this species' ability to adapt to environmental
changes. The effects of climate change are likely to further exacerbate
these threats. Because of these threats, we find that H. anthracinus is
endangered throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Yellow-Faced Bee (Hylaeus assimulans)
Historically, Hylaeus assimulans was known from numerous coastal
and lowland dry forest habitats up to 2,000 ft (610 m) in elevation on
the islands of Maui (coastal and lowland dry ecosystems), Lanai
(lowland dry ecosystem), and Oahu (coastal and lowland dry ecosystem).
There are no collections from Molokai although it is likely H.
assimulans occurred there because all other species of Hylaeus known
from Maui, Lanai, and Oahu also occurred on Molokai (Daly and Magnacca
2003, pp. 217-229). Between 1997 and 1998, surveys for Hawaiian Hylaeus
were conducted at 25 sites on Maui, Kahoolawe, Lanai, Molokai, and
Oahu. Hylaeus assimulans was absent from 6 of its historical localities
on Maui, Lanai, and Oahu, and was not observed at the remaining 19
sites with potentially suitable habitat (Daly and Magnacca 2003, pp.
56, 217; Magnacca 2005b, in litt., p. 2; Magnacca 2007, pp. 177, 181,
183; Xerces Society 2009, p. 4). Currently, H. assimulans is known from
a few small patches of coastal and lowland dry forest habitat (Magnacca
2005b, in litt., p. 2) in two locations on Maui in the lowland dry
ecosystem; one location on Kahoolawe in the coastal ecosystem; and two
locations on Lanai in the lowland dry ecosystem (Daly and Magnacca
2003, p. 58; Magnacca 2005b, in litt., p. 2). This species has likely
been extirpated from Oahu because it has not been observed since
Perkin's 1899 surveys, and was not found during recent surveys of
potentially suitable habitat on Oahu at Kaena Point, Makapuu, and
Kalaeloa (Daly and Magnacca 2003, p. 217; Magnacca 2005b, in litt., p.
2).
Habitat destruction and modification by urbanization and land use
conversion lead to fragmentation of, and eventual loss, of foraging and
nesting areas used by Hylaeus assimulans. Habitat destruction and
modification by nonnative plants (Asystasia gangetica (Chinese violet),
Atriplex semibaccata, Cenchrus ciliaris (buffelgrass), Chloris barbata
(swollen fingergrass), Digitaria insularis (sourgrass), Leucaena
leucocephala, Melinis minutiflora, Pluchea indica (Indian fleabane), P.
carolinensis, Prosopis pallida, Schinus terebinthifolius, and Verbesina
encelioides (golden crown-beard)
[[Page 67813]]
adversely impact native plant species by modifying the availability of
light, altering soil-water regimes, modifying nutrient cycling,
altering the fire characteristics, and ultimately converting native
dominated plant communities to nonnative plant communities, and results
in removal of food sources and nesting sites for H. assimulans (Xerces
Society 2009, p. 21; 76 FR 55170, September 6, 2011, p. 55184). Habitat
modification and destruction by nonnative animals such as feral pigs,
goats, axis deer, and cattle is considered one of the primary factors
underlying destruction of native vegetation in the Hawaiian Islands,
and these habitat changes also remove food sources and nesting sites of
H. assimulans (Stone 1985, pp. 262-263; Cuddihy and Stone 1990, pp. 60-
66, 73). Fire is a threat to H. assimulans, as it destroys native plant
communities on which the species depends, and opens habitat for
increased invasion by nonnative plants. Because of the greater
frequency, intensity, and duration of fires that have resulted from the
human alteration of landscapes and the introduction of nonnative
plants, especially grasses, fires are now more destructive to the
coastal and lowland dry ecosystems (Brown and Smith 2000, p. 172), and
a single grass-fueled fire often kills most native trees and shrubs in
the area (D'Antonio and Vitousek 1992, p. 74) and could destroy food
and nesting resources for H. assimulans. The numbers of wildfires and
the acreages involved are increasing in the main Hawaiian Islands;
however, their occurrences and locations are unpredictable, and could
affect habitat for yellow-faced bees at any time (Gima 1998, in litt.;
County of Maui 2009, ch. 3, p. 3; Hamilton 2009, in litt.; Honolulu
Advertiser 2010, in litt.; Pacific Disaster Center 2011, in litt.).
Random, naturally occurring events such as hurricanes, tsunami, and
drought modify and destroy habitat of H. assimulans by creating
disturbed areas conducive to invasion by nonnative plants, eliminating
food and nesting sources (Kitayama and Mueller-Dombois 1995, p. 671;
Businger 1998, pp. 1-2; Magnacca 2015, in litt.). Predation by
nonnative ants (the big-headed ant, the yellow crazy ant, Solenopsis
papuana, and S. geminata) on Hylaeus egg, larvae, and pupal stages is a
threat to H. assimulans; additionally, ants compete with H. assimulans
for their nectar food source (Howarth 1985, p. 155; Hopper et al. 1996,
p. 9; Holway et al. 2002, pp. 188, 209; Daly and Magnacca 2003, p. 9;
Lach 2008, p. 155). Predation by nonnative western yellow jacket wasps
is a threat to H. assimulans because the wasp is an aggressive,
generalist predator, and occurs in great numbers in many habitat types,
from sea level to over 8,000 ft (2,450 m), including areas where H.
assimulans and other yellow-faced bees occur (Gambino et al. 1987, p.
169). Existing regulatory mechanisms and agency policies do not address
the primary threats to the yellow-faced bees and their habitat from
nonnative ungulates. Competition with nonnative bees (honeybees,
carpenter bees, sweat bees, and alien Hylaeus bees) for nectar and
pollen is a threat to H. assimulans (Magnacca 2007, p. 188; Graham
2015, in litt; Magnacca 2015, in litt.). The small number of
populations and individuals of H. assimulans makes this species more
vulnerable to extinction because of the higher risks from genetic
bottlenecks, random demographic fluctuations, and localized
catastrophes such as hurricanes and drought (Daly and Magnacca 2003, p.
3; Magnacca 2007, p. 173). Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to H. assimulans described
above.
The remaining populations of Hylaeus assimulans and its habitat are
at risk. The known individuals are restricted to 5 locations: 2 on
Maui, 1 on Kahoolawe, and 2 on Lanai, and is likely extirpated from
Oahu. This species continues to be negatively affected by habitat
destruction and modification by urbanization and land-use conversion,
and by habitat destruction and removal of food and nesting sites by
nonnative ungulates and nonnative plants. Habitat destruction by fire
is a threat. Randomly occurring events such as hurricanes and drought
modify habitat and remove food and nesting sources for H. assimulans.
Predation by nonnative ants and wasps is a threat. Existing regulatory
mechanisms and agency policies do not address the primary threats to
the yellow-faced bees and their habitat from nonnative ungulates.
Competition with nonnative bees for food and nesting sites is a threat.
The small number of remaining populations limits this species' ability
to adapt to environmental changes. The effects of climate change are
likely to further exacerbate these threats. Because of these threats,
we find that H. assimulans is endangered throughout all of its range,
and, therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Yellow Faced Bee (Hylaeus facilis)
Historically, Hylaeus facilis was known from Maui, Lanai, Molokai,
and Oahu, in dry shrubland to wet forest from sea level to 3,000 ft
(1,000 m) (Gagne and Cuddihy 1999, p. 93; Daly and Magnacca 2003, pp.
81, 83). Perkins (1899, p. 77) remarked H. facilis was among the most
common and widespread Hylaeus species on Oahu, Maui, Lanai, and Molokai
(Magnacca 2007, p. 183). Although the species was collected in a wide
range of habitat types, it likely prefers dry to mesic forest and
shrubland (Magnacca 2005c, in litt., p. 2), which are increasingly rare
and patchily distributed habitats (Smith 1985, pp. 227-233; Juvik and
Juvik 1998, p. 124; Gagne and Cuddihy 1999, pp. 66-67, 75; Magnacca
2005c, in litt., p. 2). Researchers believe the wet forest site on Oahu
where H. facilis was observed likely had a more open understory (more
mesic conditions), and represented an outlier or residual population
(Perkins 1899, p.76; Liebherr and Polhemus 1997; p. 347). Hylaeus
facilis has almost entirely disappeared from most of its historical
range (Maui, coastal and lowland mesic; Lanai, lowland dry and lowland
mesic; and Oahu, coastal and lowland dry) (Daly and Magnacca 2003, p.
7; Magnacca 2007, p. 183). Between 1998 and 2006, 39 sites on Maui,
Lanai, Molokai, and Oahu were surveyed, including 13 historical sites.
Hylaeus facilis was absent from all 13 locations (Magnacca 2007, p.
183) and was not observed at 26 additional sites with potentially
suitable habitat (Daly and Magnacca 2003, pp. 7, 81-82; Magnacca 2007,
p. 183). Likely extirpated from Lanai, H. facilis is currently known
from only two locations, one on Molokai in the coastal ecosystem, and
one on Oahu in the coastal ecosystem (Daly and Magnacca 2003, pp. 81-
82; Magnacca 2005c, in litt., p. 2). In addition, in 1990, a single
individual was collected on Maui near Makawao at 1,500 ft (460 m);
however, this site is urbanized and devoid of native plants, and it is
likely this collection was a vagrant individual.
Habitat destruction and modification by urbanization and land use
conversion lead to fragmentation of, and eventual loss of, foraging and
nesting areas used by Hylaeus facilis. Habitat destruction and
modification by nonnative plants adversely impacts native plant species
by modifying the availability of light, altering soil-water regimes,
modifying nutrient cycling, altering the fire characteristics, and
ultimately converting native dominated plant communities to nonnative
plant communities, and results in removal of
[[Page 67814]]
food sources and nesting sites for the H. facilis. In addition to the
nonnative plant species noted above that modify and destroy habitat of
H. assimulans, Urochloa mutica, Prosopis pallida, Psidium cattleianum,
and Rubus spp. are noted to negatively affect the habitat of H. facilis
(Cuddihy and Stone 1990, p. 105; Hawaii Division of Forestry and
Wildlife (DOFAW) 2007, pp. 20-22). Habitat modification and destruction
by nonnative animals such as feral pigs, goats, axis deer, and cattle
is considered one of the primary factors underlying destruction of
native vegetation in the Hawaiian Islands, and these habitat changes
also remove food sources and nesting sites for H. facilis (Stone 1985,
pp. 262-263; Cuddihy and Stone 1990, pp. 60-66, 73). Fire is a threat
to H. facilis, as it destroys native plant communities on which the
species depends, and opens habitat for increased invasion by nonnative
plants. Because of the greater frequency, intensity, and duration of
fires that have resulted from the human alteration of landscapes and
the introduction of nonnative plants, especially grasses, fires are now
more destructive to the coastal and lowland dry ecosystems (Brown and
Smith 2000, p. 172), and a single grass-fueled fire often kills most
native trees and shrubs in the area (D'Antonio and Vitousek 1992, p.
74) and could destroy food and nesting resources for H. facilis. The
numbers of wildfires and the acreages involved are increasing in the
main Hawaiian Islands; however, their occurrences and locations are
unpredictable, and could affect habitat for yellow-faced bees at any
time (Gima 1998, in litt.; County of Maui 2009, ch. 3, p. 3; Hamilton
2009, in litt.; Honolulu Advertiser 2010, in litt.; Pacific Disaster
Center 2011, in litt.). Random, naturally occurring events such as
hurricanes, tsunami, and drought modify and destroy habitat of H.
facilis by creating disturbed areas conducive to invasion by nonnative
plants, eliminating food and nesting resources (Kitayama and Mueller-
Dombois 1995, p. 671; Businger 1998, pp. 1-2; Magnacca 2015, in litt.).
Predation by nonnative ants (the big-headed ant, the yellow crazy ant,
Solenopsis papuana, and S. geminata) on Hylaeus egg, larvae, and pupal
stages is a threat to H. facilis; additionally, ants compete with H.
facilis for their nectar food source (Howarth 1985, p. 155; Hopper et
al. 1996, p. 9; Holway et al. 2002, pp. 188, 209; Daly and Magnacca
2003, p. 9; Lach 2008, p. 155). Predation by nonnative western yellow
jacket wasps is a threat to H. facilis because the wasp is an
aggressive, generalist predator, and occurs in great numbers in many
habitat types, from sea level to over 8,000 ft (2,450 m), including
areas where H. facilis and other yellow-faced bees occur (Gambino et
al. 1987, p. 169). Existing regulatory mechanisms and agency policies
do not address the primary threats to the yellow-faced bees and their
habitat from nonnative ungulates. Competition with nonnative bees
(honeybees, carpenter bees, sweat bees, and alien Hylaeus bees) for
nectar and pollen is a threat to H. facilis (Magnacca 2007, p. 188;
Magnacca 2015, in litt.). The small number of populations and
individuals of H. facilis makes this species more vulnerable to
extinction because of the higher risks from genetic bottlenecks, random
demographic fluctuations, and localized catastrophes such as hurricanes
and drought (Daly and Magnacca 2003, p. 3; Magnacca 2007, p. 173).
Although we cannot predict the timing, extent, or magnitude of specific
impacts, we do expect the effects of climate change to exacerbate the
threats to H. facilis described above.
The remaining populations of Hylaeus facilis and its habitat are at
risk. The known individuals are restricted to one location on Molokai
and one location on Oahu, and continue to be negatively affected by
habitat destruction and modification by urbanization and land-use
conversion, and by habitat destruction and removal of food and nesting
sites by nonnative ungulates and nonnative plants. Habitat destruction
by fire is a threat. Randomly occurring events such as hurricanes and
drought modify habitat and remove food and nesting sources for H.
facilis. Predation by nonnative ants and wasps is a threat. Existing
regulatory mechanisms and agency policies do not address the primary
threats to the yellow-faced bees and their habitat from nonnative
ungulates. Competition with nonnative bees for food and nesting sites
is a threat. The small number of remaining populations limits this
species' ability to adapt to environmental changes. The effects of
climate change are likely to further exacerbate these threats. Because
of these threats, we find that H. facilis is endangered throughout all
of its range, and, therefore, find that it is unnecessary to analyze
whether it is endangered or threatened in a significant portion of its
range.
Yellow-Faced Bee (Hylaeus hilaris)
Historically, Hylaeus hilaris was known from coastal habitat on
Maui, Lanai, and Molokai; and lowland dry habitat on Maui. It is
believed to have occurred along much of the coast of these islands
because its primary hosts, H. anthracinus, H. assimulans, and H.
longiceps likely occurred throughout this habitat. First collected on
Maui in 1879, H. hilaris has only been collected twice in the last 100
years. Hylaeus hilaris was absent from three of its historical
population sites revisited by researchers between 1998 and 2006
(Magnacca 2007, p. 181). It was also not observed in 2003 at 10
additional sites with potentially suitable habitat (Daly and Magnacca
2003, pp. 103, 106). Currently, the only known population of H. hilaris
is located on Molokai, in the coastal ecosystem (Daly and Magnacca
2003, pp. 103, 106; Magnacca 2005d, in litt., p. 2; Magnacca 2007, p.
181).
Because Hylaeus hilaris is an obligate parasite on H. anthracinus,
H. assimulans, and H. longiceps, its occurrences are determined by the
remaining populations of these three species. Habitat destruction and
modification by urbanization and land use conversion leads to
fragmentation of, and eventual loss of, foraging and nesting areas of
H. hilaris, and of those Hylaeus species that H. hilaris is dependent
upon. Habitat destruction and modification by nonnative plants
adversely impacts native plant species by modifying the availability of
light, altering soil-water regimes, modifying nutrient cycling,
altering the fire characteristics, and ultimately converting native
dominated plant communities to nonnative plant communities, and results
in removal of food sources and nesting sites for the Hylaeus species
that H. hilaris is dependent upon. Nonnative plant species that modify
and destroy habitat of H. hilaris are noted in the description for H.
assimulans, above. Habitat modification and destruction by nonnative
animals such as feral pigs, goats, axis deer, and cattle is considered
one of the primary factors underlying destruction of native vegetation
in the Hawaiian Islands, and these habitat changes also remove food
sources and nesting sites for the host species of H. hilaris (Stone
1985, pp. 262-263; Cuddihy and Stone 1990, pp. 60-66, 73). Fire is a
threat to H. hilaris, as it destroys native plant communities, and
opens habitat for increased invasion by nonnative plants. Because of
the greater frequency, intensity, and duration of fires that have
resulted from the human alteration of landscapes and the introduction
of nonnative plants, especially grasses, fires are now more destructive
to the coastal and lowland dry ecosystems (Brown and Smith 2000, p.
172), and a single grass-fueled fire often kills most native trees and
shrubs in the area (D'Antonio and Vitousek
[[Page 67815]]
1992, p. 74) and could destroy food and nesting resources for H.
hilaris. The numbers of wildfires and the acreages involved are
increasing in the main Hawaiian Islands; however, their occurrences and
locations are unpredictable, and could affect habitat for yellow-faced
bees at any time (Gima 1998, in litt.; County of Maui 2009, ch. 3, p.
3; Hamilton 2009, in litt.; Honolulu Advertiser 2010, in litt.; Pacific
Disaster Center 2011, in litt.). Random, naturally occurring events
such as hurricanes, tsunami, and drought can modify and destroy habitat
of H. hilaris by creating disturbed areas conducive to invasion by
nonnative plants, eliminating food and nesting sources of its host
species (Kitayama and Mueller-Dombois 1995, p. 671; Businger 1998, pp.
1-2; Magnacca 2015, in litt.). Predation by nonnative ants (the big-
headed ant, the long-legged ant, Solenopsis papuana, and S. geminata)
on Hylaeus egg, larvae, and pupal stages is a threat to H. hilaris;
additionally, ants compete with the yellow-faced bees that H. hilaris
is dependent on for their food resources (Howarth 1985, p. 155; Hopper
et al. 1996, p. 9; Holway et al. 2002, pp. 188, 209; Daly and Magnacca
2003, p. 9; Lach 2008, p. 155). Predation by nonnative western yellow
jacket wasps is a threat to H. hilaris because the wasp is an
aggressive, generalist predator, and occurs in great numbers in many
habitat types, from sea level to over 8,000 ft (2,450 m), including
areas where H. hilaris and other yellow-faced bees occur (Gambino et
al. 1987, p. 169). Existing regulatory mechanisms and agency policies
do not address the primary threats to the yellow-faced bees and their
habitat from nonnative ungulates. Competition with nonnative bees
(honeybees, carpenter bees, sweat bees, and alien Hylaeus bees) for
nectar and pollen is a threat to the host yellow-faced bees of H.
hilaris (Magnacca 2007, p. 188; Graham 2015, in litt.; Magnacca 2015,
in litt.). The small number of populations and individuals of H.
hilaris makes this species more vulnerable to extinction because of the
higher risks from genetic bottlenecks, random demographic fluctuations,
and localized catastrophes such as hurricanes and drought (Daly and
Magnacca 2003, p. 3; Magnacca 2007, p. 173). Although we cannot predict
the timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to H. hilaris
described above.
The remaining populations of Hylaeus hilaris and its habitat are at
risk. There is one known occurrence on Molokai. Hylaeus hilaris and its
host species continue to be negatively affected by habitat destruction
and modification by urbanization and land-use conversion, and by
habitat destruction and removal of food and nesting sites (for host
species) by nonnative ungulates and nonnative plants. Habitat
destruction by fire is a threat. Randomly occurring events such as
hurricanes and drought modify habitat and remove food and nesting
sources for H. hilaris and its host species. Predation by nonnative
ants and wasps is a threat. Existing regulatory mechanisms and agency
policies do not address the primary threats to the yellow-faced bees
and their habitat from nonnative ungulates. Competition with nonnative
bees for food and nesting sites is a threat. The small number of
remaining populations limits this species' ability to adapt to
environmental changes, especially because it is an obligate parasite of
other rare Hylaeus bees. Because of these threats, we find that H.
hilaris is endangered throughout all of its range, and, therefore, find
that it is unnecessary to analyze whether it is endangered or
threatened in a significant portion of its range.
Yellow-Faced Bee (Hylaeus kuakea)
Because the first collection of Hylaeus kuakea was not made until
1997, its historical range is unknown (Magnacca 2005e, in litt., p. 2;
Magnacca 2007, p. 184). Phylogenetically, H. kuakea belongs in a
species-group primarily including species inhabiting mesic forests
(Magnacca and Danforth 2006, p. 405). Only four individuals (all males)
have been collected from two different sites in the Waianae Mountains
of Oahu in the lowland mesic ecosystem (Magnacca 2007, p. 184). The
species has never been collected in any other habitat type or area,
including some sites that have been more thoroughly surveyed (Magnacca
2011, in litt.). Not all potentially suitable habitat has been surveyed
due to the remote and rugged locations, small size, rareness, and
distant spacing among large areas of nonnative forest (Smith 1985, pp.
227-233; Juvik and Juvik 1998, p. 124; Wagner et al. 1999, pp. 66-67,
75).
Habitat destruction and modification by feral pigs leads to
fragmentation of, and eventual loss of, foraging and nesting areas of
Hylaeus kuakea. Habitat destruction and modification by nonnative
plants adversely impacts native plant species by modifying the
availability of light, altering soil-water regimes, modifying nutrient
cycling, altering the fire characteristics, and ultimately converting
native dominated plant communities to nonnative plant communities, and
results in removal of food sources and nesting sites for H. kuakea.
Nonnative plant species that modify and destroy habitat of H. kuakea
are noted in the descriptions for H. assimulans and H. facilis, above.
Fire is a threat to H. kuakea because it destroys native plant
communities and opens habitat for increased invasion by nonnative
plants. Because of the greater frequency, intensity, and duration of
fires that have resulted from the human alteration of landscapes and
the introduction of nonnative plants, especially grasses, fires are now
more destructive, including in lowland mesic areas (Brown and Smith
2000, p. 172), and a single grass-fueled fire often kills most native
trees and shrubs in the area (D'Antonio and Vitousek 1992, p. 74) and
could destroy food and nesting resources for H. kuakea. The numbers of
wildfires and the acreages involved are increasing in the main Hawaiian
Islands; however, their occurrences and locations are unpredictable,
and could affect habitat for yellow-faced bees at any time (Gima 1998,
in litt.; County of Maui 2009, ch. 3, p. 3; Hamilton 2009, in litt.;
Honolulu Advertiser 2010, in litt.; Pacific Disaster Center 2011, in
litt.). Random, naturally occurring events such as hurricanes and
drought can modify and destroy habitat of H. kuakea by creating
disturbed areas conducive to invasion by nonnative plants, eliminating
food and nesting resources (Kitayama and Mueller-Dombois 1995, p. 671;
Businger 1998, pp. 1-2). Predation by nonnative ants (the big-headed
ant, the long-legged ant, Solenopsis papuana, and S. geminata) on
Hylaeus egg, larvae, and pupal stages is a threat to H. kuakea;
additionally, ants compete with H. kuakea for their nectar food source
(Howarth 1985, p. 155; Hopper et al. 1996, p. 9; Holway et al. 2002,
pp. 188, 209; Daly and Magnacca 2003, p. 9; Lach 2008, p. 155).
Predation by nonnative western yellow jacket wasps is a threat to H.
kuakea because the wasp is an aggressive, generalist predator, and
occurs in great numbers in many habitat types, from sea level to over
8,000 ft (2,450 m), including areas where H. kuakea and other yellow-
faced bees occur (Gambino et al. 1987, p. 169). Existing regulatory
mechanisms and agency policies do not address the primary threats to
the yellow-faced bees and their habitat from nonnative ungulates.
Competition with nonnative bees (honeybees, carpenter bees, sweat bees,
and alien Hylaeus bees) for nectar and pollen is a threat to H. kuakea
(Magnacca 2007, p. 188; Graham 2015, in litt.; Magnacca 2015, in
litt.). The small number of populations and individuals of H. kuakea
makes this
[[Page 67816]]
species more vulnerable to extinction because of the higher risks from
genetic bottlenecks, random demographic fluctuations, and localized
catastrophes such as hurricanes and drought (Daly and Magnacca 2003, p.
3; Magnacca 2007, p. 173). Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to H. kuakea described above.
The remaining populations of Hylaeus kuakea and its habitat are at
risk. The known individuals are restricted to mesic forest in one area
of one island (Oahu), and continue to be negatively affected by habitat
destruction and removal of food and nesting sites by nonnative
ungulates and nonnative plants. Habitat destruction by fire is a
threat. Randomly occurring events such as hurricanes and drought modify
habitat and remove food and nesting sources for H. kuakea. Predation by
nonnative ants and wasps is a threat. Existing regulatory mechanisms
and agency policies do not address the primary threats to the yellow-
faced bees and their habitat from nonnative ungulates. Competition with
nonnative bees for food and nesting sites is a threat. The small number
of remaining populations limits this species' ability to adapt to
environmental changes. The effects of climate change are likely to
further exacerbate these threats. Because of these threats, we find
that H. kuakea is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Yellow-faced bee (Hylaeus longiceps)
Hylaeus longiceps is historically known from coastal and lowland
dry shrubland habitat up to 2,000 ft (610 m) in numerous locations on
the islands of Maui, Lanai, Molokai, and Oahu. Perkins (1899, p. 98)
noted H. longiceps was locally abundant, and probably occurred
throughout much of the leeward and lowland areas on these islands.
Hylaeus longiceps is now restricted to small populations in patches of
coastal and lowland dry habitat on the Maui, Lanai, Molokai, and Oahu
(Magnacca 2005f, in litt., p. 2; Magnacca and King 2013, pp. 13, 16).
Twenty-five sites that were either historical collecting localities or
contained potentially suitable habitat for this species were surveyed
between 1997 and 2008 (Magnacca and King 2013, p. 16). Hylaeus
longiceps was observed at only seven of the surveyed sites: Three sites
on Lanai (in the coastal and lowland dry ecosystems), two sites on Oahu
(in the coastal ecosystem), and one site on each of the islands of Maui
(in the coastal ecosystem) and Molokai (in the coastal ecosystem) (Daly
and Magnacca 2003, p. 135; Magnacca and King 2013, pp. 11-12).
Most of the coastal and lowland habitat of Hylaeus longiceps has
been developed or degraded, and is no longer suitable (Liebherr and
Polhemus 1997, pp. 346-347; Magnacca 2007, pp. 186-188). Habitat
destruction and modification by axis deer (Lanai) and urbanization
(Maui and Molokai) leads to fragmentation of, and eventual loss of,
foraging and nesting areas of H. longiceps (Daly and Magnacca 2003, pp.
217-229). Habitat modification and destruction by human impacts in
areas accessible by four-wheel drive vehicles on Lanai is a threat
because these vehicles can destroy plants used as food sources and
destroy ground nesting sites for H. longiceps (Daly and Magnacca 2003,
p. 135). Habitat destruction and modification by nonnative plants
adversely affects native plant species used by H. longiceps as a food
source by modifying the availability of light, altering soil-water
regimes, modifying nutrient cycling, altering the fire characteristics,
and ultimately converting native-dominated plant communities to
nonnative plant communities. Nonnative plant species that modify and
destroy habitat of H. longiceps are noted in the descriptions for H.
assimulans and H. facilis, above. Fire is a threat to H. longiceps
because it destroys native plant communities, and opens habitat for
increased invasion by nonnative plants. Because of the greater
frequency, intensity, and duration of fires that have resulted from the
human alteration of landscapes and the introduction of nonnative
plants, especially grasses, fires are now more destructive to the
coastal and lowland dry ecosystems (Brown and Smith 2000, p. 172), and
a single grass-fueled fire often kills most native trees and shrubs in
the area (D'Antonio and Vitousek 1992, p. 74) and could destroy food
and nesting resources for H. longiceps. The numbers of wildfires and
the acreages involved are increasing in the main Hawaiian Islands;
however, their occurrences and locations are unpredictable, and could
affect habitat for yellow-faced bees at any time (Gima 1998, in litt.;
County of Maui 2009, ch. 3, p. 3; Hamilton 2009, in litt.; Honolulu
Advertiser 2010, in litt.; Pacific Disaster Center 2011, in litt.).
Random, naturally occurring events such as hurricanes, tsunami, and
drought modify and destroy habitat of H. longiceps by creating
disturbed areas conducive to invasion by nonnative plants, eliminating
food and nesting resources (Kitayama and Mueller-Dombois 1995, p. 671;
Businger 1998, pp. 1-2; Magnacca 2015, in litt.). Predation by, and
competition for food sources, by nonnative ants and the nonnative
western yellow jacket wasp is a threat to H. longiceps (see H. kuakea,
above) (Gambino et al. 1987, p. 169; Howarth 1985, p. 155; Hopper et
al. 1996, p. 9; Holway et al. 2002, pp. 188, 209; Daly and Magnacca
2003, p. 9; Lach 2008, p. 155). Existing regulatory mechanisms and
agency policies do not address the primary threats to the yellow-faced
bees and their habitat from nonnative ungulates. Competition with
nonnative bees (honeybees, carpenter bees, sweat bees, and alien
Hylaeus bees) for nectar and pollen is a threat to H. longiceps
(Magnacca 2007, p. 188; Graham 2015, in litt.; Magnacca 2015, in
litt.). The small number of populations and individuals of H. longiceps
makes this species more vulnerable to extinction because of the higher
risks from genetic bottlenecks, random demographic fluctuations, and
localized catastrophes such as hurricanes and drought (Daly and
Magnacca 2003, p. 3; Magnacca 2007, p. 173). Although we cannot predict
the timing, extent, or magnitude of specific impacts, we do expect the
effects of climate change to exacerbate the threats to H. longiceps
described above.
The remaining population of Hylaeus longiceps and its habitat are
at risk. The known individuals are restricted to seven locations, three
on Lanai, two on Oahu, and one each on Maui and Molokai, and continue
to be negatively affected by habitat destruction and modification by
urbanization and land-use conversion, by habitat destruction and
removal of food and nesting sites by nonnative ungulates and nonnative
plants, and by recreational use vehicles on Lanai. Habitat destruction
by fire is a threat. Randomly occurring events such as hurricanes and
drought may modify habitat and remove food and nesting sources for H.
longiceps. Predation by nonnative ants and wasps is a threat. Existing
regulatory mechanisms and agency policies do not address the primary
threats to the yellow-faced bees and their habitat from nonnative
ungulates. Competition with nonnative bees for food and nesting sites
is a threat. The small number of remaining populations limits this
species' ability to adapt to environmental changes. Because of these
threats, we find that H. longiceps is endangered throughout all of its
range, and, therefore, find that it is
[[Page 67817]]
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Yellow-Faced Bee (Hylaeus mana)
Hylaeus mana is known only from lowland mesic forest dominated by
native Acacia koa in the Koolau Mountains of Oahu, at 1,400 ft (430 m).
Few other Hylaeus species have been found in this type of forest on
Oahu (Daly and Magnacca 2003, p. 138). This type of native forest is
increasingly rare and patchily distributed because of competition and
encroachment into habitat by nonnative plants (Smith 1985, pp. 227-233;
Juvik and Juvik 1998, p. 124; Wagner et al. 1999, pp. 66-67, 75).
Decline of this forest type could lead to decline in populations and
numbers of H. mana. Three additional population sites were discovered
on Oahu in 2012, including a new observation of the species at the
original site (Magnacca and King 2013, pp. 17-18). The three new sites
are within a narrow range of lowland mesic forest at 1,400 ft (430 m),
bordered by nonnative plant habitat at lower elevations and wetter
native forest habitat above (Magnacca and King 2013, pp. 17-18).
Hylaeus mana was most often observed on Santalum freycinetianum var.
freycinetianum, which suggests that H. mana may be closely associated
with this plant species (Magnacca and King 2013, p. 18). Additional
surveys may reveal more populations; however, the extreme rarity of
this species, its absence from many survey sites, the fact that it was
not discovered until very recently, and the limited range of its
possible host plant, all suggest that few populations remain (Magnacca
2005g, in litt., p. 2; Magnacca and King 2013, pp. 17-18).
Habitat destruction and modification by feral pigs leads to
fragmentation of, and eventual loss of, foraging and nesting areas of
Hylaeus mana (Daly and Magnacca 2003, pp. 217-229). Habitat destruction
and modification by nonnative plants adversely impacts native plant
species used by H. mana as a food source by modifying the availability
of light, altering soil-water regimes, modifying nutrient cycling,
altering the fire characteristics, and ultimately converting native
dominated plant communities to nonnative plant communities. Nonnative
plant species that modify and destroy habitat of H. mana are noted in
the descriptions for H. assimulans and H. facilis, above, and can
outcompete native canopy species such as Acacia koa, the known
preferred native canopy type of H. mana (GISD 2011, in litt.; State of
Hawaii 2013, in litt. (S.C.R. No. 74)). Fire is a threat to H. mana, as
it destroys native plant communities on which the species depends, and
opens habitat for increased invasion by nonnative plants. Because of
the greater frequency, intensity, and duration of fires that have
resulted from the human alteration of landscapes and the introduction
of nonnative plants, especially grasses, fires are now more
destructive, including in lowland mesic ecosystems (Brown and Smith
2000, p. 172). A single grass-fueled fire often kills most native trees
and shrubs in the area (D'Antonio and Vitousek 1992, p. 74) and could
destroy food and nesting resources for H. mana. The numbers of
wildfires and the acreages involved are increasing in the main Hawaiian
Islands; however, their occurrences and locations are unpredictable,
and could affect habitat for yellow-faced bees at any time (Gima 1998,
in litt.; County of Maui 2009, ch. 3, p. 3; Hamilton 2009, in litt.;
Honolulu Advertiser 2010, in litt.; Pacific Disaster Center 2011, in
litt.). Random, naturally occurring events such as hurricanes and
drought can modify and destroy habitat of H. mana by creating disturbed
areas conducive to invasion by nonnative plants (Kitayama and Mueller-
Dombois 1995, p. 671; Businger 1998, pp. 1-2). Predation and
competition for food sources by nonnative ants and the nonnative
western yellow jacket wasp are threats to H. mana (see H. kuakea,
above) (Howarth 1985, p. 155; Gambino et al. 1987, p. 169; Hopper et
al. 1996, p. 9; Holway et al. 2002, pp. 188, 209; Daly and Magnacca
2003, p. 9; Lach 2008, p. 155). Existing regulatory mechanisms and
agency policies do not address the primary threats to the yellow-faced
bees and their habitat from nonnative ungulates. Competition with
nonnative bees (honeybees, carpenter bees, sweat bees, and alien
Hylaeus bees) for nectar and pollen is a threat to H. mana (Magnacca
2007, p. 188; Graham 2015, in litt.; Magnacca 2015, in litt.). The
small number of populations and individuals of H. mana makes this
species more vulnerable to extinction because of the higher risks from
genetic bottlenecks, random demographic fluctuations, and localized
catastrophes such as hurricanes and drought (Daly and Magnacca 2003, p.
3; Magnacca 2007, p. 173). Although we cannot predict the timing,
extent, or magnitude of specific impacts, we do expect the effects of
climate change to exacerbate the threats to H. mana described above.
The remaining populations of Hylaeus mana and its habitat are at
risk. The known individuals are restricted to three locations of native
koa forest on Oahu, and continue to be negatively affected by habitat
destruction and removal of food and nesting sites by nonnative
ungulates and nonnative plants. Habitat destruction by fire is a
threat. Randomly occurring events such as hurricanes and drought may
modify habitat and remove food and nesting sources for H. mana.
Predation by nonnative ants and wasps is a threat. Existing regulatory
mechanisms and agency policies do not address the primary threats to
the yellow-faced bees and their habitat from nonnative ungulates.
Competition with nonnative bees for food and nesting sites is a threat.
The small number of remaining populations limits this species' ability
to adapt to environmental changes. The effects of climate change are
likely to further exacerbate these threats. Because of these threats,
we find that H. mana is endangered throughout all of its range, and,
therefore, find that it is unnecessary to analyze whether it is
endangered or threatened in a significant portion of its range.
Orangeblack Hawaiian Damselfly (Megalagrion xanthomelas)
The orangeblack Hawaiian damselfly was once Hawaii's most abundant
damselfly species likely because of its ability to use a variety of
aquatic habitats for breeding sites. Historically, the orangeblack
Hawaiian damselfly probably occurred on all of the main Hawaiian
Islands (except Kahoolawe) in suitable aquatic habitat within the
anchialine pool, coastal, lowland dry, and lowland mesic ecosystems
(Perkins 1913, p. clxxviii; Zimmerman 1948, p. 379; Polhemus 1996, p.
30). Its historical range on Kauai is unknown. On Oahu, it was recorded
from Honolulu, Kaimuki, Koko Head, Pearl City, Waialua, the Waianae
Mountains, and Waianae (Polhemus 1996, pp. 31, 33). On Molokai, it was
known from Kainalu, Meyer's Lake (Kalaupapa Peninsula), Kaunakakai,
Mapulehu, and Palaau (Polhemus 1996, pp. 33-41). On Lanai, small
populations occurred on Maunalei Gulch, and in ephemeral coastal ponds
at the mouth of Maunalei Gulch drainage, at Keomuku, and in a
mixohaline (brackish water) habitat at Lopa (Polhemus 1996, pp. 37-41;
HBMP 2010). On Maui, this species was recorded from an unspecified
locality in the west Maui Mountains (Polhemus 1996, pp. 41-42; Polhemus
et al. 1999, pp. 27-29). On Hawaii Island, it was known from Hilo,
Kona, and Naalehu (Polhemus 1996, pp. 42-47).
Currently, the orangeblack Hawaiian damselfly occurs on Oahu,
Molokai, Lanai, Maui, and Hawaii Island. In 1994, on Oahu, a very small
population
[[Page 67818]]
was discovered in pools of an intermittent stream (Englund 2001, p.
256). On Molokai, populations occur at the mouths of two streams, and
in wetlands on the south coast (Polhemus 1996, p. 47). On Lanai, a
large population occurs in an artificial pond (Polhemus 1996, p. 47).
The species is present on west Maui at a stream and near anchialine
pools on east Maui (Polhemus et al. 1999, p. 29). Several large
populations exist in coastal wetlands on Hawaii Island at 14 locations
(Polhemus 1996, pp. 42-47; Orlando 2015, in litt.). The species is
believed to be extirpated from Kauai (Asquith and Polhemus 1996, p.
91).
Past and present land use and water management practices, including
agriculture, urban development, ground water development, and
destruction of perched aquifer and surface water resources, and feral
ungulates (pigs, goats, axis deer), modify and destroy habitat of the
orangeblack Hawaiian damselfly (Harris et al. 1993, pp. 9-13; Meier et
al. 1993, pp. 181-183). Nonnative plant species such as Urochloa mutica
form dense, monotypic stands that can completely eliminate any open
water habitat of the orangeblack Hawaiian damselfly (Smith 1985, p.
186). Stochastic events such as drought, flooding, and hurricanes can
also modify and destroy habitat, and kill individuals. Predation of the
orangeblack Hawaiian damselfly by nonnative fish and nonnative aquatic
invertebrates on the orangeblack Hawaiian damselfly is a significant
threat; predation by Jackson's chameleons (Trioceros jacksonii) may
occur as well (Sailer 2015, in litt.). Hawaiian damselflies evolved
with few, if any, predatory fish, and the reduced defensive and evasive
behaviors of most of the fully aquatic species, including the
orangeblack Hawaiian damselfly, makes them particularly vulnerable to
predation by nonnative fish (Englund 1999, pp. 225-225, 235; Haines
2015, in litt.). The damselfly is not observed in any bodies of water
that support nonnative fish (Henrickson 1988, p. 183; McPeek 1990a, pp.
92-96). Nonnative backswimmers (aquatic true bugs; Heteroptera) are
voracious predators and frequently feed on prey much larger than
themselves, such as tadpoles, small fish, and other aquatic
invertebrates and may be a potential threat to damselfly's aquatic
larvae (naiads) (Borror et al. 1989, p. 296). In addition, the
nonnative bullfrog (Rana catesbeiana, Lithobates catesbeianus), found
in ponds and along streams, is a generalist predator, and eats insects
and crustaceans as well as a wide variety of small vertebrates (Bury
and Whelan 1985, p. 4). Predation by the bullfrog is a threat to the
orangeblack Hawaiian damselfly (Englund et al. 2007, pp. 215, 219;
Haines 2015, in litt.). Also, caddisflies (Trichoptera spp.) compete
with native aquatic invertebrates for resources and space (Flint et al.
2003, p. 38; Haines 2015, in litt.) and reduce prey abundance for
orangeblack Hawaiian damselfly larvae.
Hawaii State law (State Water Code) does not provide for permanent
or minimal instream flow standards, and channel modifications or
revisions to flow standards can be undertaken at any time by the Water
Commission, without regard for changes that degrade or destroy habitat,
food resources, or aquatic life stages of the orangeblack Hawaiian
damselfly. Therefore, existing regulatory mechansims do not adequately
address the threat of modification and destruction of the aquatic
habitat of the orangeblack Hawaiian damselfly (Hawaii Administrative
Rule (HAR)-State Water Code, title 13, chapter 169-36; Tango 2010, in
litt.).
The remaining populations and habitat of the orangeblack Hawaiian
damselfly are at risk; numbers are decreasing on Oahu, Molokai, Lanai,
Maui, and Hawaii Island, and both the species and its habitat continue
to be negatively affected by modification and destruction by
development and water management practices, drought, feral ungulates,
and by nonnative plants, combined with predation by nonnative fish and
other nonnative vertebrates. Competition with caddisflies is a
potential threat to the orangeblack Hawaiian damselfly. The orangeblack
damselfly was once the most common Hawaiian damselfly in the State, and
occurred in any suitable aquatic habitat. Populations no longer occur
on Kauai. The Oahu populations were described from seven locations, and
this species now only occurs at one location. The populations on
Molokai have declined from five to three. Populations on Lanai have
declined from four to one in an artificial pond. On Maui, there are
only two populations, one on east Maui, and one on west Maui. Of the 21
known populations on Hawaii Island, only 14 remain. Because of the
dramatic decline in numbers and populations, and because of the ongoing
threats described above, we find that this species is endangered
throughout all of its range, and, therefore, find that it is
unnecessary to analyze whether it is endangered or threatened in a
significant portion of its range.
Anchialine Pool Shrimp (Procaris hawaiana)
The shrimp family Procarididae is represented by a small number of
species globally, with only two species within the genus Procaris
(Magnacca 2015, in litt.). Procaris hawaiana is an endemic anchialine
pool shrimp species known only from the islands of Maui and Hawaii. The
second species, P. ascensionis, is restricted to similar habitat on
Ascension Island in the South Atlantic Ocean. Of the anchialine pools
on Hawaii Island, only 25 are known to contain P. hawaiana. During
nocturnal-diurnal surveys conducted from 2009 to 2010, 19 pools within
Manuka NAR were found to contain P. hawaiana. Five additional pools
located on unencumbered State land adjacent to Manuka NAR also
contained P. hawaiana. An additional separate pool also contains P.
hawaiana, along with the endangered anchialine pool shrimp Vetericaris
chaceorum (Holthuis 1973, pp. 12-19; Maciolek 1983, pp. 607-614; Brock
2004, pp. 30-57). On Maui, P. hawaiana occurs in two anchialine pools
(Holthuis 1973, pp. 12-19; Maciolek 1983, pp. 607-614; Brock 2004, pp.
30-57).
Like other anchialine pool shrimp species, Procaris hawaiana
inhabits extensive networks of water-filled interstitial spaces (cracks
and crevices) leading to and from the open pools where they can be
detected, a trait which has precluded accurate estimates of population
size (Holthuis 1973, p. 36; Maciolek 1983, pp. 613-616). Surveys for
many rare species of anchialine pool shrimp, including P. hawaiana,
often involve baiting in likely habitat to determine presence or
absence. Absence, and presumably extirpation, of shrimp species from
suitable habitat is the best or only measure of species decline as
population sizes are not easily determined or monitored (Holthuis 1973,
pp. 7-12; Maciolek 1983, pp. 613-616), but owing to the potential for
shrimp to move between pools through subterranean connections, the lack
of sighting on one or several visits to a site is not definitive
evidence that the species is extirpated (Kinzie 2015, in litt.).
Extirpation of anchialine pool shrimp has been documented definitively
in some cases; for example, Halocaridina rubra disappeared from an
anchialine pool at Honokohau Harbor (Hawaii Island) as a result of the
use of the pool for dumping of used oil, grease, and oil filters (Brock
2004, p. 14). To date, however, P. hawaiana is not known to have been
extirpated from any of the pools where
[[Page 67819]]
it has been documented to occur (Wada 2016, in litt.).
Habitat modification and destruction by human activities is a
significant threat to Procaris hawaiana. It is estimated that up to 90
percent of existing anchialine pools in Hawaii have been destroyed by
filling and bulldozing (Baily-Brock and Brock 1993, p. 354; Brock 2004,
p. i). Anchialine pools are used as dumping pits for bottles, cans, and
used oil and grease, and these activities are a known cause of the
disappearance of other anchialine pool shrimp species from the pools.
Trampling damage from use of anchialine pools for swimming and bathing
has been documented (Brock 2004, pp. 13-17). Although a permit from the
State is required to collect anchialine pool shrimp, unpermitted
collection of shrimp is ongoing (Fuku-Bonsai 2015, in litt.). A single
person with a handnet could do irreparable damage to a population of P.
hawaiana (Yamamoto 2015, in litt.), but collection by permitted
individuals is not prohibited at State Parks or City and County
property where some anchialine pools occur. Predation by nonnative fish
is a direct threat to P. hawaiana. Nonnative fish (tilapia, Oreochromis
mossambica) also outcompete native herbivorous species of shrimp that
serve as a prey-base for P. hawaiana, disrupting the delicate
ecological balance in the anchialine pool system, and leading to
decline of the pools and the shrimp inhabiting them (Brock 2004, pp.
13-17). Although anchialine pools within State of Hawaii NARs are
provided some protection, these areas are remote and signage does not
prevent human use and damage of the pools (see Factor B). The
persistence of P. hawaiana is hampered by the small number of extant
populations and the small geographic range of the known populations.
The populations of P. hawaiana are at risk of extinction because of
their increased vulnerability to loss of individuals from disturbance,
habitat destruction, and the effects of invasive species and because of
the reduction in genetic variability that may make the species less
able to adapt to changes in the environment (Harmon and Braude 2010,
pp. 125-128). In addition, large-scale water extraction from
underground water sources negatively affects the habitat and P.
hawaiana directly (Conry 2012, in litt.). A threat from development
upslope of anchialine pool habitat is infiltration of waste water or
application of fertilizer and pesticides that may enter the ground
water system of the anchialine pools and consequently affect the pool's
ecosystem health, food sources of the pool shrimp, or the pool shrimp
directly (Kinzie 2015, in litt.; Yamamoto et al. 2015, pp. 75-83). Sea-
level rise and coastal inundation resulting from the effects of climate
change is a threat to P. hawaiana (Sakihara 2015, in litt.). Sea-level
rise would increase surface connectivity between isolated anchialine
pools, and exacerbate the spread of nonnative fish into pools not yet
occupied by nonnative fish (Sakihara 2015, in litt.).
Procaris hawaiana and its habitat are at risk. There are a total of
700 known anchialine pools in the State of Hawaii. Procaris hawaiana is
restricted to 25 anchialine pools out of 600 on Hawaii Island and to 2
anchialine pools on Maui. These 27 anchialine pools continue to be
negatively affected by habitat destruction and modification by human
use of the pools for bathing and for dumping of trash and nonnative
fish; filling and bulldozing; water extraction; contamination;
predation by and competition with nonnative fish; and collection for
the aquarium trade. The small number of populations (27) limits this
species' ability to adapt to environmental changes. Because of these
threats, we find that this species is endangered throughout all of its
range, and, therefore, find that it is unnecessary to analyze whether
it is endangered or threatened in a significant portion of its range.
Distinct Population Segment (DPS) Analysis
Band-Rumped Storm-Petrel (Oceanodroma castro)
Under the Act, we have the authority to consider for listing any
species, subspecies, or, for vertebrates, any distinct population
segment (DPS) of these taxa if there is sufficient information to
indicate that such action may be warranted. To guide the implementation
of the DPS provisions of the Act, we, and the National Marine Fisheries
Service (National Oceanic and Atmospheric Administration--Fisheries),
published the Policy Regarding the Recognition of Distinct Vertebrate
Population Segments Under the Endangered Species Act (DPS Policy) in
the Federal Register on February 7, 1996 (61 FR 4722). Under our DPS
Policy, we use two elements to assess whether a population segment
under consideration for listing may be recognized as a DPS: (1) The
population segment's discreteness from the remainder of the species to
which it belongs, and (2) the significance of the population segment to
the species to which it belongs. If we determine that a population
segment being considered for listing is a DPS, then the population
segment's conservation status is evaluated based on the five listing
factors established by the Act to determine if listing it as either
endangered or threatened is warranted. In the proposed rule (80 FR
58820; September 30, 2015), we evaluated the Hawaii population of the
band-rumped storm-petrel to determine whether it meets the definition
of a DPS under our DPS Policy.
Discreteness
Under the DPS Policy, a population segment of a vertebrate taxon
may be considered discrete if it satisfies either one of the following
conditions: (1) It is markedly separated from other populations of the
same taxon as a consequence of physical, physiological, ecological, or
behavioral factors (quantitative measures of genetic or morphological
discontinuity may provide evidence of this separation); or (2) it is
delimited by international governmental boundaries within which
differences in control of exploitation, management of habitat,
conservation status, or regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D) of the Act. The Hawaii
population of the band-rumped storm-petrel meets the first criterion:
it is markedly separated from other populations of this species by
physical (geographic) and physiological (genetic) factors, as described
below.
The band-rumped storm-petrel is widely distributed in the tropics
and subtropics, with breeding populations in numerous island groups in
the Atlantic and in Hawaii, Galapagos, and Japan in the Pacific
(Harrison 1983, p. 274; Carboneras et al. 2014, p. 1 and Fig. 2). The
geographic separation of these breeding populations is widely
recognized, with strong genetic differentiation between the two ocean
basins and among individual populations (Friesen et al. 2007a, p. 1768;
Smith et al. 2007, p. 768). Whether individual populations merit
taxonomic separation remains unclear, and further study is needed
(Friesen et al. 2007b, p. 18591; Smith et al. 2007, p. 770; reviewed in
Howell 2012, pp. 349, 369-370); some populations, such as those in the
Galapagos and Cape Verde islands, may warrant full species status
(Smith et al. 2007, p. 770). Like other storm-petrels, the band-rumped
storm-petrel is a highly pelagic (open-ocean) seabird (Howell 2012, p.
349). In addition, like other species in the seabird order
Procellariiformes, band-rumped storm-petrels exhibit strong philopatry,
or fidelity to their natal sites
[[Page 67820]]
(Allan 1962, p. 274; Harris 1969, pp. 96, 113, 120; Harrison et al.
1990, p. 49; Smith et al. 2007, pp. 768-769). Both of these
characteristics contribute to isolation of breeding populations, in
spite of the absence of physical barriers such as land masses within
ocean basins (Friesen et al. 2007a, pp. 1777-1778).
Band-rumped storm-petrels from Hawaii are likely to encounter
individuals from other populations only very rarely. The approximate
distances from Hawaii to other known breeding sites are much greater
than the birds' average foraging range of 860 mi (1,200 km): 4,000 mi
(6,600 km) to Japan and 4,600 mi (7,400 km) to Galapagos (the two other
Pacific populations), and 7,900 mi (12,700 km) to Madeira, 7,300 mi
(11,700 km) to the Azores, and 9,700 mi (15,600 km) to Ascension Island
(in the Atlantic). Data from at-sea surveys of the eastern tropical
Pacific conducted since 1988 show that the density of band-rumped
storm-petrels attenuates north and northwest of Galapagos and that the
species rarely occurs in a broad area southeast of Hawaii (Pitman,
Ballance, and Joyce 2015, unpublished). This pattern suggests a gap in
the at-sea distribution of this species, and low likelihood of
immigration on an ecological timescale, between Hawaii and Galapagos.
We are not aware of any data describing the at-sea distribution of this
species between Hawaii and Japan, but the absence of breeding records
from western Micronesia (Pyle and Engbring 1985, p. 59) indicates a
distributional gap between these two archipelagoes as well. Other than
occasional encounters in their foraging habitat, the vast expanses of
ocean between Japan, Hawaii, and Galapagos provide for no other sources
of potential connectivity between band-rumped storm-petrel populations
in the Pacific, such as additional breeding sites.
Even those disparate breeding populations of pelagic seabirds that
do overlap at sea may remain largely isolated otherwise and exhibit
genetic differentiation (e.g., Walsh and Edwards 2005, pp. 290, 293).
Despite the birds' capacity to move across large areas of ocean,
genetic differentiation among breeding populations of band-rumped
storm-petrels is high (Friesen et al. 2007b, p. 18590; Smith et al.
2007, p. 768), even between populations nesting in different seasons on
the same island (in Galapagos; Smith and Friesen 2007, p. 1599).
Genetic analysis found low relatedness (1) between Atlantic and Pacific
populations; (2) among Japan, Hawaii, and Galapagos populations; or (3)
among Cape Verde, Ascension, and northeast Atlantic breeding
populations (Smith et al. 2007, p. 768). Hawaiian birds have not been
well-sampled for genetic analysis, but the few individuals from Hawaii
included in a rangewide analysis showed that Hawaiian birds differed
from all other populations, and were most closely related to birds from
Japan (Friesen et al. 2007b, p. 18590).
We have determined that the Hawaii population of the band-rumped
storm-petrel is discrete from the rest of the taxon because its
breeding and foraging range are markedly separated from those of other
populations. The Hawaii population is geographically isolated from
populations in Japan and Galapagos, as well as from populations in very
distant island groups in the central and western Atlantic Ocean.
Molecular evidence indicates that the genetic structure of the species
reflects the spatial or temporal separation of individual populations;
the scant molecular data from Hawaii suggest that this holds for the
Hawaii population as well.
Significance
Under our DPS Policy, once we have determined that a population
segment is discrete, we consider its biological and ecological
significance to the larger taxon to which it belongs. This
consideration may include, but is not limited to: (1) Evidence of the
persistence of the discrete population segment in an ecological setting
that is unusual or unique for the taxon, (2) evidence that loss of the
population segment would result in a significant gap in the range of
the taxon, (3) evidence that the population segment represents the only
surviving natural occurrence of a taxon that may be more abundant
elsewhere as an introduced population outside its historical range, or
(4) evidence that the discrete population segment differs markedly from
other populations of the species in its genetic characteristics. We
have found substantial evidence that the Hawaii population of the band-
rumped storm-petrel meets two of the significance criteria listed
above: the loss of this population would result in a significant gap in
the range of the taxon, and this population persists in a unique
ecological setting. As described above, the physical isolation that
defines the discreteness of Hawaii population is likely reflected in
genetic differentiation from other populations, but at this time we
lack sufficient data to consider genetic characteristics as an
independent factor in our determination of the Hawaii population's
significance to the rest of the taxon. Genetic patterns on an ocean-
basin or species-wide scale, however, have implications for
connectivity and potential gaps in the band-rumped storm-petrel's range
(described below).
Dispersal between populations of seabird species with ranges
fragmented by large expanses of ocean may play a vital role in the
persistence of individual populations (Bicknell et al. 2012, p. 2872).
No evidence currently exists of such dispersal among Pacific
populations of band-rumped storm-petrels at frequencies or in numbers
that would change the population status between years, for example, by
providing immigrants that compensate for breeding failure or adult
mortality resulting from predation, as has been hypothesized for
Leach's storm-petrel in the Atlantic (Bicknell et al. 2012, p. 2872).
Given the remnant population of band-rumped storm-petrels in Hawaii and
recently documented decline in Japan (Biodiversity Center of Japan
2014, p. 1), we would not expect to see exchange on such short
timescales. However, genetic evidence is suggestive of exchange between
these two populations on an evolutionary timescale (Friesen et al.
2007b, p. 18590).
The loss of this population would result in a significant gap in
the range of the band-rumped storm-petrel. As noted above, seabirds in
the order Procellariiformes, including the band-rumped storm-petrel,
exhibit very high natal site fidelity, and so are slow to recolonize
extirpated areas or range-gaps (Jones 2010, p. 1214), and may lack
local adaptations; they thus face a potentially increased risk of
extinction with the loss of individual populations (Smith et al. 2007,
p. 770). The Hawaii population of the band-rumped storm petrel
constitutes the entire Central Pacific distribution of the species,
located roughly half-way between the populations in Galapagos and Japan
(see Figure 1, below), and its loss would create a gap of approximately
8,500 mi (13,680 km) between them and significantly reducing the
likelihood of connectivity and genetic exchange. Such exchange would be
reliant on chance occurrences, such as severe storms that could result
in birds being displaced to the opposite side of the Pacific Ocean
basin, and such chance dispersal events would not necessarily result in
breeding.
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The Hawaii population of the band-rumped storm-petrel is
significant also because it persists in a unique ecological setting.
This is the only population of the species known to nest at high-
elevation sites (above 6,000 ft (1,800 m)) (Banko et al. 1991, pp. 651-
653; Athens et al. 1991, p. 95). In prehistory, the species likely
nested in lowland habitats and more accessible habitats in Hawaii as
well as in the high-elevation and otherwise remote areas where the
species is found today; archaeological evidence suggests that band-
rumped storm-petrels were once sufficiently common at both high (5,260
and 6,550 ft (1,600 and 2,000 m)) and low elevations on Hawaii Island
to be used as a food source by humans (Ziegler pers. comm. in Harrison
et al. 1990, pp. 47-48; Athens et al. 1991, pp. 65, 78-80; Banko et al.
1991, p. 650). In lowland areas, the species was common enough for the
Hawaiians to name it and to identify it by its call (Harrison et al.
1990, p. 47; Banko et al. 1991, p. 650). In addition to the impacts of
harvest by humans in prehistory, seabirds in Hawaii, including the
band-rumped storm-petrel, were negatively affected by the proliferation
of nonnative predators such as rats and pigs, and, later, cats and
mongoose, and by loss of habitat (reviewed in Duffy 2010, pp. 194-196).
Predation and habitat loss combined likely led to the extirpation of
the band-rumped storm-petrel from coastal and lowland habitats and
other accessible nesting areas, as occurred in the
[[Page 67822]]
endangered Hawaiian petrel (Pterodroma sandwichensis) and threatened
Newell's shearwater (Puffinus newelli), which have similar nesting
habits and life histories (Olson and James 1982, p. 43; Slotterback
2002, p. 6; Troy et al. 2014, pp. 315, 325-326). The band-rumped storm-
petrel's persistence in sites such as the Southwest Rift Zone (6,900 ft
(2,100 m)) on Mauna Loa (Hawaii Island) has required them to surmount
physiological challenges posed by nesting in high-elevation conditions
(cold temperatures, low humidity, and less oxygen). They may possess
special adaptations for this, such as reduction in porosity and other
eggshell modifications to reduce the loss of water and carbon dioxide
during incubation at high elevation (Rahn et al. 1977, p. 3097; Carey
et al. 1982, p. 716; Carey et al. 1983, p. 349). In sum, the remnant
distribution of band-rumped storm-petrel breeding sites in only the
most remote and rugged terrain in Hawaii reflects the conditions
necessary for the species' persistence in Hawaii (relatively
undisturbed habitat in areas least accessible to predators) and also
reflects unique adaptations that facilitate the species' persistence in
high-elevation areas.
We have determined that the Hawaii population of band-rumped storm-
petrel is significant to the rest of the taxon. Its loss would result
in a gap in the range of the species of more than 8,500 mi (13,680 km),
reducing and potentially precluding connectivity between the two
remaining populations in the Pacific Basin. In addition, the Hawaii
population nests at high elevation on some islands, constituting a
unique ecological setting represented nowhere else in the species'
breeding range.
DPS Conclusion
We have evaluated the Hawaii population of band-rumped storm-petrel
to determine if it meets the definition of a DPS, considering its
discreteness and significance as required by our policy. We have found
that this population is markedly separated from other populations by
geographic distance, and this separation is likely reflected in the
population's genetic distinctiveness. The Hawaii population is
significant to the rest of the species because its loss would result in
a significant gap in the species' range; Hawaii is located roughly
half-way between the other two populations in the Pacific Ocean, and
little or no evidence exists of current overlap at sea between the
Hawaii population and either the Japan or Galapagos populations. The
Hawaii population of band-rumped storm-petrel also nests at high
elevation in Hawaii--conditions at high elevation constitute an
ecological setting unique to the species. We conclude that the Hawaii
population of band-rumped storm-petrel is a distinct vertebrate
population segment under our February 7, 1996, DPS Policy (61 FR 4722),
and that it warrants review for listing under the Act. Therefore, we
have incorporated the Hawaii DPS of the band-rumped storm-petrel in our
evaluation of threats affecting the other 48 species addressed in this
rule (summarized above; see also Summary of Factors Affecting the 49
Species From the Hawaiian Islands, below).
Summary of Factors Affecting the 49 Species From the Hawaiian Islands
Section 4 of the Act (16 U.S.C. 1533), and its implementing
regulations (50 CFR part 424), set forth the procedures for adding
species to the Federal Lists of Endangered and Threatened Wildlife and
Plants. A species may be determined to be an endangered or threatened
species due to one or more of the five factors described in section
4(a)(1) of the Act: (A) The present or threatened destruction,
modification, or curtailment of its habitat or range; (B)
overutilization for commercial, recreational, scientific, or
educational purposes; (C) disease or predation; (D) the inadequacy of
existing regulatory mechanisms; and (E) other natural or manmade
factors affecting its continued existence. Listing actions may be
warranted based on any of the above threat factors, singly or in
combination. Each of these factors is discussed below.
In considering factors that might constitute threats to a species,
we must look beyond the exposure of the species to a factor to evaluate
whether the species responds to the factor in a way that causes impacts
to the species or is likely to cause impacts in the future. If a
species responds negatively to such exposure, the factor may be a
threat and, during the status review, our aim is to determine whether
impacts are or will be of an intensity or magnitude to place the
species at risk. The factor is a threat if it drives, or contributes
to, the risk of extinction of the species such that the species
warrants listing as an endangered or threatened species as those terms
are defined by the Act. This does not necessarily require empirical
proof of a threat. The combination of exposure and some corroborating
evidence of how the species is likely affected could suffice. In sum,
the mere identification of factors that could affect a species
negatively is not sufficient to compel a finding that listing is
appropriate; we require evidence that these factors act on the species
to the point that the species meets the definition of an endangered or
threatened species.
If we determine that the threats posed to a species by one or more
of the five listing factors are, or are likely to become, of such
magnitude and/or intensity that the species meets the definition of
either endangered or threatened under section 3 of the Act, that
species may then be listed as endangered or threatened. The Act defines
an endangered species as ``in danger of extinction throughout all or a
significant portion of its range,'' and a threatened species as
``likely to become an endangered species within the foreseeable future
throughout all or a significant portion of its range.'' The threats to
each of the individual 49 species are summarized in Table 2, and
discussed in detail below.
We acknowledge that the specific threats to the individual species
in this final rule are not all completely understood. Scientific study
of each of the 49 species is limited because of their rarity and the
challenging logistics associated with conducting field work in Hawaii
(areas are typically remote, difficult to access, challenging work
environments, and expensive to survey in a comprehensive manner).
However, information is available on many of the threats that act on
Hawaiian ecosystems, and, for some ecosystems, these threats are well
studied and understood. Each of the native species that occurs in
Hawaiian ecosystems suffers from exposure to those threats to differing
degrees. For the purposes of our listing determination, the best
available scientific information leads us to conclude that the threats
that act at the ecosystem level also act on each of the species that
occurs in those ecosystems. In some cases we have additionally
identified species-specific threats, such as loss of host plants.
The following threats affect the 49 species in one or more of the
ecosystems addressed in this rule:
(1) Modification and destruction of habitat, including streams,
ponds, and anchialine pools, by urban development and water extraction.
Human activities also contribute to increased sedimentation in
anchialine pools.
(2) Habitat destruction and modification by feral ungulates
including pigs, goats, axis deer, black-tailed deer, mouflon, sheep,
and cattle. The disturbance of soils by these animals causes erosion
and creates fertile seedbeds for nonnative plants, leading to further
habitat degradation. Ungulates also trample seedlings.
[[Page 67823]]
(3) Habitat destruction and modification by nonnative plants.
Nonnative plants modify availability of light, alter soil-water
regimes, modify nutrient cycling, alter fire regimes, and ultimately
convert native dominated plant communities to nonnative plant
communities. They also cause or contribute to loss of host plants used
for food and nesting by the yellow-faced bees.
(4) Habitat destruction by wildfires caused naturally or by humans.
Fires also destroy the native plant seedbank, and contribute to habitat
conversion of native forest to nonnative grasslands (grass/fire cycle).
(5) Habitat destruction and modification, or direct damage and
death, by stochastic events including drought, erosion, flooding, tree
falls, rock falls, landslides, hurricanes, and tsunamis.
(6) Illegal collection of anchialine pool shrimp for personal use
or commercial trade.
(7) Herbivory or defoliation of native plants by ungulates, rats,
slugs, and black twig borers, which have been observed to contribute to
the decline or death of 35 the 39 plant species (except for Cyperus
neokunthianus, Cyrtandra hematos, Lepidium orbiculare, and Stenogyne
kaalae ssp. sherffii). Herbivory also destroys seeds and fruit and
contributes to lack of reproduction in the wild and low genetic
diversity compounding the decline of native plants.
(8) Predation of the band-rumped storm-petrel by rats, barn owls,
cats, and mongoose.
(9) Predation of the orangeblack Hawaiian damselfly by bullfrogs,
backswimmers, Jackson's chameleons, and nonnative fish.
(10) Predation of the anchialine pool shrimp by nonnative fish.
(11) Predation of Hylaeus bees by ants and wasps.
(12) Competition for food and nesting sites of the Hylaeus yellow-
faced bees by nonnative ants, wasps, and bees, and competition for food
and habitat of the orangeblack Hawaiian damselfly by caddisflies.
Competition for space and food resources of the anchialine pool shrimp
by nonnative fish.
(13) Injury and mortality of the band-rumped storm-petrel caused by
artificial lighting, communication towers, and power lines.
(14) Injury and mortality of the band-rumped storm-petrel by the
activities of fisheries and encounters with marine debris.
(15) Low numbers and/or no reproduction of all 49 species
exacerbated by one or more of the above threats, combined with
inability of the species to adapt to sea-level rise or other factors
associated with climate change.
Existing regulatory mechanisms do not ameliorate these threats for
any of the 49 species such that listing is not warranted. Each of the
threats listed above is discussed in more detail below, and summarized
in Table 2.
[[Page 67824]]
Table 2--Summary of Primary Threats Identified for Each of the 49 Hawaiian Islands Species
----------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Factor A Factor B Factor C Factor D Factor E
-----------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Predation/ Inadequate Other
Species Ecosystem Agriculture and Non- native Stochastic Over- Predation/ herbivory by Predation/ existing species- Climate
urban Ungulates plants Fire events utilization herbivory by other NN herbivory by NN regulatory specific change
development ungulates vertebrates invertebrates mechanisms threats
----------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Plants:
Asplenium diellaciniatum... MM................. ............... P, G, BTD... X.......... ............ ............... .............. X............. .............. ................ X............ LN........ X.
Calamagrostis expansa...... MW................. ............... P........... X.......... ............ ............... .............. X............. R............. ................ X............ LN........ X, Ft.
Cyanea kauaulaensis........ LW................. ............... ............ X.......... ............ E, F, L........ .............. .............. R............. S............... X............ LN, NR.... X.
Cyclosorus boydiae......... LW, MW............. X, WE.......... P........... X.......... ............ DR, F, L....... .............. X............. .............. ................ X............ LN........ X, Ft.
Cyperus neokunthianus...... LW................. ............... P........... X.......... ............ ............... .............. .............. .............. ................ X............ LN........ X.
Cyrtandra hematos.......... MW................. ............... P, G........ X.......... ............ ............... .............. .............. .............. ................ X............ HY, LN, NR X, Ft.
Deparia kaalaana........... LM, LW............. ............... P........... X.......... ............ DR, F.......... .............. X............. .............. S............... X............ LN........ X.
Dryopteris glabra var. MW................. ............... P, BTD...... X.......... ............ L.............. .............. .............. R............. S............... X............ LN........ X, Ft.
pusilla.
Exocarpos menziesii........ LM, MM, MD......... ............... G, M, SH.... X.......... X........... ............... .............. X............. .............. ................ X............ LN........ X, Ft.
Festuca hawaiiensis........ MD................. ............... G, SH....... X.......... X........... ............... .............. X............. .............. ................ X............ LN........ X, Ft.
Gardenia remyi............. LM, LW............. ............... P, G, D..... X.......... ............ L.............. .............. X............. R............. ................ X............ LN, NR.... X, Ft.
Huperzia stemmermanniae.... MW................. ............... P, G, D, C.. X.......... ............ DR............. .............. X............. .............. ................ X............ LN........ X, Ft.
Hypolepis hawaiiensis var. MW................. ............... ............ X.......... ............ ............... .............. .............. .............. S............... X............ LN........ X, Ft.
mauiensis.
Joinvillea ascendens ssp. LM, LW, MW, MM..... ............... P, G, D, BTD X.......... X........... L.............. .............. X............. R............. ................ X............ LN, NR.... X, Ft.
ascendens.
Kadua fluviatilis.......... LM, LW............. ............... P, G........ X.......... ............ L.............. .............. X............. .............. ................ X............ LN........ X, Ft.
Kadua haupuensis........... LM................. ............... P........... X.......... ............ L.............. .............. .............. R............. ................ X............ LN........ X.
Labordia lorenciana........ MM................. ............... P, G, BTD... X.......... X........... F, L, TF....... .............. X............. R............. BTB............. X............ LN, NR.... X.
Lepidium orbiculare........ LM................. ............... P, G........ X.......... ............ L.............. .............. .............. .............. ................ X............ LN........ X.
Microlepia strigosa var. LM, MW, MM......... ............... P, G........ X.......... ............ ............... .............. X............. .............. ................ X............ HY, LN.... X.
mauiensis.
Myrsine fosbergii.......... LM, LW, MW......... ............... P, G........ X.......... ............ ............... .............. X............. .............. ................ X............ HY, LN.... X, Ft.
Nothocestrum latifolium.... LD, LM, DC......... ............... P, G, D, X.......... X........... ............... .............. X............. .............. BTB............. X............ LN, NR.... X, Ft.
BTD, M, C.
Ochrosia haleakalae........ LM, MM, DC......... ............... P, G, C..... X.......... X........... ............... .............. X............. R............. S............... X............ LN, NR.... X, Ft.
Phyllostegia brevidens..... LW, MW, WC......... ............... P........... X.......... ............ E, L........... .............. X............. .............. S............... X............ LN........ X, Ft.
Phyllostegia helleri....... LW, MW, WC......... ............... P, G........ X.......... ............ L.............. .............. .............. R............. ................ X............ LN........ X, Ft.
Phyllostegia stachyoides... MW, MM............. ............... P, G, D..... X.......... ............ DR, E, F, L, RF .............. X............. R............. S............... X............ LN........ X, Ft.
Portulaca villosa.......... CO, LD, MD......... ............... G, D, M, C.. X.......... X........... L, RF.......... .............. X............. .............. ................ X............ LN........ X, Ft.
Pritchardia bakeri......... LM................. ............... P........... X.......... ............ HUR............ .............. .............. R............. ................ X............ LN........ X.
Pseudognaphalium CO................. ............... G, D........ X.......... ............ L, RF.......... .............. X............. .............. ................ X............ LN........ X, Ft.
sandwicensium var.
molokaiense.
Ranunculus hawaiensis...... MM, MD, SA......... ............... P, M, C..... X.......... ............ DR, E.......... .............. X............. .............. ................ X............ LN........ X, Ft.
Ranunculus mauiensis....... MW, MM, WC......... ............... P, G, D, X.......... X........... DR, E, L....... .............. X............. R............. S............... X............ LN........ X, Ft.
BTD, C.
Sanicula sandwicensis...... MM, MD, SA......... ............... P, G........ X.......... X........... DR, E, F....... .............. X............. R............. ................ X............ LN........ X, Ft.
Santalum involutum......... LM, LW............. ............... P, G........ X.......... X........... ............... .............. X............. R............. ................ X............ LN........ X.
Schiedea diffusa ssp. LW, MW............. ............... P........... X.......... ............ ............... .............. .............. R............. S............... X............ LN........ X.
diffusa.
Schiedea pubescens......... LW, MW, MM, WC..... ............... P, G, D, C.. X.......... X........... DR, E, F....... .............. X............. R............. S............... X............ LN........ X, Ft.
Sicyos lanceoloideus....... LM, MM, DC......... ............... P, G, BTD... X.......... X........... DR............. .............. X............. .............. ................ X............ LN........ X, Ft.
Sicyos macrophyllus........ MW, MM, MD......... ............... P, M, C..... X.......... X........... ............... .............. X............. R............. ................ X............ LN........ X, Ft.
Solanum nelsonii........... CO................. ............... D, C........ X.......... X........... DR, E, F, TS... .............. X............. R............. ................ X............ LN, SL.... X, Ft.
Stenogyne kaalae ssp. LW................. ............... P........... X.......... ............ ............... .............. .............. .............. ................ X............ LN........ X, Ft.
sherffii.
Wikstroemia skottsbergiana. LW................. ............... P, G........ X.......... ............ L.............. .............. .............. R............. ................ X............ LN........ X.
ANIMALS:
Band-rumped storm-petrel CO, SA, DC, WC..... ............... G........... X.......... ............ E, L, HUR...... .............. .............. R, O, CA, MO.. ................ X............ LI, ST, H, X.
(Oceanodroma castro). LN.
Orangeblack Hawaiian AP, CO, LD, LM..... X, WE.......... P, G, D..... X.......... ............ DR, F, HUR..... .............. .............. FS, BF, JC.... BS.............. X............ LN, CD.... X.
damselfly (Megalagrion
xanthomelas).
Anchialine pool shrimp AP................. X, WE.......... ............ ........... ............ ............... X............. .............. FS............ ................ X............ FS, H, LN, X.
(Procaris hawaiana). RU, SD,
SL.
Yellow-faced bee (Hylaeus CO, LD............. X.............. P, G, D, SH, X.......... X........... DR, HUR, TS.... .............. .............. .............. A, W,........... X............ LN, W, B, X.
anthracinus). C. LHP.
Yellow-faced bee (Hylaeus CO, LD............. X.............. P, G, D, M, X.......... X........... DR, HUR, TS.... .............. .............. .............. A, W............ X............ LN, W, B, X.
assimulans). C. LHP.
Yellow-faced bee (Hylaeus CO, LD, LM......... X.............. P, G, D, C.. X.......... X........... DR, HUR, TS.... .............. .............. .............. A, W............ X............ LN, W, B, X.
facilis). LHP.
[[Page 67825]]
Yellow-faced bee (Hylaeus CO, LD............. X.............. P, G, D, C.. X.......... X........... DR, HUR, TS.... .............. .............. .............. A, W............ X............ LN, W, B, X.
hilaris). LHP.
Yellow-faced bee (Hylaeus LM................. ............... P........... X.......... X........... DR, HUR........ .............. .............. .............. A, W............ X............ LN, W, B, X.
kuakea). LHP.
Yellow-faced bee (Hylaeus CO, LD............. X.............. D........... X.......... X........... DR, HUR, TS.... .............. .............. .............. A, W............ X............ LN, W, B, X.
longiceps). LHP, RU.
Yellow-faced bee (Hylaeus LM................. ............... P........... X.......... X........... DR, HUR........ .............. .............. .............. A, W............ X............ LN, W, B, X.
mana). LHP.
----------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
Factor A = Habitat Modification; Factor B = Overutilization; Factor C = Disease or Predation; Factor D = Inadequacy of Regulatory Mechanisms: the Xs in this column indicate that existing regulatory mechanisms do not ameliorate the
threats to the species such that listing is not warranted (we do not identify Factor D, in and of itself, as a threat to the species); Factor E = Other Species-Specific Threats.
AP = Anchialine Pools; CO = Coastal; LD = Lowland Dry; LM = Lowland Mesic; LW = Lowland Wet; MW = Montane Wet; MM = Montane Mesic; MD = Montane Dry; SA = Subalpine; DC = Dry Cliff; WC = Wet Cliff.
A = Ants, B = Bees (competition); BF = Bullfrog; BS = Backswimmer; BTB = Black Twig Borer; BTD = Black-tailed Deer; C = Cattle; CA = Cats; CD = Caddisflies; D = Axis Deer; ; FS = Fish; G = Goats; JC = Jackson's Chameleons; M =
Mouflon; MO = Mongoose; O = Barn Owls; P = Pigs; R = Rats; S = Slugs; SH = Sheep; W = Wasps (competition, predation).
DR = Drought; E = Erosion; F = Flooding; Ft = assessed in Fortini et al. (2013) climate change vulnerability analysis; H = Human (fisheries, marine debris, contamination); HUR = Hurricanes; HY = Hybridization; L = Landslides; LHP =
Loss of Host Plants; LI = Lights; LN = Low Numbers; NR = No Regeneration; RF = Rockfalls; RU = Recreational Use (swimming, fishing, dumping trash and nonnative fish); SD = Sedimentation; SL = Sea Level Rise; ST = Structures; TF =
Tree Fall; TS = Tsunami; WE = Water Extraction.
[[Page 67826]]
Factor A. The Present or Threatened Destruction, Modification, or
Curtailment of Their Habitat or Range
The Hawaiian Islands are located over 2,000 mi (3,200 km) from the
nearest continent. This isolation has allowed the few plants and
animals transported to the islands by wind, water, or birds to evolve
into many varied and endemic species. The only native terrestrial
mammals on the Hawaiian Islands include two bat taxa, the Hawaiian
hoary bat (Lasiurus cinereus semotus), and an extinct, unnamed
insectivorous bat (Ziegler 2002, p. 245). The native plants of the
Hawaiian Islands therefore evolved in the absence of mammalian
predators, browsers, or grazers, and subsequently, many native species
lost unneeded defenses against threats typical of continental
environments such as herbivory and competition with aggressive, weedy
plant species (Loope 1992, p. 11; Gagne and Cuddihy 1999, p. 45; Wagner
et al. 1999, pp. 3-6). For example, Carlquist (in Carlquist and Cole
1974, p. 29) notes, ``Hawaiian plants are notably nonpoisonous, free
from armament, and free from many characteristics thought to be
deterrents to herbivores (oils, resins, stinging hairs, coarse
texture).'' In addition, species restricted to highly specialized
habitats (e.g., Hawaiian damselflies) or food and nesting sources
(e.g., Hawaiian yellow-faced bees) are particularly vulnerable to
changes in their habitat (Carlquist and Cole 1974, pp. 28-29; Loope
1992, pp. 3-6).
Habitat Destruction and Modification by Agriculture and Urban
Development
Past land use practices such as agriculture or urban development
have resulted in little or no native vegetation below 2,000 ft (600 m)
throughout the Hawaiian Islands (TNC 2006). These land use practices
negatively affect the anchialine pool, coastal, lowland dry, and
lowland mesic ecosystems, including streams and wetlands that occur
within these areas. Hawaii's agricultural industries (e.g., sugar cane,
pineapple) have been declining in importance, and large tracts of
former agricultural lands are being converted into residential areas or
left fallow (TNC 2007). In addition, Hawaii's population has increased
almost 10 percent in the past 10 years, further increasing demands on
limited land and water resources in the islands (Hawaii Department of
Business, Economic Development and Tourism 2013, in litt.).
Development and urbanization of anchialine pool, coastal, lowland
dry, and lowland mesic ecosystems on Oahu, Molokai, Maui, Lanai, and
Hawaii Island are a threat to some species:
On Oahu, the plant Cyclosorus boydiae, the orangeblack
Hawaiian damselfly, and the yellow-faced bees Hylaeus anthracinus, H.
assimulans, H. facilis, and H. longiceps.
On Molokai, the orangeblack Hawaiian damselfly and the
yellow-faced bees Hylaeus anthracinus, H. facilis, H. hilaris, and H.
longiceps.
On Maui, the plant Cyclosorus boydiae, the orangeblack
Hawaiian damselfly, the anchialine pool shrimp Procaris hawaiana, and
the yellow-faced bees Hylaeus anthracinus, H. assimulans, H. facilis,
H. hilaris, and H. longiceps.
On Lanai, the orangeblack Hawaiian damselfly, and the
yellow-faced bees Hylaeus anthracinus, H. assimulans, H. facilis, H.
hilaris, and H. longiceps.
On Hawaii Island, the orangeblack Hawaiian damselfly, the
anchialine pool shrimp Procaris hawaiana, and the yellow-faced bee
Hylaeus anthracinus. (Daly and Magnacca 2003, pp. 55, 173; Palmer 2003,
p. 88; Magnacca 2007, p. 188; Magnacca and King 2013, pp. 22-25).
Although we are unaware of any comprehensive, site-by-site
assessment of wetland development in Hawaii (Erikson and Puttock 2006,
p. 40), Dahl (1990, p. 7) estimated that at least 12 percent of lowland
to upper-elevation wetlands in Hawaii had been converted to non-wetland
habitat by the 1980s. If only coastal plain (below 1,000 ft (300 m))
marshlands and wetlands are considered, it is estimated that 30 percent
were developed or converted to agricultural use (Kosaka 1990, in
litt.). Records show the modification and reduction in area of these
marshlands and wetlands that provided habitat for many damselfly
species, including the orangeblack Hawaiian damselfly (Englund 2001, p.
256; Rees and Reed 2013, Fig 2S). Once modified, these areas then lack
the aquatic habitat features that the orangeblack Hawaiian damselfly
requires for essential life-history needs, such as pools of
intermittent streams, ponds, and coastal springs (Polhemus 1996 pp. 30-
31, 36). Although the filling of wetlands is regulated by section 404
of the Clean Water Act (33 U.S.C. 1251 et seq.), the loss of riparian
or wetland habitats utilized by the orangeblack Hawaiian damselfly may
still occur due to Hawaii's population growth and development, with
concurrent demands on limited developable land and water resources. The
State's Commission of Water Resource Management (CWRM) recognizes the
need to update the 2008 water resource protection plan, and an update
is currently under development (CWRM 2014, in litt.). In addition,
marshes have been slowly filled and converted to meadow habitat as a
result of sedimentation from increased storm water runoff from upslope
development, the accumulation of uncontrolled growth of invasive
vegetation, and blockage of downslope drainage (Wilson Okamoto &
Associates, Inc. 1993, pp. 3-4-3-5). Agriculture and urban development
have thus contributed to habitat destruction and modification, and
continue to be a threat to the habitat of the orangeblack Hawaiian
damselfly and the fern, Cyclosorus boydiae.
On Hawaii Island, it is estimated that up to 90 percent of the
anchialine pools have been destroyed or altered by human activities,
including bulldozing and filling of pools (Brock 2004, p. i; Bailey-
Brock and Brock 1993, p. 354). Dumping of trash and nonnative fish has
affected anchialine pools on this island (Brock 2004, pp. 13-17) (see
Factor E. Other Natural or Manmade Factors Affecting Their Continued
Existence, below). Brock also noted that garbage like bottles and cans
appear to have no net negative impact, while the dumping of used oil,
oil filters, and grease has resulted in the disappearance of the
anchialine pool shrimp Halaocaridina rubra from a pool adjacent to
Honokohau Harbor on Hawaii Island. Lua O Palahemo (where Procaris
hawaiana occurs) on Hawaii Island is accessible to the public, and
dumping has occurred there (Brock 2004, pp. 13-17). We are not aware of
any dumping activities within the two Maui anchialine pools known to be
occupied by P. hawaiana; however, this threat remains a possibility
(Brock 2004, pp. 13-17).
Destruction and modification of Hylaeus habitat by urbanization and
land use conversion, including agriculture, has led to the
fragmentation of foraging and nesting habitat of these species. In
particular, because native host plant species are known to be essential
to the yellow-faced bees for foraging of nectar and pollen, any further
loss of this habitat may reduce their long-term chances for recovery.
Additionally, further destruction and modification of Hylaeus habitat
is also likely to facilitate the introduction and spread of nonnative
plants within these areas (see ``Habitat Destruction and Modification
by Nonnative Plants,'' below).
Habitat Destruction and Modification by Nonnative Ungulates
Nonnative ungulates have greatly affected the native vegetation, as
well as the native fauna, of the Hawaiian
[[Page 67827]]
Islands. Impacts to the native species and ecosystems accelerated
following the arrival of Captain James Cook in 1778. The Cook
expedition and subsequent explorers intentionally introduced a European
race of pigs (i.e., boars) and other livestock such as goats to serve
as food sources for seagoing explorers (Tomich 1986, pp. 120-121; Loope
1998, p. 752). The mild climate of the islands, combined with lack of
competitors or predators, led to the successful establishment of large
populations of these feral mammals, to the detriment of native Hawaiian
species and ecosystems (Cox 1992, pp. 116-117). The presence of
introduced mammals is considered one of the primary factors underlying
the modification and destruction of native vegetation and habitats of
the Hawaiian Islands (Cox 1992, pp. 118-119). All of the 11 ecosystems
on the main islands (except Kahoolawe) are currently affected by
habitat destruction resulting from the activities of various
combinations of nonnative ungulates, including pigs (Sus scrofa), goats
(Capra hircus), axis deer (Axis axis), black-tailed deer (Odocoileus
hemionus columbianus), sheep (Ovis aries), mouflon (Ovis gmelini
musimon) and mouflon-sheep hybrids, and cattle (Bos taurus). Habitat
destruction or modification by ungulates is a threat to 37 of the 39
plant species, the band-rumped storm-petrel, the orangeblack Hawaiian
damselfly, and the seven yellow-faced bees (see Table 2).
Pigs (Sus scrofa)
The destruction or modification of habitat by pigs is currently a
threat to four of the ecosystems (lowland mesic, lowland wet, montane
wet, and montane mesic) in which these species occur. Feral pigs are
known to cause deleterious impacts to ecosystem processes and functions
throughout their worldwide distribution (Campbell and Long 2009, p.
2319). Pigs have been described as having the most pervasive and
disruptive nonnative influences on the unique ecosystems of the
Hawaiian Islands and are widely recognized as one of the greatest
current threats (Aplet et al. 1991. p. 56; Anderson and Stone 1993, p.
195; Anderson et al. 2007, in litt.). Introduced European pigs
hybridized with smaller, domesticated Polynesian pigs, became feral,
and invaded forested areas, especially mesic and wet forests, from low
to high elevations; they are present on all the main Hawaiian Islands
except Lanai and Kahoolawe, where they have been eradicated (Tomich
1986, pp. 120-121; Munro (1911-1930) 2007, p. 85). By the early 1900s,
feral pigs were already recognized as a serious threat to these areas,
and an eradication project was conducted by the Hawaii Territorial
Board of Agriculture and Forestry, which removed 170,000 pigs from
forests Statewide (Diong 1982, p. 63).
Feral pigs are extremely destructive and have both direct and
indirect impacts on native plant communities. While rooting in the
earth in search of invertebrates and plant material, pigs directly
affect native plants by disturbing and destroying vegetative cover and
by trampling plants and seedlings. It has been estimated that at a
conservative rooting rate of 2 square yards (sq yd) (1.7 square meters
(sq m)) per minute and only 4 hours of foraging per day, a single pig
could disturb over 1,600 sq yd (1,340 sq m) (or approximately 0.3 acres
(ac) (0.1 hectares (ha)) of groundcover per week (Anderson et al. 2007,
in litt.). Feral pigs are a major vector for the establishment and
spread of invasive nonnative plant species, such as Passiflora
tarminiana and Psidium cattleianum, by dispersing seeds carried on
their hooves and coats and in their feces (which also serve to
fertilize disturbed soil) (Diong 1982, pp. 169-170; Matson 1990, p.
245; Siemann et al. 2009, p. 547). Pigs also feed directly on native
plants such as Hawaiian tree ferns. Pigs preferentially eat the core of
tree-fern trunks, and these cored trunks then fill with rainwater and
serve as breeding sites for introduced mosquitoes that spread avian
malaria, with devastating consequences for Hawaii's native forest birds
(Baker 1975, p. 79). Additionally, rooting pigs contribute to erosion,
especially on slopes, by clearing vegetation and creating large areas
of disturbed soil (Smith 1985, pp. 190, 192, 196, 200, 204, 230-231;
Stone 1985, pp. 254-255, 262-264; Medeiros et al. 1986, pp. 27-28;
Scott et al. 1986, pp. 360-361; Tomich 1986, pp. 120-126; Cuddihy and
Stone 1990, pp. 64-65; Aplet et al. 1991, p. 56; Loope et al. 1991, pp.
1-21; Gagne and Cuddihy 1999, p. 52; Nogueira-Filho et al. 2009, pp.
3677-3682; Dunkell et al. 2011, pp. 175-177). The resulting erosion
alters native plant communities by damaging individual plants,
contributing to watershed degradation, and changing nutrient
availability for plants; erosion also affects aquatic animals by
increasing sedimentation in streams and pools (Vitousek et al. 2009,
pp. 3074-3086; Nogueira-Filho et al. 2009, p. 3681; Cuddihy and Stone
1992, p. 667). The following 15 plants are at risk from erosion and
landslides resulting from the activities of feral pigs: Cyclosorus
boydiae, Dryopteris glabra var. pusilla, Gardenia remyi, Joinvillea
ascendens ssp. ascendens, Kadua fluviatilis, K. haupuensis, Labordia
lorenciana, Lepidium orbiculare, Phyllostegia brevidens, P. helleri, P.
stachyoides, Ranunculus hawaiensis, R. mauiensis, Sanicula
sandwicensis, and Schiedea pubescens. Thirty-two of the 39 plants (all
except for Cyanea kauaulaensis, Exocarpos menziesii, Festuca
hawaiiensis, Hypolepis hawaiiensis var. mauiensis, Portulaca villosa,
Pseudognaphalium sandwicensium var. molokaiense, and Solanum nelsonii)
are at risk of habitat destruction and modification by feral pigs, and
the orangeblack Hawaiian damselfly and six of the seven yellow-faced
bees (all except Hylaeus longiceps) are at risk of habitat destruction
and modification by feral pigs (see Table 2).
Goats (Capra hircus)
Feral goats currently destroy and modify habitat in 10 of the 11
ecosystems (coastal, lowland dry, lowland mesic, lowland wet, montane
wet, montane mesic, montane dry, subalpine, dry cliff, and wet cliff)
in which these species occur. Goats, native to the Middle East and
India, were successfully introduced to the Hawaiian Islands in the late
1700s. Actions to control feral goat populations began in the 1920s
(Tomich 1986, pp. 152-153). However, goats still occupy a wide variety
of habitats on all the main islands (except for Kahoolawe; see below),
where they consume native vegetation, trample roots and seedlings,
strip tree bark, accelerate erosion, and promote the invasion of
nonnative plants (van Riper and van Riper 1982, pp. 34-35; Stone 1985,
p. 261; Kessler 2010, pers. comm.). Kahoolawe was negatively affected
by ungulates beginning in 1793, with introduction of goats and the
addition of sheep (up to 15,000) and cattle (about 900) by ranchers
between 1858 and 1941, with the goat population estimated to be as high
as 50,000 individuals by 1988 (KIRC 2014, in litt.; KIRC 2015, in
litt.). Beginning in 1941, the U.S. military used the entire island as
a bombing range, and in 1994, control of Kahoolawe was returned to the
State and the Kahoolawe Island Reserve Commission. The remaining
ungulates were eradicated in 1993 (McLeod 2014, in litt.). Because they
are able to access extremely rugged terrain, and have a high
reproductive capacity (Clark and Cuddihy 1980, pp. C-19-C2-20; Culliney
1988, p. 336; Cuddihy and Stone 1990, p. 64), goats are believed to
have completely eliminated some plant species from certain islands
(Atkinson and Atkinson 2000, p. 21). Goats are
[[Page 67828]]
highly destructive to native vegetation and contribute to erosion by:
(1) Eating young trees and young shoots of plants before they become
established; (2) creating trails that damage native vegetative cover;
(3) destabilizing substrate and creating gullies that convey water; and
(4) dislodging stones from ledges that results in rockfalls and
landslides that damage or destroy native vegetation below (Cuddihy and
Stone 1990, pp. 63-64). Feral goats forage along some cliffs where
band-rumped storm-petrels nest on Kauai, and may trample nests and
increase erosion (Scott et al. 1986, pp. 8, 352-357; Tomich 1986, pp.
152-153). The band-rumped storm-petrel and the following 12 plants are
at risk from landslides or erosion caused by feral goats: Gardenia
remyi, Joinvillea ascendens ssp. ascendens, Kadua fluviatilis, Labordia
lorenciana, Lepidium orbiculare, Phyllostegia helleri, P. stachyoides,
Portulaca villosa, Pseudognaphalium sandwicensium var. molokaiense,
Ranunculus mauiensis, Sanicula sandwicensis, and Schiedea pubescens.
Twenty-four of the 39 plants (all except for Calamagrostis expansa,
Cyanea kauaulaensis, Cyclosorus boydiae, Cyperus neokunthianus, Deparia
kaalaana, Dryopteris glabra var. pusilla, Hypolepis hawaiiensis var.
mauiensis, Kadua haupuensis, Phyllostegia brevidens, Pritchardia
bakeri, Ranunculus hawaiensis, Schiedea diffusa ssp. diffusa, Sicyos
macrophyllus, Solanum nelsonii, and Stenogyne kaalae ssp. sherffii),
the band-rumped storm-petrel, the orangeblack Hawaiian damselfly, and
four of the yellow-faced bees (Hylaeus anthracinus, H. assimulans, H.
facilis, and H. hilaris) are at risk of habitat destruction and
modification by feral goats.
Axis Deer (Axis axis)
Axis deer destroy and modify 6 of the 11 ecosystems (coastal,
lowland dry, lowland mesic, lowland wet, montane wet, and montane
mesic) in which these species are found. Axis deer were introduced to
the Hawaiian Islands for hunting opportunities on Molokai in 1868, on
Lanai in 1920, and on Maui in 1959 (Hobdy 1993, p. 207; Erdman 1996,
pers. comm. in Waring 1996, in litt., p. 2; Hess 2008, p. 2). Axis deer
are primarily grazers, but also browse numerous palatable plant species
including those grown as commercial crops (Waring 1996, p. 3; Simpson
2001, in litt.). They prefer the low, openly vegetated areas for
browsing and grazing, but during episodes of drought (e.g., from 1998
to 2001 on Maui (Medeiros 2010, pers. comm.)), axis deer move into
urban and forested areas in search of food (Waring 1996, p. 5;
Nishibayashi 2001, in litt.). Like goats, axis deer are highly
destructive to native vegetation and contribute to erosion by eating
young trees and young shoots of plants before they can become
established. Other axis deer impacts include stripping bark from mature
trees, creating trails, promoting erosion by destabilizing substrate,
creating gullies that convey water, and dislodging stones from ledges
that can result in rockfalls and landslides that directly damage
vegetation (Cuddihy and Stone 1990, pp. 63-64).
On Molokai, axis deer likely occur at all elevations from sea level
to almost 5,000 ft (1,500 m) at the summit area (Kessler 2011, pers.
comm.). The most current population estimate for axis deer on the
island of Molokai is between 4,000 and 5,000 individuals (Anderson
2003, p. 119). Little management for deer control has been implemented
on Molokai, and this figure from more than a decade ago is likely an
underestimate of the axis deer population on this island today (Scott
et al. 1986, p. 360; Anderson 2003, p. 30; Hess 2008, p. 4). On Lanai,
axis deer were reported to number approximately 6,000 to 8,000
individuals in 2007 (The Aloha Insider 2008, in litt; WCities 2010, in
litt.). On Maui, five adult axis deer were released east of Kihei in
1959 (Hobdy 1993, p. 207; Hess 2008, p. 2). In 2013, the Maui Axis Deer
Working Group estimated that there may be 8,000 deer on southeast Maui
alone, based on helicopter surveys (Star Advertiser 2015, in litt.;
Hawaii News Now 2014, in litt.) According to Medeiros (2010, pers.
comm.) axis deer can be found in all but high-elevation ecosystems
(subalpine and alpine) and montane bogs on Maui, and are increasing in
numbers at such high rates that native forests are changing in
unprecedented ways. Additionally, Medeiros (2010, pers. comm.) asserted
that native plants will only survive in habitat that is fenced or
otherwise protected from the browsing and trampling effects of axis
deer. Kessler (2010, pers. comm.) and Hess (2010, pers. comm.) reported
the presence of axis deer up to 9,000 ft (2,700 m) on Maui, and Kessler
suggests that no ecosystem is safe from the negative impacts of these
animals. Montane bogs are also susceptible to impacts from axis deer.
As the native vegetation is removed by browsing and trampling, the soil
dries out, and nonnative plants invade. Eventually, the bog habitat and
its associated native plants and animals are replaced by grassland or
shrubland dominated by nonnative plants (Mitchell et al. 2005, p. 6-
32).
While axis deer are allowed as game animals on these three islands,
the State does not permit their introduction to other Hawaiian Islands.
In 2010-2011, axis deer were illegally introduced to Hawaii Island as a
game animal (Kessler 2011, pers. comm.; Aila 2012, in litt.), and deer
have now been observed across the southern portion of the island
including in Kohala, Kau, Kona, and Mauna Kea (HDLNR 2011, in litt.).
The Hawaii Department of Lands and Natural Resources (HDLNR) Division
of Forestry and Wildlife (HDOFAW) has developed a response-and-removal
plan, including a partnership now underway with the Hawaii Department
of Agriculture (HDOA), the Big Island Invasive Species Committee
(BIISC), Federal natural resource management agencies, ranchers,
farmers, private landowners, and concerned citizens (BigIsland.com,
June 6, 2011). Also, in response to the introduction of axis deer to
Hawaii Island, the Hawaii Invasive Species Council drafted House Bill
2593 to amend House Revised Statutes (H.R.S.) 91, which allows agencies
to adopt emergency rules in the instances of imminent peril to public
health, including to livestock and poultry health (BigIsland.com 2011,
in litt.; Martin 2012, in litt.). This emergency rule became permanent
on June 21, 2012, when House Bill 2593 was enacted into law as Act 194
(State of Hawaii 2012, in litt.).
The following 16 species in this rule are at risk from the
activities of axis deer: Gardenia remyi, Huperzia stemmermanniae,
Joinvillea ascendens ssp. ascendens, Nothocestrum latifolium,
Phyllostegia stachyoides, Portulaca villosa, Pseudognaphalium
sandwicensium var. molokaiense, Ranunculus mauiensis, Schiedea
pubescens, Solanum nelsonii, the orangeblack Hawaiian damselfly, and
five of the yellow-faced bees (Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, and H. longiceps).
Black-Tailed Deer (Odocoileus hemionus columbianus)
Black-tailed deer destroy and modify habitat in 5 of the 11
ecosystems (lowland mesic, lowland wet, montane wet, montane mesic, and
dry cliff) in which these species occur. The black-tailed deer is one
of nine subspecies of mule deer (Natural History Museum 2015, in
litt.). Black-tailed deer were first introduced to Kauai in 1961, for
the purpose of sport hunting (Tomich 1986, pp. 131-134). Currently,
these deer are only known from the western side of the island, where
they feed on a variety of
[[Page 67829]]
native (e.g., Acacia koa and Coprosma spp.) and nonnative plants (van
Riper and van Riper 1982, pp. 42-46; Tomich 1986, p. 134). In addition
to their direct impacts on native plants (browsing), black-tailed deer
likely affect native plants indirectly by serving as a primary vector
for the spread of introduced plants by carrying their seeds or other
propagules on their coats and hooves and in feces. Black-tailed deer
have been noted as a cause of habitat alteration in the Kauai
ecosystems (NTBG 2007, in litt.; HBMP 2010). Seven of the 39 plants
(Asplenium diellaciniatum, Dryopteris glabra var. pusilla, Joinvillea
ascendens ssp. ascendens, Labordia lorenciana, Nothocestrum latifolium,
Ranunculus mauiensis, and Sicyos lanceoloideus) are at risk of habitat
destruction and modification by black-tailed deer.
Sheep (Ovis aries)
Four of the ecosystems on Hawaii Island (lowland dry, lowland
mesic, montane mesic, and montane dry) in which these species occur are
currently threatened by habitat destruction and modification due to the
activities of feral sheep. Sheep were introduced to Hawaii Island in
1791, when Captain Vancouver brought five rams and two ewes from
California (Tomich 1986, pp. 156-163). Soon after, stock was brought
from Australia, Germany, and the Mediterranean for sheep production
(Tomich 1986, pp. 156-163; Cuddihy and Stone 1990, pp. 65-66), and by
the early 1930s, herds reached close to 40,000 individuals (Scowcroft
and Conrad 1992, p. 627). Capable of acquiring the majority of their
water needs by consuming vegetation, sheep can inhabit dry forests in
remote regions of the mountains of Mauna Kea and Mauna Loa, including
the saddle between the two volcanoes. Feral sheep browse and trample
native vegetation and have decimated large areas of native forest and
shrubland on Hawaii Island (Tomich 1986, pp. 156-163; Cuddihy and Stone
1990, pp. 65-66). Browsing results in the erosion of top soil that
alters moisture regimes and micro-environments, leading to the loss of
native plants and animals (Tomich 1986, pp. 156-163; Cuddihy and Stone
1990, pp. 65-66). In addition, nonnative plant seeds are dispersed into
native forest by adhering to sheep's wool coats (DOFAW 2002, p. 3). In
1962, game hunters intentionally crossbred feral sheep with mouflon
sheep and released them on Mauna Kea, where they have done extensive
damage to the montane dry ecosystem (Tomich 1986, pp. 156-163). Over
the past 30 years, attempts to protect the vegetation of Mauna Kea and
the saddle area between the two volcanoes have been only sporadically
effective (Hess 2008, pp. 1, 4). Currently, a large population of sheep
(and mouflon hybrids) extends from Mauna Kea into the saddle and
northern part of Mauna Loa, including State forest reserves, where they
trample and browse all vegetation, including endangered plants (Hess
2008, p. 1). One study estimated as many as 2,500 mouflon within just
the Kau district of the Kahuku Unit (Volcanoes National Park) in 2006
(Hess et al. 2006, p. 10). Two of the 39 plants, Exocarpos menziesii
and Festuca hawaiiensis, and the yellow-faced bee Hylaeus anthracinus,
are reported to be at risk of habitat destruction and modification by
feral sheep (see Table 2).
Mouflon (Ovis gmelini musimon)
Mouflon destroy and modify habitat in 6 of the 11 ecosystems on
Maui, Lanai, and Hawaii Island (lowland dry, lowland mesic, montane
mesic, montane dry, subalpine, and dry cliff) in which these species
occur. Native to central Asia, mouflon were introduced to the islands
of Lanai and Hawaii in the 1950s as game species, and are now widely
established on these islands (Tomich 1986, pp. 163-168; Cuddihy and
Stone 1990, p. 66; Hess 2008, p. 1). Due to their high reproductive
rate, the original population of 11 mouflon on the island of Hawaii
increased to more than 2,500 individuals in 36 years, even though they
were hunted for game (Hess 2008, p. 3). Mouflon have decimated vast
areas of native shrubland and forest through grazing, browsing, and
bark stripping (Stone 1985, p. 271; Cuddihy and Stone 1990, pp. 63, 66;
Hess 2008, p. 3). Mouflon also create trails and pathways through
vegetation, resulting in soil compaction and increased runoff and
erosion. In some areas, the interaction of browsing and soil compaction
has led to a shift from native forest to grassy scrublands (Hess 2008,
p. 3). Mouflon only gather in herds when breeding, thus complicating
control techniques and hunting efficiency (Hess 2008, p. 3; Ikagawa
2011, in litt.). Currently, many of the current and proposed fence
exclosures on Hawaii Island constructed to protect rare species and
habitat are designed to exclude feral pigs, goats, and sheep and are
only 4 ft (1.3 m) in height; a fence height of at least 6 ft (2 m) is
necessary to exclude mouflon (Ikagawa 2011, in litt.). Five of the 39
plant species (Exocarpos menziesii, Nothocestrum latifolium, Portulaca
villosa, Ranunculus hawaiensis, and Sicyos macrophyllus), and the
yellow-faced bee Hylaeus assimulans, are at risk from habitat
destruction and modification resulting from the activities of mouflon
(see Table 2).
Cattle (Bos taurus)
Cattle destroy and modify habitat in 7 of the 11 ecosystems on Maui
and Hawaii Island (coastal, lowland dry, lowland mesic, lowland wet,
montane wet, montane mesic, and montane dry) in which these species
occur. Cattle, the wild progenitors of which were native to Europe,
northern Africa, and southwestern Asia, were introduced to the Hawaiian
Islands in 1793, and large feral herds (as many as 12,000 on the island
of Hawaii) developed as a result of restrictions on killing cattle
decreed by King Kamehameha I (Cuddihy and Stone 1990, p. 40). While
small cattle ranches were developed on Kauai, Oahu, Molokai, west Maui,
and Kahoolawe, very large ranches of tens of thousands of acres were
created on east Maui and Hawaii Island (Stone 1985, pp. 256, 260;
Broadbent 2010, in litt.). Feral cattle can be found today on the
islands of Molokai, Maui, and Hawaii. Feral cattle eat native
vegetation, trample roots and seedlings, cause erosion, create
disturbed areas into which alien plants invade, and spread seeds of
alien plants carried in their feces and on their bodies. The forest in
areas grazed by cattle rapidly degrades into grassland pasture, and
plant cover remains reduced for many years following removal of cattle
from an area. Increased nitrogen availability through the feces of
cattle contributes to the ingress of nonnative plant species (Kohala
Mountain Watershed Partnership (KMWP) 2007, pp. 54-55; Laws et al.
2010, in litt.). Furthermore, several alien grasses and legumes
purposely introduced for cattle forage have become invasive weeds
(Tomich 1986, pp. 140-150; Cuddihy and Stone 1990, p. 29). According to
Kessler (2011, pers. comm.) approximately 300 individuals roam east
Maui as high as the subalpine ecosystem (i.e., to 9,800 ft (3,000 m)),
and feral cattle are occasional observed on west Maui. Feral cattle
(more than 100 individuals) are reported from remote regions of Hawaii
Island, including the back of Pololu and Waipio Valleys in the Kohala
Mountains, and the Kona Unit of the Hakalau Forest National Wildlife
Refuge (NWR) (KMWP 2007, p. 55; USFWS 2010, pp. 3-15, 4-86). Nine of
the 39 plant species (Huperzia stemmermanniae, Nothocestrum latifolium,
Ochrosia haleakalae, Portulaca villosa, Ranunculus hawaiensis, R.
mauiensis, Schiedea pubescens, Sicyos macrophyllus, and Solanum
nelsonii) and four of the
[[Page 67830]]
yellow-faced bees (Hylaeus anthracinus, H. assimulans, H. facilis, and
H. hilaris) are currently at risk of habitat destruction or
modification due to the activities of feral cattle.
In summary, 37 of the 39 plant species (all except Cyanea
kauaulaensis and Hypolepis hawaiiensis var. mauiensis), and 9 of the 10
animals (except for the anchialine pool shrimp Procaris hawaiana), are
at risk of habitat destruction and modification by ungulates including
pigs, goats, axis deer, black-tailed deer, sheep, mouflon, and cattle
(see Table 2). The effects of these nonnative animals include the
destruction of vegetative cover, trampling of plants and seedlings,
direct consumption of native vegetation, soil disturbance and
sedimentation (erosion and landslides), dispersal of nonnative plant
seeds by animals, alteration of soil nitrogen availability, and
creation of open, disturbed areas conducive to further invasion by
nonnative pest plant species. All of these impacts also can lead to the
conversion of a native plant community to one dominated by nonnative
species (see ``Habitat Destruction and Modification by Nonnative
Plants,'' below). In addition, because these animals inhabit terrain
that is often steep and remote, foraging and trampling contributes to
severe erosion of watersheds and degradation of streams and wetlands
(Cuddihy and Stone 1990, p. 59; Dunkell et al. 2011, pp. 175-194).
Habitat Destruction and Modification by Nonnative Plants
Ten of the 11 ecosystems (excluding anchialine pool ecosystem) and
the species in this rule that are associated with them are currently at
risk of habitat destruction and modification by nonnative plants.
Native vegetation on all of the main Hawaiian Islands has undergone
extreme alteration because of past and present land management
practices, including ranching, deliberate introduction of nonnative
plants and animals, and agriculture (Cuddihy and Stone 1990, pp. 27,
58). The original native flora of Hawaii (present before human arrival)
consisted of about 1,000 taxa, 89 percent of which are endemic (Wagner
et al. 1999, pp. 3-6). Over 800 plant taxa have been introduced to the
Hawaiian Islands. These were brought to Hawaii for food or for cultural
reasons, to reforest areas destroyed by grazing feral and domestic
animals, or for horticultural or agricultural purposes; some were
introduced unintentionally (Scott et al. 1986, pp. 361-363; Cuddihy and
Stone 1990, p. 73). Individual descriptions of 114 nonnative plant
species that negatively affect the 49 species are provided in our
proposed rule (80 FR 58820, September 30, 2015; see pp. 58869-58881).
Fourteen of these nonnative plants are included in the Hawaii Noxious
Weed List (Hawaii Department of Agriculture HAR 1981-title 4, subtitle
6, chapter 68).
Nonnative plants adversely affect native habitat in Hawaii by (1)
modifying the availability of light, (2) altering soil-water regimes,
(3) modifying nutrient cycling, and (4) altering fire regimes of native
plant communities (i.e., the ``grass/fire cycle'' that converts native-
dominated plant communities to nonnative plant communities; see below)
(Smith 1985, pp. 180-181; Cuddihy and Stone 1990, p. 74; D'Antonio and
Vitousek 1992, p. 73; Vitousek et al. 1997, p. 6). The contribution of
nonnative plants to the extinction of native species in the lowland and
upland habitats of Hawaii is well-documented (Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74). The most commonly observed effect of
nonnative plants on native species is displacement through competition.
Competition occurs for water or nutrients, or it may involve
allelopathy (chemical inhibition of growth of other plants), shading,
or precluding sites for seedling establishment (Vitousek et al. 1987 in
Cuddihy and Stone 1990, p. 74).
Alteration of fire regimes represents an ecosystem-level change
caused by the invasion of nonnative plants, primarily grasses
(D'Antonio and Vitousek 1992, p. 73). Grasses generate standing dead
material that burns readily, and grass tissues with large surface-to-
volume ratios dry out quickly, contributing to flammability (D'Antonio
and Vitousek 1992, p. 73). The finest size classes of grass material
ignite and spread fires under a broader range of conditions than do
woody fuels or even surface litter (D'Antonio and Vitousek 1992, p.
73). The grass life form allows rapid recovery following fire because
there is little above-ground vegetative structure. Grasslands also
support a microclimate in which surface temperatures are hotter,
contributing to drier vegetative conditions that favor fire (D'Antonio
and Vitousek 1992, p. 73). In summary, nonnative plants directly and
indirectly affect the 39 plants and 9 of the 10 animals in this rule
(except the anchialine pool shrimp) by destroying and modifying their
habitat, by removing their native host plants, or by direct
competition.
Habitat Destruction and Modification by Fire
Seven of the 11 ecosystems (coastal, lowland dry, lowland mesic,
montane mesic, montane dry, subalpine, and dry cliff) and the species
in this rule that are associated with them are at risk of destruction
and modification by fire. Fire is an increasing, human-exacerbated
threat to native species and ecosystems in Hawaii. The pre-settlement
fire regime in Hawaii was characterized by infrequent, low-severity
events, as few natural ignition sources existed (Cuddihy and Stone
1990, p. 91; Smith and Tunison 1992, pp. 395-397). It is believed that
prior to human colonization fuel was sparse in wet plant communities
and only seasonally flammable in mesic and dry plant communities. The
only ignition sources were volcanism and lightning (Baker et al. 2009,
p. 43). Although Vogl (1969, in Cuddihy and Stone 1990, p. 91) proposed
that naturally occurring fires may have been important in the
development of some of the original Hawaiian flora, Mueller-Dombois
(1981, in Cuddihy and Stone 1990, p. 91) asserts that most natural
vegetation types of Hawaii would not carry fire before the introduction
of alien grasses. Smith and Tunison (in Cuddihy and Stone 1990, p. 91)
state that native plant fuels typically have low flammability. Existing
fuel loads were often discontinuous, and rainfall in many areas on most
islands was moderate to high. Fires inadvertently or intentionally set
by the Polynesian settlers probably contributed to the initial decline
of native vegetation in the drier plains and foothills. These early
settlers practiced slash-and-burn agriculture that created open lowland
areas suitable for the opportunistic invasion and colonization of
nonnative, fire-adapted grasses (Kirch 1982, pp. 5-6, 8; Cuddihy and
Stone 1990, pp. 30-31). Beginning in the late 18th century, Europeans
and Americans introduced plants and animals that further degraded
native Hawaiian ecosystems. Ranching and the creation of pasturelands
in particular created highly fire-prone areas of nonnative grasses and
shrubs (D'Antonio and Vitousek 1992, p. 67). Although fires were
infrequent in mountainous regions, extensive fires have recently
occurred in lowland dry and lowland mesic areas, leading to grass/fire
cycles that convert native dry forest and native wet forest to
nonnative grassland (D'Antonio and Vitousek 1992, p. 77).
Because of the greater frequency, intensity, and duration of fires
that have resulted from the human alteration of landscapes and the
introduction of nonnative plants, especially grasses, fires are now
more destructive to native Hawaiian ecosystems (Brown and Smith 2000,
p. 172), and a single grass-fueled
[[Page 67831]]
fire often kills most native trees and shrubs in the area (D'Antonio
and Vitousek 1992, p. 74). Fire destroys dormant seeds of native
plants, as well as individual plants and animals themselves, even in
steep, inaccessible areas or near streams and ponds. Successive fires
remove habitat for native species by altering microclimate conditions,
creating conditions more favorable to nonnative plants. Nonnative
grasses (e.g., Cenchrus setaceus; fountain grass), many of which may be
fire-adapted, produce a high fuel load that allow fire to burn areas
that would not otherwise burn easily, regenerate quickly after fire,
and establish rapidly in burned areas (Fujioka and Fujii 1980 in
Cuddihy and Stone 1990, p. 93; D'Antonio and Vitousek 1992, pp. 70, 73-
74; Tunison et al. 2002, p. 122). Native woody plants may recover to
some degree, but fire tips the competitive balance toward nonnative
species (National Park Service 1989 in Cuddihy and Stone 1990, p. 93).
During a post-burn survey on Hawaii Island, in an area of native
Diospyros forest with undergrowth of the nonnative grass Pennisetum
setaceum [Cenchrus setaceus], Takeuchi (1991, p. 2) noted that ``no
regeneration of native canopy is occurring within the Puuwaawaa burn
area.'' Takeuchi also stated that ``burn events served to accelerate a
decline process already in place, compressing into days a sequence
which would ordinarily have taken decades'' (Takeuchi 1991, p. 4), and
concluded that, in addition to increasing the number of fires, the
nonnative Pennisetum acted to suppress establishment of native plants
after a fire (Takeuchi 1991, p. 6).
For many decades, fires have affected rare or endangered species
and their habitats on Molokai, Lanai, and Maui (Gima 1998, in litt.;
Hamilton 2009, in litt.; Honolulu Advertiser 2010, in litt.; Pacific
Disaster Center 2011, in litt.). These three islands experienced
approximately 1,290 brush fires between 1972 and 1999 that burned a
total of 64,250 ac (26,000 ha) (County of Maui 2009, ch. 3, p. 3;
Pacific Disaster Center 2011, in litt.). Between 2000 and 2003, the
annual number of wildfires on these islands jumped from 118 to 271; of
these, several burned more than 5,000 ac (2,023 ha) each (Pacific
Disaster Center 2011, in litt.). On Molokai, between 2003 and 2004,
three wildfires each burned 10,000 ac (4,050 ha) (Pacific Disaster
Center 2011, in litt.). From August through early September 2009, a
wildfire burned approximately 8,000 ac (3,237 ha), including 600 ac
(243 ha) of the remote Makakupaia section of the Molokai Forest
Reserve, a small portion of The Nature Conservancy's (TNC's) Kamakou
Preserve, and encroached on Onini Gulch, Kalamaula, and Kawela
(Hamilton 2009, in litt.). Species at risk because of wildfire on
Molokai include the plants Joinvillea ascendens ssp. ascendens,
Nothocestrum latifolium, Portulaca villosa, Ranunculus mauiensis,
Schiedea pubescens, and Solanum nelsonii, and the yellow-faced bees
Hylaeus anthracinus, H. facilis, H. hilaris, and H. longiceps.
Several wildfires have occurred on Lanai in the last decade. In
2006, a wildfire burned 600 ac (243 ha) between Manele Road and the
Palawai Basin, about 3 mi (4 km) south of Lanai City (The Maui News
2006, in litt.). In 2007, a brush fire at Mahana burned about 30 ac (12
ha), and in 2008, another 1,000 ac (405 ha) were burned by wildfire in
the Palawai Basin (The Maui News 2007, in litt.; KITV Honolulu 2008, in
litt.). Species at risk because of wildfire on Lanai include Exocarpos
menziesii, Nothocestrum latifolium, Portulaca villosa, Schidea
pubescens; and the yellow-faced bees Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H. longiceps.
On west Maui, wildfires burned more than 8,650 ac (3,501 ha)
between 2007 and 2010 (Honolulu Advertiser 2010, in litt.; Shimogawa
2010, in litt.). These fires encroached into the West Maui Forest
Reserve, on the ridges of Olowalu and Kealaloloa, which is habitat for
several endangered plants. In 2007, on east Maui, a fire consumed over
600 ac (240 ha), increasing invasion of the area by nonnative plants
(Pinus spp.) (Pacific Disaster Center 2007, in litt.; The Maui News
2011, in litt.). Species at risk because of wildfire on west and east
Maui include the plants Festuca hawaiiensis, Joinvillea ascendens ssp.
ascendens, Nothocestrum latifolium, Ochrosia haleakalae, Portulaca
villosa, Ranunculus mauiensis, Sanicula sandwicensis, Schiedea
pubescens, Sicyos macrophyllus, and Solanum nelsonii, and the yellow-
faced bees Hylaeus anthracinus, H. assimulans, H. facilis, H. hilaris,
and H. longiceps.
Several recent fires on Oahu in the Waianae Mountain range have
affected rare and endangered species. Between 2004 and 2005, wildfires
burned more than 360 ac (146 ha) in Honouliuli Preserve, habitat of
more than 90 rare and endangered plants and animals (TNC 2005). In
2006, a fire at Kaena Point State Park burned 60 ac (24 ha), and
encroached on endangered plants in Makua Military Training Area. In
2007, there was a significant fire at Kaukonahua that crossed 12
gulches, eventually encompassing 5,655 ac (2,289 ha) that negatively
affected eight endangered plant species and their habitat (Abutilon
sandwicense, Bonamia menziesii, Colubrina oppositifolia, Eugenia
koolauensis, Euphorbia haeleeleana, Hibiscus brackenridgei ssp.
mokuleianus, Nototrichium humile, and Schiedea hookeri) (U.S. Army
Garrison 2007b, Appendices pp. 1-5). This fire provided ingress for
nonnative ungulates (cattle, goats, and pigs) into previously
undisturbed areas, and opened dense native vegetation to the invasive
grass Urochloa maxima (Panicum maximum, guinea grass), also a food
source for cattle and goats. The grass was observed to generate blades
over 2 ft (0.6 m) in length only 2 weeks following the fire (U.S. Army
Garrison 2007b, Appendices pp. 1-5). In 2009, two smaller fires burned
200 ac (81 ha) at Manini Pali (Kaena Point State Park) and almost 4 ac
(1.5 ha) at Makua Cave. Both of these fires burned into area designated
as critical habitat, although no individual plants were directly
affected (U.S. Army Natural Resource Program 2009, Appendix 2, 17 pp.).
Most recently, in 2014, two fires affected native forest, one in the
Oahu Forest National Wildlife Refuge (350 ac, 140 ha), on the leeward
side of the Koolau Mountains (DLNR 2014, in litt.), and one above
Makakilo, in the Waianae Mountains, just below Honouliuli FR, that
burned more than 1,000 ac (400 ha) (KHON 2014, in litt.). The Makakilo
fire took over 2 weeks to contain. Species at risk because of wildfire
on Oahu include the plants Joinvillea ascendens ssp. ascendens,
Nothocestrum latifolium, Portulaca villosa, Ranunculus mauiensis, and
Sicyos lanceoloideus, and the yellow-faced bees Hylaeus anthracinus, H.
assimulans, H. facilis, H. kuakea, H. longiceps, and H. mana.
In 2012, on Kauai, a wildfire that was possibly started by an
unauthorized camping fire burned 40 ac (16 ha) in the Na Pali-Kona
Forest Reserve on Milolii Ridge, forcing closure of a hiking trail.
Fortunately, several endangered and threatened plants in the adjacent
Kula NAR were not impacted (KITV 2012, in litt.). The same year,
another wildfire burned over 650 ac (260 ha) on Hikimoe Ridge, and
threatened the Puu Ka Pele section of Waimea Canyon State Park (Hawaii
News Now 2012, in litt.; Star Advertiser 2012, in litt.). Species at
risk of because wildfire on Kauai include the plants Joinvillea
ascendens ssp. ascendens, Labordia lorenciana, Nothocestrum latifolium,
Ranunculus mauiensis, Santalum involutum, and Sicyos lanceoloideus.
[[Page 67832]]
In the driest areas on the island of Hawaii, wildfires are
exacerbated by the uncontrolled growth of nonnative grasses such as
Cenchrus setaceus (Fire Science Brief 2009, in litt.). Since its
introduction to the island in 1917, this grass now covers more than 200
square mi (500 square km) of the leeward areas of the island (Joint
Fire Science Brief (JFSB) 2009, in litt.). In the past 50 years, three
wildfires on the leeward side encompassed a total of 30,000 ac (12,140
ha) (JFSB 2009, in litt.). These wildfires traveled great distances at
rates of 4 to 8 miles per hour (mph) (7 to 12 kilometers per hour
(kph)), burning 2.5 ac (1 ha) to 6 ac (2.5 ha) per minute (the
equivalent of 6 to 8 football fields per minute) (Burn Institute 2009,
p. 4). Between 2002 and 2003, three successive lava-ignited wildfires
in the east rift zone of Hawaii Volcanoes National Park affected native
forests in lowland dry, lowland mesic, and lowland wet ecosystems (JFSB
2009, p. 3), cumulatively burning an estimated 11,225 ac (4,543 ha)
(Wildfire News, June 9, 2003; JFSP 2009, p. 3). These fires destroyed
over 95 percent of the canopy cover and encroached upon forest areas
that were previously thought to have low susceptibility to wildfires.
After the fires, nonnative ferns were observed in higher elevation
rainforest where they had not previously been seen, and were believed
to inhibit the recovery of the native Metrosideros polymorpha (ohia)
trees (JFSP 2003, pp. 1-2). Nonnative grasses invaded the burn area,
increasing the risk of fire encroaching into the surrounding native
forest (Ainsworth 2011, in litt.). Extreme drought conditions also
contributed to the number and intensity of wildfires on Hawaii Island
(Armstrong and Media 2010, in litt.; Loh 2010, in litt.). This
``extreme'' drought classification for Hawaii was recently lifted to
``moderate''; however, drier than average conditions persist, and
another extreme drought event may occur (NOAA 2015, in litt.). In
addition, El Ni[ntilde]o conditions in the Pacific (see ``Climate
Change'' under Factor E, below), a half-century of decline in annual
rainfall, and intermittent dry spells have contributed to the
conditions favoring wildfires in all the main Hawaiian Islands (Marcus
2010, in litt.). Species at risk because of wildfire on Hawaii Island
include the plants Exocarpos menziesii, Festuca hawaiiensis, Joinvillea
ascendens ssp. ascendens, Ochrosia haleakalae, Portulaca villosa,
Ranunculus mauiensis, Sanicula sandwicensis, Sicyos macrophyllus, and
Solanum nelsonii, and the yellow-faced bee Hylaeus anthracinus.
In summary, fire is a threat to 14 plant species and their habitat
(Exocarpos menziesii, Festuca hawaiiensis, Joinvillea ascendens ssp.
ascendens, Labordia lorenciana, Nothocestrum latifolium, Ochrosia
haleakalae, Portulaca villosa, Ranunculus mauiensis, Sanicula
sandwicensis, Santalum involutum, Schiedea pubescens, Sicyos
lanceoloideus, S. macrophyllus, and Solanum nelsonii), and all seven
yellow-faced bees because these species and their habitat are located
in or near areas that were burned previously, or in areas considered at
risk because of fire due to the cumulative and compounding effects of
drought and the presence of highly flammable nonnative grasses.
Habitat Destruction and Modification by Hurricanes
Ten of the 11 ecosystems (all except the anchialine pool ecosystem)
where these species occur are at risk of habitat destruction and
modification by hurricanes. Hurricanes exacerbate the impacts of other
threats such as habitat destruction and modification by ungulates and
competition with nonnative plants. By destroying native vegetation,
hurricanes open the forest canopy, modify the availability of light,
and create disturbed areas conducive to invasion by nonnative pest
species (see ``Habitat Destruction and Modification by Nonnative
Plants'', above) (Asner and Goldstein 1997, p. 148; Harrington et al.
1997, pp. 539-540). In addition, hurricanes adversely affect native
Hawaiian stream habitat by defoliating and toppling vegetation, thus
loosening the surrounding soil and increasing erosion. Along with
catastrophic flooding, this soil and vegetative debris can be washed
into streambeds (by hurricane-induced rain or subsequent rain storms),
resulting in the scouring of stream bottoms and channels (Polhemus
1993a, 88 pp.). Natural disasters such as hurricanes can be
particularly devastating to Hawaiian plant and animal species that
persist in low numbers and in restricted ranges (Mitchell et al. 2005,
p. 4-3).
Hurricanes affecting Hawaii were only rarely reported from ships in
the area from the 1800s until 1949. Between 1950 and 1997, 22
hurricanes passed near or over the Hawaiian Islands, 5 of which caused
serious damage (Businger 1998, pp. 1-2). In November 1982, Hurricane
Iwa struck the Hawaiian Islands with wind gusts exceeding 100 miles per
hour (mph) (160 kilometers per hour (kmh)), causing extensive damage,
especially on the islands of Kauai, Niihau, and Oahu (Businger 1998,
pp. 2, 6). Many forest trees were destroyed (Perlman 1992, pp. 1-9),
which opened the canopy and facilitated the invasion of native forest
by nonnative plants (Kitayama and Mueller-Dombois 1995, p. 671).
Hurricanes therefore exacerbate the threats posed by nonnative plants,
as described in ``Habitat Destruction and Modification by Nonnative
Plants,'' above. In September 1992, Hurricane Iniki, a category 4
hurricane with maximum sustained winds of 130 mph (209 kmh, 113 knots),
passed directly over the island of Kauai and close to the island of
Oahu, causing significant damage to Kauai and along Oahu's southwestern
coast (Blake et al. 2007, pp. 20, 24). Biologists documented damage to
the habitat of six endangered plant species on Kauai, and one plant on
Oahu. Polhemus (1993a, pp. 86-87) documented the extirpation of the
scarlet Kauai damselfly (Megalagrion vagabundum) (a species related to
M. xanthomelas), from the entire Hanakapiai Stream system on the island
of Kauai as a result of the impacts of Hurricane Iniki. Damage by
future hurricanes will further alter the remaining native-plant
dominated habitat for rare plants and animals in native ecosystems of
Kauai, Oahu, and other Hawaiian Islands (Bellingham et al. 2005, p.
681) (see ``Climate Change'' under Factor E. Other Natural or Manmade
Factors Affecting Their Continued Existence, below).
In summary, hurricanes exacerbate other habitat threats, such as
competition with nonnative plants, as well as result in direct habitat
destruction. This is a particular problem for the plant Pritchardia
bakeri, the band-rumped storm-petrel, the orangeblack Hawaiian
damselfly, and all seven yellow-faced bees.
Habitat Modification and Destruction Due to Landslides, Rockfalls,
Treefall, Flooding, Erosion, Drought, and Tsunamis
Habitat destruction and modification by landslides, rockfalls,
treefall, flooding, erosion, and drought (singly or in combination) is
a threat to all 11 ecosystems in which these species occur. Landslides,
rockfalls, treefall, flooding, and erosion change native plant and
animal communities by destabilizing substrates, damaging or destroying
individual plants, and altering hydrological patterns. In the open sea
near Hawaii, rainfall averages 25 to 30 inches (in) (630 to 760
millimeters (mm)) per year, yet the islands may receive up to 15 times
this amount in some places, caused by
[[Page 67833]]
orographic features (topography) (Wagner et al. 1999, adapted from
Price (1983) and Carlquist (1980), pp. 38-39). During storms, rain may
fall at rates of 3 in (76 mm) per hour or more, and sometimes may reach
nearly 40 in (1,000 mm) in 24 hours, resulting in destructive flash-
flooding in streams and narrow gulches (Wagner et al. 1999, adapted
from Price (1983) and Carlquist (1980), pp. 38-39). Due to the steep
topography in many mountainous areas on the Hawaiian Islands,
disturbance caused by introduced ungulates exacerbates erosion and
increases the potential for landslides, rockfalls, or flooding, which
in turn damages or destroys native plants and disturbs habitat of the
band-rumped storm-petrel (see Table 2). These events could eliminate
one or more isolated occurrences of species that persist in low numbers
and a limited geographic range, resulting in reduced redundancy and
resilience of the species.
Landslides, rockfalls, treefall, flooding, and erosion are threats
to 20 plant species (Cyanea kauaulaensis, Cyclosorus boydiae, Deparia
kaalaana, Dryopteris glabra var. pusilla, Gardenia remyi, Joinvillea
ascendens ssp. ascendens, Kadua fluviatilis, K. haupuensis, Labordia
lorenciana, Lepidium orbiculare, Phyllostegia brevidens, P. helleri, P.
stachyoides, Portulaca villosa, Pseudognaphalium sandwicensium var.
molokaiense, Ranunculus hawaiensis, R. mauiensis, Sanicula
sandwicensis, Schiedea pubescens, and Solanum nelsonii) and to the
band-rumped storm-petrel and the orangeblack Hawaiian damselfly.
Landslides, rockfalls, and erosion can directly affect nests and
nesting habitat of the band-rumped storm-petrel. Destabilization of
cliff habitat leads to additional landslides and alteration of
hydrological patterns, affecting the availability of soil moisture.
Landslides also destroy and modify riparian and stream habitat by
direct physical damage, and create disturbed areas leading to invasion
by nonnative plants, as well as damaging or destroying plants directly.
Kadua haupuensis, Labordia lorenciana, Lepidium orbiculare,
Phyllostegia brevidens, and P. helleri are known only from a few
individuals in single occurrences on cliffs or steep-walled stream
valleys, and one landslide could extirpate a species by direct
destruction. Monitoring data presented by the Plant Extinction
Prevention Program (PEPP) and botanical surveys suggest that flooding
is a likely threat to eight plant species, Cyanea kauaulaensis,
Cyclosorus boydiae, Deparia kaalaana, Labordia lorenciana, Phyllostegia
stachyoides, Sanicula sandwicensis, Schiedea pubescens, and Solanum
nelsonii, as some individuals occur on stream banks (Wood et al. 2007,
p. 198; PEPP 2011, pp. 162-164; Oppenheimer and Lorence 2012, pp. 20-
21; PEPP 2013, p. 54; PEPP 2014, pp. 95, 142). The naiad life stage of
the orangeblack Hawaiian damselfly would be destroyed by flooding if an
individual is carried out of suitable habitat or into areas occupied by
nonnative fish.
Drought is reported to be a threat to 10 plants (Cyclosorus
boydiae, Deparia kaalaana, Huperzia stemmermanniae, Phyllostegia
stachyoides, Ranunculus hawaiensis, R. mauiensis, Sanicula
sandwicensis, Schiedea pubescens, Sicyos lanceoloideus, and Solanum
nelsonii), the orangeblack Hawaiian damselfly (directly or by
desiccation of streams and ponds), and all seven yellow-faced bees
(Magnacca 2007, pp. 181, 183; Polhemus 2008, p. 26; Chu et al. 2010,
pp. 4887, 4891, 4898; PEPP 2011, pp. 162-164; Fortini et al. 2013, p.
2; PEPP 2013, p. 177; PEPP 2014, pp. 140-142, 154-156, 162, 166-167).
Between 1860 and 2002, there were 49 periods of drought on Oahu, 30
periods of drought on Molokai, Lanai, and Maui, and at least 18 serious
or severe drought events on Hawaii Island (Giambelluca et al. 1991, pp.
3-4; Hawaii Commission on Water Resource Management (CWRM) 2009, in
litt.; Hawaii Civil Defense 2011, pp. 14-1-14-12). The most severe
drought events over the past 15 years were associated with the El
Ni[ntilde]o phenomenon (Hawaii Civil Defense 2011, p. 14-3). In 1998,
the city of Hilo had the lowest January total rainfall (0.014 in) ever
observed for any month since records have been kept, with average
rainfall being almost 10 in for January (Hawaii Civil Defense 2011, p.
14-3). Currently, the State remains under abnormally dry to moderate
drought conditions, with the onset of another El Ni[ntilde]o event
(U.S. Drought Monitor 2015, in litt., National Weather Service 2015, in
litt.). Drought events dry up streams, irrigation ditches, and
reservoirs, and deplete groundwater supplies (Hawaii CWRM 2009, in
litt.). Recent episodes of drought have driven axis deer farther into
forested areas in search of food, increasing their negative impacts on
native vegetation from herbivory, bark stripping, and trampling (see
Factor C. Disease or Predation, below) (Waring 1996, in litt;
Nishibayashi 2001, in litt.). Drought events could eliminate one or
more isolated populations of a species that currently persists in low
numbers and a limited geographic range, resulting in reduced redundancy
and resilience of the species or extinction.
Tsunamis destroy and modify habitat for species in Northwestern
Hawaiian Islands and in low-lying coastal areas of the main Hawaiian
Islands. Tsunamis in Hawaii are caused by earthquakes, submarine
landslides, and volcanic eruptions that may occur within the
archipelago or in distant parts of the Pacific. These events disturb
the ocean's surface, and gravity combined with the water's motion
produces a series of long-period waves that travel quickly and can
reach heights of 32 ft (10 m) or more when reaching land. Major
tsunamis occur worldwide about once every 10 years, on average, and
almost 60 percent of those occur in the Pacific Ocean (Pacific Tsunami
Warning Center, https://ptwc.weather.gov/ptwc/faq.php#8, accessed June
2016). In 2011, a tsunami caused by an earthquake in Japan reached
Hawaii and the Northwestern Hawaiian Islands. This tsunami swept over
Midway Atoll's Eastern Island and Kure Atoll's Green Island, where it
inundated plants, spread plastic debris, killed thousands of seabirds,
and destroyed seabird nesting areas as it traveled about 500 ft (150 m)
inland (DOFAW 2011, in litt.; Starr 2011, in litt.; USFWS 2011, in
litt.). This threat could occur at any time and negatively affect
occurrences and habitat of the plant Solanum nelsonii and the yellow-
faced bees Hylaeus anthracinus, H. assimulans, H. facilis, H. hilaris,
and H. longiceps.
Habitat Destruction and Modification by Water Extraction
Freshwater habitats on all the main Hawaiian Islands have been
severely altered and degraded because of past and present land and
water management practices, including agriculture, urban development,
and development of ground water, perched aquifer, and surface water
resources (Harris et al. 1993, p. 11; Meier et al. 1993, p. 181).
Extensive modification of lentic (standing water) habitat in the
Hawaiian Islands began about 1100 A.D. with a rapid increase in the
human population (Harris et al. 1993, p. 9; Kirch 1982, pp. 5-6).
Hawaiians cultivated Colocasia esculenta (kalo, taro) by creating
shallow, walled ponds, or loi, in marshes and riparian areas (Meier et
al. 1993, p. 181; Handy and Handy 1972, p. 58). By 1778, virtually all
valley bottoms with permanent stream flow and most basin marshes were
converted to taro cultivation (Handy and Handy 1972, pp. 396, 411).
Hawaiians also modified wetlands by constructing fishponds, many of
which were primarily fresh water, fed by streams or springs (Meier
[[Page 67834]]
et al. 1993, p. 181). Despite this habitat modification by early
Hawaiians, many areas of extensive marshland remained intact and were
used by the native damselflies. Over time, however, many of the
wetlands formerly used for taro were drained and filled for dry-land
agriculture or development (Stone 1989, p. 129; Meier et al. 1993, pp.
181-182). In addition, marshes are slowly filled and converted to
meadow habitat due to increased sedimentation resulting from increased
storm water runoff from upslope development and blockage of downslope
drainage (Wilson Okamoto and Associates, Inc. 1993, p. 3-5). Presently
the most significant threat to the remaining natural ponds and marshes
in Hawaii, habitat for the orangeblack Hawaiian damselfly, is the
nonnative grass species Urochloa mutica. This sprawling, perennial
grass was first observed on Oahu in 1924, and now occurs on all the
main islands (O'Connor 1999, p. 1504). This species forms dense,
monotypic stands that can completely eliminate any open water by
layering of its trailing stems (Smith 1985, p. 186).
Similar to the loss of wetlands in Hawaii, the loss of streams has
been significant and began with the early Hawaiians who modified stream
systems by diverting water to irrigate taro. However, these Hawaiian-
made diversions were closely regulated and were not permitted to take
more than half the stream flow, and were typically used to flood taro
loi only periodically (Handy and Handy 1972, pp. 58-59). The advent of
sugarcane plantations in 1835 led to more extensive stream diversions.
These systems were typically designed to tap water at upper elevation
sources (above 980 ft (300 m)) by means of concrete weirs. All or most
of the stream flow was diverted into fields or reservoirs (Takasaki et
al. 1969, p. 65; Harris et al. 1993, p. 10). By the 1930s, major water
diversions had been developed on all the main islands, and currently
one-third of Hawaii's perennial streams are diverted (Harris et al.
1993, p. 10). In addition to diverting water for agriculture and
domestic water supply, streams have been diverted for use in producing
hydroelectric power (Hawaii Stream Assessment 1990, p. 96). Surface
flow has also been diverted into channels, and the perched aquifers
which fed the streams have been tapped by means of tunnels (Stearns and
Vaksvik 1935, pp. 365, 378-434; Stearns 1985, p. 291, 301-303). Many of
these aquifers are the sources of springs, which contribute flow to
streams. The draining of these aquifers causes springs to become dry
(Stearns and Vaksvik 1935, pp. 380, 388; USGS 2000, in litt.). Most
remaining streams that are not already diverted have been, and continue
to be, seriously degraded by the activities of feral ungulates and by
nonnative plants. Channelization has not been restricted to lower
reaches, and it results in the loss of riparian vegetation, increasing
flow velocity, illumination, and water temperature (Parrish et al.
1984, pp. 83-84). These conditions make the channels unsuitable as
habitat for the orangeblack Hawaiian damselfly.
Water extraction (e.g., withdrawal of subsurface fresh water for
development and human use) from underground fresh water sources
increases salinity levels of anchialine pools and negatively affect the
anchialine pool shrimp, Procaris hawaiana, which relies on the delicate
balance of mixohaline (brackish water) habitats (Conry 2012, in litt.;
National Park Service 2016, in litt.). Water extraction also negatively
affects the plant Cyclosorus boydiae and the orangeblack Hawaiian
damselfly by degrading or destroying habitat for these species (Harris
et al. 1993, pp. 9-13; Medeiros et al. 1993, p. 88; Meier et al. 1993,
pp. 181-183; Palmer 2003, p. 88).
Habitat Destruction and Modification by Climate Change
Climate change affects the habitat of the 49 species. Discussion of
climate change impacts is included in our complete discussion of
climate change under Factor E. Other Natural or Manmade Factors
Affecting Their Continued Existence, below.
Summary of Factor A
Destruction and modification of the habitat of each of the 49
species addressed in this rule is occurring throughout the entire range
of each of the species. These impacts include the effects of
agriculture and urban development, introduced ungulates, nonnative
plants, fire, hurricanes, landslides, rockfalls, treefall, flooding,
erosion, drought, tsunamis, and water extraction.
Habitat destruction and modification by agriculture and urban
development is an ongoing and serious threat to the plant Cyclosorus
boydiae, the orangeblack Hawaiian damselfly, the anchialine pool shrimp
Procaris hawaiana, and the yellow-faced bees Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H. longiceps. Conversion of
wetland and other aquatic habitat (i.e., water extraction) for
agriculture and urban development is ongoing, is expected to continue
into the future, and affects the orangeblack Hawaiian damselfly by
removing habitat required for hunting and breeding. Water extraction
affects the orangeblack Hawaiian damselfly because it (1) Reduces the
amount and distribution of stream habitat; (2) reduces stream flow and
habitat; and (3) leads to an increase in water temperature, which
causes physiological stress to the damselfly naiads. Water extraction
affects the delicate balance of the anchialine pool ecosystem,
including salinity and biota, negatively affecting the anchialine pool
shrimp, Procaris hawaiana. Loss of stream-course habitat affects
Cyclosorus boydiae because this is the only habitat where this riparian
species occurs.
The threat of habitat destruction and modification by ungulates is
ongoing as ungulates currently occur in all ecosystems on which these
species depend except the anchialine pool system. Introduced ungulates
pose a threat to 37 of the 39 plants (except for Cyanea kauaulaensis
and Hypolepis hawaiiensis var. mauiensis), and 9 of the 10 animal
species (all except for the anchialine pool shrimp) in this rule that
occur in these 10 ecosystems because ungulates: (1) Directly affect the
species by trampling and grazing (see Factor C discussion, below); (2)
increase soil disturbance and erosion; (3) create open, disturbed areas
conducive to nonnative plant invasion by dispersing fruits and seeds,
which results in conversion of a native-dominated plant community to a
nonnative-dominated plant community; and (4) increase marsh and stream
disturbance and sedimentation, which negatively affects these aquatic
habitats.
Habitat destruction and modification by nonnative plants is a
serious and ongoing current threat to all 39 plant species because
nonnative plants: (1) Adversely affect microhabitat by modifying the
availability of light; (2) alter soil-water regimes; (3) modify
nutrient cycling processes; (4) alter fire ecology, leading to
incursions of fire-tolerant nonnative plant species into native
habitat; (5) outcompete, and possibly directly inhibit (through
allelopathy) the growth of native plant species; and (6) alter habitat
and substrate such that erosion leading to rockfalls and landslides may
increase. Each of these processes can convert native-dominated plant
communities to nonnative plant communities (Cuddihy and Stone 1990, p.
74; Vitousek 1992, pp. 33-35).
The threat of habitat destruction and modification by fire to 14
plant species (Exocarpos menziesii, Festuca hawaiiensis, Joinvillea
ascendens ssp. ascendens, Labordia lorenciana, Nothocestrum latifolium,
Ochrosia haleakalae, Portulaca villosa, Ranunculus mauiensis, Sanicula
[[Page 67835]]
sandwicensis, Santalum involutum, Schiedea pubescens, Sicyos
lanceoloideus, S. macrophyllus, and Solanum nelsonii) and all seven
yellow-faced bee species is serious and ongoing because fires occur
frequently and damage and destroy native vegetation, including dormant
seeds, seedlings, and juvenile and adult plants, including host plants
for the bees. Many nonnative, invasive plants, particularly fire-
tolerant grasses, create more destructive fires, invade burned areas,
and can out-compete native plants and inhibit their regeneration
(D'Antonio and Vitousek 1992, pp. 70, 73-74; Tunison et al. 2002, p.
122). Successive fires that burn farther and farther into native
habitat destroy the ecosystem and its components upon which these
species depend.
Habitat destruction and modification by natural disasters such as
hurricanes represent a serious threat to the plant Pritchardia bakeri,
the band-rumped storm-petrel, the orangeblack Hawaiian damselfly, and
all seven yellow-faced bee species. Hurricanes open the forest canopy,
modifying available light and creating disturbed areas that are
conducive to invasion by nonnative plants (Asner and Goldstein 1997, p.
148; Harrington et al. 1997, pp. 346-347). The discussion under
``Habitat Destruction and Modification by Nonnative Plants'' provides
additional information related to canopy gaps, light availability, and
the establishment of nonnative plant species. In addition, hurricanes
cause mortality of birds, including adults and chicks drowned when nest
sites are flooded (Schreiber 2002, p. 186; Hass et al. 2012, pp. 252-
253). Hurricanes also destroy nesting habitat, a particular problem for
species like storm-petrels that return to the same nest site each year
(Schreiber 2002, p. 186). These hurricane impacts are likely for the
band-rumped storm-petrel. Finally, hurricanes can alter and directly
damage streams and wetlands used by the orangeblack Hawaiian damselfly
(Polhemus 1993a, pp. 86-87). The impacts from hurricanes can be
particularly devastating to these species because they persist in low
numbers in restricted ranges and are therefore less resilient to such
disturbances. A single destructive hurricane holds the potential of
driving to extinction species that persist as one or several small,
isolated populations.
Landslides, rockfalls, treefalls, flooding, and erosion (singly or
combined) are a threat to 20 plant species (Cyanea kauaulaensis,
Cyclosorus boydiae, Deparia kaalaana, Dryopteris glabra var. pusilla,
Gardenia remyi, Joinvillea ascendens ssp. ascendens, Kadua fluviatilis,
K. haupuensis, Labordia lorenciana, Lepidium orbiculare, Phyllostegia
brevidens, P. helleri, P. stachyoides, Portulaca villosa,
Pseudognaphalium sandwicensium var. molokaiense, Ranunculus hawaiensis,
R. mauiensis, Sanicula sandwicensis, Schiedea pubescens, and Solanum
nelsonii), the band-rumped storm-petrel, and the orangeblack Hawaiian
damselfly by destabilizing substrates, damaging and killing
individuals, altering hydrological patterns, and destroying or
modifying habitat--all resulting in changes to native plant and animal
communities. Drought is a threat to 10 plant species (Cyclosorus
boydiae, Deparia kaalaana, Huperzia stemmermanniae, Phyllostegia
stachyoides, Ranunculus hawaiensis, R. mauiensis, Sanicula
sandwicensis, Schiedea pubescens, Sicyos lanceoloideus, and Solanum
nelsonii), the orangeblack Hawaiian damselfly (directly or by
desiccation of streams and ponds), and all seven yellow-faced bee
species (and the host plants upon which all seven yellow-faced bees
depend).
Habitat destruction and modification by over-washing of low-lying
areas by tsunamis is a threat to coastal species, including Solanum
nelsonii, Hylaeus anthracinus, H. assimulans, H. facilis, H. hilaris,
and H. longiceps.
Factor B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Plants
We are not aware of any threats to the 39 plant species that would
be attributed to overutilization for commercial, recreational,
scientific, or educational purposes.
Animals
Anchialine Pool Shrimp
Illegal collection is a threat to the anchialine pool shrimp
Procaris hawaiana because of inadequate monitoring and enforcement at
the pools where this species occurs. All terrestrial and aquatic
invertebrates (including anchialine pool shrimp) are protected under
(1) the State of Hawaii Revised Statutes (1993) chapter 195D-4-f
license; and (2) DLNR chapter 124: Indigenous Wildlife, Endangered and
Threatened Wildlife, and Introduced Wild Birds. Collection of plants
and animals is prohibited in the State Natural Area Reserves (NARs)
Ahihi-Kinau (Maui) and Manuka (Hawaii Island), but enforcement of
prohibitions is insufficient to prevent illegal collecting at these
remote sites. Collection is not prohibited in State Parks or City and
County property where some anchialine pools occur, and is not expressly
prohibited at Lua O Palahemo (Hawaii Island), and thus no regulatory
protection of these shrimp exists at the remaining five anchialine
pools outside of Manuka NAR that are known to contain P. hawaiana. A
Native Invertebrate Research and Collecting permit issued by DLNR's
Division of Forestry and Wildlife is required to collect anchialine
pool shrimp for research or commercial purposes, and the commercial
market is supported by legal, permitted collection. We expect that
permit holders, whether they are collecting for scientific or
commercial purposes, adhere to the conditions of their permit and do
not pose a threat to P. hawaiana. However, we consider illegal
collection of this anchialine pool shrimp, P. hawaiana, to be an
ongoing threat because, despite the prohibition on collecting within
the NARs and the permitting process for collection elsewhere,
collection can occur at any time owing to insufficient patrolling or
other monitoring or enforcement at the pools where P. hawaiana occurs.
Factor C. Disease or Predation
Disease
We are not aware of any current threats to the 49 species that
would be attributable to disease. Disease may be a potential threat to
the yellow-faced bee Hylaeus anthracinus, as pathogens carried by
nonnative bees, wasps, and ants could be transmitted through shared
food sources (Graham 2015, in litt.); however, we have no evidence of
this type of disease transmission at this time.
Predation
Hawaii's plants and animals evolved in nearly complete isolation
from continental influence. Successful, natural colonization of these
remote volcanic islands is infrequent, and many organisms never
succeeded in establishing populations. As an example, Hawaii lacks
native ants and conifers, has very few families of birds, and has only
had two native species of land mammal, both insectivorous bats (Loope
1998, p. 748; Ziegler 2002, pp. 244-245). In the absence of grazing or
browsing mammals, plants that became established did not need
mechanical or chemical defenses against mammalian herbivory such as
thorns, prickles, and toxins. Because the evolutionary pressure to
either produce or maintain such defenses was lacking, Hawaiian plants
either lost or never developed these adaptations (Carlquist 1980, p.
173). Likewise, native Hawaiian birds
[[Page 67836]]
and insects experienced no evolutionary pressure to develop defense
mechanisms against mammalian or invertebrate predators that were not
historically present on the islands. The native flora and fauna are
thus particularly vulnerable to the impacts of introduced nonnative
species, as discussed below.
Introduced Ungulates
In addition to the habitat impacts discussed above (see ``Habitat
Destruction and Modification by Introduced Ungulates'' under Factor A.
The Present or Threatened Destruction, Modification, or Curtailment of
Their Habitat or Range), grazing and browsing (predation) by introduced
ungulates are a threat to the following 27 plant species in this
proposal (see Table 2, above): Asplenium diellaciniatum (black-tailed
deer); Calamagrostis expansa (pigs), Cyclosorus boydiae (pigs), Deparia
kaalaana (pigs), Exocarpos menziesii (goats, mouflon, sheep), Festuca
hawaiiensis (goats, sheep), Gardenia remyi (pigs, goats, axis deer),
Huperzia stemmermanniae (goats, axis deer, cattle), Joinvillea
ascendens ssp. ascendens (pigs, goats, axis deer, black-tailed deer),
Kadua fluviatilis (pigs, goats), Labordia lorenciana (goats, black-
tailed deer), Microlepia strigosa var. mauiensis (pigs), Myrsine
fosbergii (pigs, goats), Nothocestrum latifolium (pigs, goats, axis
deer, black-tailed deer, mouflon, cattle), Ochrosia haleakalae (goats,
cattle), Phyllostegia brevidens (pigs), P. stachyoides (pigs, goats),
Portulaca villosa (goats, axis deer, mouflon), Pseudognaphalium
sandwicensium var. molokaiense (axis deer), Ranunculus hawaiensis
(pigs, mouflon, cattle), R. mauiensis (pigs, goats, axis deer, black-
tailed deer, cattle), Sanicula sandwicensis (goats), Santalum involutum
(goats), Schiedea pubescens (axis deer, cattle), Sicyos lanceoloideus
(goats, black-tailed deer), S. macrophyllus (mouflon, cattle), and
Solanum nelsonii (axis deer, cattle).
Feral Pigs
We have direct evidence of ungulate damage to some of the 39 plant
species, but for many, due to their remote locations or lack of study,
ungulate damage is presumed based on the known presence of these
introduced ungulates in the areas where these species occur and the
results of studies involving similar species or ecosystems conducted in
Hawaii and elsewhere (Diong 1982, p. 160; Mueller-Dombois and Spatz,
1975, pp. 1-29; Hess 2008, 4 pp.; Weller et al. 2011, p. 8). For
example, in a study conducted by Diong (1982, p. 160) on Maui, feral
pigs were observed browsing on young shoots, leaves, and fronds of a
wide variety of plants, of which over 75 percent were endemic species.
A stomach-content analysis in this study showed that most of the pigs'
food source consisted of the endemic Cibotium (hapuu, tree fern). Pigs
were observed to fell native plants and remove the bark from standing
plants of species in the genera Cibotium, Clermontia, Coprosma,
Hedyotis [Kadua], Psychotria, and Scaevola, resulting in larger trees
and shrubs dying after a few months of repeated feeding (Diong 1982, p.
144). Beach (1997, pp. 3-4) found that feral pigs in Texas spread
disease and parasites, and their rooting and wallowing behavior led to
spoilage of watering holes and loss of soil through leaching and
erosion. Rooting activity by pigs also decreased the survivability of
some plant species through disruption at root level of mature plants
and seedlings (Beach 1997, pp. 3-4; Anderson et al. 2007, in litt.). In
Hawaii, pigs dig up forest ground cover consisting of delicate and rare
species of orchids, ferns, mints, lobeliads, and other taxa, including
their roots, tubers, and rhizomes (Stone and Anderson 1988, p. 137).
The following plants are particularly at risk of herbivory by feral
pigs: Calamagrostis expansa on Maui and Hawaii Island (HBMP 2010);
Cyclosorus boydiae on Oahu (HBMP 2010); Deparia kaalaana on Maui (HBMP
2010); Gardenia remyi on Hawaii Island (PEPP 2011, pp. 113-114; PEPP
2012, p. 102), west Maui (HBMP 2010), Molokai (HBMP 2010) and Kauai
(HBMP 2010); Joinvillea ascendens ssp. ascendens on Hawaii Island (PEPP
2011, pp. 120-121; PEPP 2012, p. 113; HBMP 2010), Kauai (PEPP 2014, p.
109; HBMP 2010), Maui (HBMP 2010), Molokai (HBMP 2010), and Oahu (HBMP
2010); Kadua fluviatilis on Kauai (HBMP 2010) and Oahu (HBMP 2010);
Microlepia strigosa var. mauiensis on Maui (Bily 2009, in litt.;
Oppenheimer 2007, in litt.); Myrsine fosbergii on Kauai (HBMP 2010);
Nothocestrum latifolium on Maui (PEPP 2011, p. 140; HBMP 2010) and
Molokai (HBMP 2010); Phyllostegia brevidens on Maui and Hawaii Island
(PEPP 2014, p. 36); P. stachyoides on Molokai (PEPP 2014, pp. 140-141);
Ranunculus hawaiensis on Hawaii Island (HBMP 2010); and R. mauiensis on
Kauai (PEPP 2011, p. 161; PEPP 2013, p. 177; PEPP 2014, p. 156; HBMP
2010), Maui (PEPP 2011, p. 144; PEPP 2013, pp. 177-178; PEPP 2014, p.
155; HBMP 2010), and Molokai (HBMP 2010). Feral pigs occur in 10 of the
11 ecosystems (all except anchialine pool) discussed here; the results
of the studies described above suggest that foraging by pigs can
directly damage and destroy these plants through herbivory. Feral pigs
may also consume native host plants of the yellow-faced bees Hylaeus
anthracinus, H. assimulans, H. facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana.
Feral Goats
Feral goats are able to forage in extremely rugged terrain and are
instrumental in the decline of native vegetation in many areas of the
Hawaiian Islands (Cuddihy and Stone 1990, p. 64; Clarke and Cuddihy
1980, p. C-20; van Riper and van Riper 1982, pp. 34-35; Tomich 1986,
pp. 153-156). Feral goats consume a variety of plants for food and have
been observed to browse on (but are not limited to) native plant
species in the following genera: Argyroxiphium, Canavalia, Chamaesyce,
Erythrina, Plantago, Schiedea, and Stenogyne (Cuddihy and Stone 1990,
p. 64; Warren 2004, p. 462; Wood 2007, pers. comm.). A study conducted
on the island of Hawaii demonstrated that native Acacia koa seedlings
are unable to survive due to browsing and grazing by goats (Spatz and
Mueller-Dombois 1973, p. 874). If goats remained in the area in high
numbers, mature trees eventually died and with them the root systems
that supported suckers and vegetative reproduction. When feral goats
were excluded by fences for 3 years, there was a positive height-growth
response of A. koa suckers (Spatz and Mueller-Dombois 1973, p. 873).
Another study at Puuwaawaa on Hawaii Island demonstrated that prior to
management actions in 1985, regeneration of endemic shrubs and trees in
a goat-grazed area was almost totally lacking, contributing to the
invasion of forest understory by exotic grasses and weeds. After the
removal of goats, A. koa and native Metrosideros seedlings were
observed germinating by the thousands (HDLNR 2002, p. 52). Based on
these studies, and other comparisons of fenced and unfenced areas, it
is clear that goats devastate native Hawaiian ecosystems (Loope et al.
1988, p. 277). Because feral goats occur in 10 of the 11 ecosystems
(all except anchialine pool) discussed in this proposal, the results of
the studies described above indicate that goats likely also alter these
ecosystems and directly damage or destroy native plants. Browsing or
grazing by feral goats poses a particular threat to the following plant
species: Exocarpos menziesii on Hawaii Island (NTBG Herbarium Database
2014, in litt.), Festuca hawaiiensis on Hawaii Island (Wood 2001b, in
litt.), Gardenia remyi
[[Page 67837]]
on Kauai (PEPP 2011, p. 114; PEPP 2013, p. 107; Kishida 2011, in
litt.), Huperzia stemmermanniae on Hawaii Island (HBMP 2010),
Joinvillea ascendens ssp. ascendens on Kauai (PEPP 2010, p. 80), Kadua
fluviatilis on Kauai (HBMP 2010), Labordia lorenciana on Kauai (PEPP
2011, p. 124; PEPP 2013, p. 126), Myrsine fosbergii on Kauai (HBMP
2010), Nothocestrum latifolium on Maui (HBMP 2010), Ochrosia haleakalae
on Maui and Hawaii Island (HBMP 2010), Phyllostegia stachyoides on
Molokai (HBMP 2010), Portulaca villosa on Hawaii Island (PEPP 2012, p.
140), Ranunculus mauiensis on Kauai and Maui (PEPP 2011, p. 161; PEPP
2012, p. 144; PEPP 2013, pp. 177-178; PEPP 2014, pp. 155-156; Kishida
2011, in litt.), Sanicula sandwicensis on Maui (PEPP 2011, p. 163), and
Sicyos lanceoloideus on Kauai (PEPP 2012, p. 154; PEPP 2013, p. 189).
In addition, browsing by feral goats may also damage or destroy native
host plants of the yellow-faced bees Hylaeus anthracinus, H.
assimulans, H. facilis, and H. hilaris.
Axis Deer
Axis deer are known to consume a wide range of forage items
throughout their native range and in areas where they have been
introduced (Anderson 1999, p. 3). Although they prefer to graze on
grass, axis deer have been documented to eat over 75 species of plants,
including all plant parts (Anderson 1999, p. 3). They exhibit a high
degree of opportunism regarding their choice of forage, and consume
progressively less palatable plants until no edible vegetation remains
(Dinerstein 1987, in Anderson 1999, p. 5; Medeiros 2010, pers. comm.).
Axis deer on Maui follow a cycle of grazing and browsing in open
lowland grasslands during the rainy season (November through March) and
then migrating to the lava flows of montane mesic forest during the dry
summer months to graze and browse on many native plant species, for
example, Abutilon menziesii (kooloaula, listed endangered), Erythrina
sandwicensis (wiliwili), and Sida fallax (Medeiros 2010, pers. comm.).
During the El Ni[ntilde]o drought cycles from 1988 through 2001, Maui
experienced an 80 to 90 percent decline in native shrub species caused
by axis deer browsing on and girdling young saplings (Medeiros 2010,
pers. comm.). On Lanai, grazing by axis deer has been reported as a
major threat to the endangered Gardenia brighamii (nanu), and Swedberg
and Walker (1978, in Anderson 2003, pp. 124-25) reported that the
native plants Osteomeles anthyllidifolia (uulei) and Leptecophylla
tameiameiae (pukiawe) comprised more than 30 percent of axis deer rumen
volume. During the driest summer months, axis deer are observed in
coastal areas in search of food (Medeiros 2010, pers. comm.). Because
axis deer occur in 10 of the 11 ecosystems on Molokai, Lanai, and Maui
(all except anchialine pool), the results from the studies above, in
addition to direct observations from field biologists, suggest that
axis deer can also alter these ecosystems and directly damage or
destroy native plants. Browsing or grazing by axis deer poses a threat
to the following plant species: Gardenia remyi on Molokai (HBMP 2010),
Huperzia stemmermanniae on Maui (HBMP 2010), Joinvillea ascendens ssp.
ascendens on Maui (PEPP 2014, pp. 108-109), Nothocestrum latifolium on
Lanai (PEPP 2012, p. 129), Portulaca villosa on Lanai (HBMP 2010),
Pseudognaphalium sandwicensium var. molokaiense on Molokai (Wood 2005c,
in litt.; Kallstrom 2008, in litt.; MNTF 2010), Ranunculus mauiensis on
Maui (PEPP 2013, p. 178; PEPP 2014, pp. 154-155), Schiedea pubescens on
Molokai and Lanai (Wood 2004, in litt.; Rowland 2006, in litt.;
Oppenheimer 2001, in litt.), and Solanum nelsonii on Molokai (PEPP
2012, p. 156; PEPP 2013, pp. 190-191; PEPP 2014, p. 167). Axis deer may
also damage or destroy native host plants of the yellow-faced bees
Hylaeus anthracinus, H. assimulans, H. facilis, H. hilaris, and H.
longiceps.
Black-Tailed Deer
Black-tailed deer are extremely adaptable, and in their native
range (U.S. Pacific coast) inhabit every principal ecosystem including
open grasslands, agricultural land, shrubland, woodland, mountain
forests, semi-deserts, and high mountain ecosystems (NRCS 2005, in
litt.). Their home range size varies in the continental United States,
but has been estimated to from 1 to 4 sq mi (2.5 to 10 km) and
sometimes as large as 30 sq mi (78 sq km), with adults defending small
areas when caring for fawns (NRCS 2005, in litt.). We do not know their
home range size on Kauai; however, the island is only 562 sq mi (1,456
sq km) in size. Black-tailed deer are primarily browsers, but as they
have a smaller rumen compared to other browsers in relation to their
body size, they must select the most nutritious plants and parts of
plants (Mule Deer Foundation 2011, in litt.). Their diet consist of a
diversity of living, wilted, dry, or decaying vegetation, including
leaves, needles, succulent stems, fruits, nuts, shrubs, herbaceous
undergrowth, domestic crops, and grasses (NRCS 2005, in litt.). Black-
tailed deer consume native vegetation on the island of Kauai (van Riper
and van Riper 1982, pp. 42-43; Stone 1985, pp. 262-263; Tomich 1986,
pp. 132-134; Cuddihy and Stone 1990, p. 67). In the 1990s, it was
estimated there were about 350 animals in and near Waimea Canyon;
however, in 2013, the population was estimated to be 1,000 to 1,200
animals in public hunting areas (not including private lands), and was
expanding into the southern and eastern sections of the island (Mule
Deer Working Group 2013, in litt.). According to State records, black-
tailed deer are feeding largely on the introduced species Psidium
cattleianum and Rubus rosifolius, as well as the native species Alyxia
stellata (maile), Dodonaea viscosa (aalii), Dianella sandwicensis
(ukiuki), Coprosma sp. (pilo), and Acacia koa (Cuddihy and Stone 1990,
p. 67). Browsing by black-tailed deer is a threat to the Kauai plant
species Asplenium diellaciniatum, Joinvillea ascendens ssp. ascendens,
Labordia lorenciana, Nothocestrum latifolium, Ranunculus mauiensis, and
Sicyos lanceoloideus.
Mouflon and Sheep
Mouflon, feral domestic sheep, and mouflon-sheep hybrids browse
native vegetation on Lanai and Hawaii Island. Domestic sheep have been
raised on Kauai, Lanai, Kahoolawe, and Hawaii, but today sheep farming
only occurs on Hawaii Island on Mauna Kea and Hualalai (Pratt and
Jacobi in Pratt et al. 2009, p. 151). Sheep browse (eating shoots,
leaves, flowers, and bark) on the native Sophora chrysophylla (mamane),
the primary food source of the endangered forest bird, the palila
(Loxioides bailleui) (Scowcroft and Sakai 1983, p. 495). Feral sheep
reductions were initiated in palila habitat; however, even after most
were removed, tree bark stripping continued and some mamane populations
did not recover (Pratt and Jacobi in Pratt et al. 2009, p. 151). On
Hawaii Island, vegetation browsing by mouflon led to the decline of the
largest population of the endangered Argyroxiphium kauense (kau
silversword, Mauna Loa silversword, or ahinahina), reducing it from a
``magnificent population of several thousand'' (Degener et al. 1976,
pp. 173-174) to fewer than 2,000 individuals in a period of 10 years
(unpublished data in Powell 1992, p. 312). Mamane is also preferred
browse for mouflon, and according to Scowcroft and Sakai (1983, p.
495), mouflon eat the shoots, leaves, flowers, and bark of this
species. Mouflon are also reported to strip bark from native koa trees
and to seek out the native plants Geranium
[[Page 67838]]
cuneatum (hinahina) and Silene hawaiiensis, and Lanai occurrences of
Gardenia brighamii (Benitez et al. 2008, p. 57; Mehrhoff 1993, p. 11).
While mouflon were introduced to Lanai and Hawaii Island as game
mammals, a private game ranch on Maui has added mouflon to its stock,
and it is likely that over time some individuals may escape (Hess 2010,
pers. comm.; Kessler 2010, pers. comm.). Browsing and grazing by
mouflon, feral domestic sheep, and mouflon-sheep hybrids poses a threat
to the plant species Exocarpos menziesii on Hawaii Island (Keitt and
Island Conservation 2008, pp. 90, 92; NPS 2013, pp. i, 124); Festuca
hawaiiensis on Hawaii Island (Oppenheimer 2001, in litt.; HBMP 2007, in
litt.); Nothocestrum latifolium on Lanai (PEPP 2012, p. 129); Portulaca
villosa on Lanai (HBMP 2010); Ranunculus hawaiensis on Hawaii Island
(HBMP 2010); and Sicyos macrophyllus on Hawaii Island (HBMP 2010).
Because feral sheep and mouflon occur in all of the described
ecosystems except for the anchialine pool ecosystem, the data from
studies above suggest that in addition to consuming the host plants of
the yellow-faced bees Hylaeus anthracinus and H. assimulans on Lanai,
herbivory by feral sheep and mouflon also may consume host plants of
the other species on Lanai: H. facilis, H. hilaris, and H. longiceps.
Feral Cattle
Grazing by cattle is considered one of the most important factors
in the destruction of Hawaiian forests (Baldwin and Fagerlund 1943, pp.
118-122). Feral cattle are currently found only on the islands of
Molokai, Maui, and Hawaii (Tomich 1986, pp. 140-144; de Sa et al. 2013,
29 pp.). Cattle consume tree seedlings and browse saplings (Cuddihy
1984, p. 16). In Hawaii Volcanoes National Park (Hawaii Island),
Cuddihy reported that there were twice as many native plant species as
nonnatives in areas that had been fenced to exclude cattle (Cuddihy
1984, pp. 16, 34). Loss of the native sandalwood forest on Lanai is
attributed to cattle (Skottsberg 1953 in Cuddihy 1984, p. 16). Browsing
and grazing by feral cattle poses a threat to the following plant
species: Huperzia stemmermanniae on Maui and Hawaii Island (Medeiros et
al. 1996b, p. 96); Nothocestrum latifolium on Molokai and Maui (HBMP
2010); Ochrosia haleakalae on Maui (HBMP 2010); Ranunculus hawaiensis
on Hawaii Island (HBMP 2010); R. mauiensis on Maui and Hawaii Island
(PEPP 2012, p. 144; PEPP 2013, p. 178; PEPP 2014, pp. 154-155; HBMP
2010); Schiedea pubescens on Maui (Wood 2005d, in litt.; HBMP 2010);
Sicyos macrophyllus on Hawaii Island (PEPP 2010, p. 111; HBMP 2010);
and Solanum nelsonii on Molokai (Wood 1999, in litt.; HBMP 2010).
Because feral cattle occur in 6 of the 11 ecosystems (lowland dry,
lowland mesic, lowland wet, montane wet, montane mesic, and subalpine)
in which these species occur on Molokai, Maui, and Hawaii Island, the
results from the studies cited above, in addition to direct
observations from field biologists, indicate that grazing by feral
cattle can directly damage or destroy these plants.
Blackbuck
The blackbuck antelope (Antelope cervicapra) is an endangered
species from India brought to a private game reserve on Molokai about
15 years ago from an Indian zoo (Kessler 2010, pers. comm.). According
to Kessler (2010, pers. comm.), a few individuals escaped captivity and
established a wild population of unknown size on the low, dry plains of
western Molokai. Blackbuck primarily use grassland habitat for grazing.
In India, foraging consumption and nutrient digestibility are high in
the moist winter months and low in the dry summer months (Jhala 1997,
pp. 1348, 1351). Although most plant species are grazed intensely when
they are green, some are grazed only after they are dry (Jhala 1997,
pp. 1348, 1351). Because the foraging dynamics of blackbuck antelope in
Hawaii and possible habitat effects are unknown at this time, we do not
currently consider this ungulate a threat to the four native plant
species known from dry areas on Molokai: Gardenia remyi, Nothocestrum
latifolium, Portulaca villosa, and Pseudognaphalium sandwicensium var.
molokaiense, or to the yellow-faced bees Hylaeus anthracinus, H.
facilis, H. hilaris, and H. longiceps (which rely on host plants that
ungulates consume).
Other Introduced Vertebrates
Rats
Three species of introduced rats occur in the Hawaiian Islands.
Studies of Polynesian rat (Rattus exulans) DNA suggest they first
appeared in the islands along with emigrants from the Marquesas Islands
(French Polynesia) in about 400 A.D., with a second introduction around
1100 A.D. (Ziegler 2002, p. 315). The black rat (R. rattus) and the
Norway rat (R. norvegicus) arrived in the islands more recently, as
stowaways on ships sometime in the late 19th century (Atkinson and
Atkinson 2000, p. 25). The Polynesian rat and the black rat are
primarily found in rural and remote areas of Hawaii, in dry to wet
habitats, while the Norway rat is typically found in urban areas or
agricultural fields (Tomich 1986, p. 41). The black rat is widely
distributed throughout the main Hawaiian Islands and can be found in a
range of ecosystems and as high as 9,000 ft (2,700 m), but it is most
common at low- to mid-elevations (Tomich 1986, pp. 38-40). Sugihara
(1997, p. 194) found both the black and Polynesian rats up to 7,000 ft
(2,000 m) on Maui, but found the Norway rat only at lower elevations.
Rats are omnivorous and eat almost any type of food (Nelson 2012, in
litt.). Rats occur in 7 of the 11 ecosystems (coastal, lowland mesic,
lowland wet, montane wet, montane mesic, montane dry, and wet cliff),
and predation or herbivory by rats is a threat to 19 plants
(Calamagrostis expansa (Maui and Hawaii Island; HBMP 2010), Cyanea
kauaulaensis (Maui; PEPP 2012, pp. 71-72; PEPP 2014, p. 73), Dryopteris
glabra var. pusilla (Kauai; Wood 2015, in litt.); Gardenia remyi
(Kauai, Molokai, Maui, and Hawaii Island; Wood 2004, in litt.; HBMP
2010); Joinvillea ascendens ssp. ascendens (Kauai, Oahu, Molokai, Maui,
and Hawaii Island; PEPP 2014, p. 109), Kadua haupuensis (Kauai; Lorence
et al. 2010, p. 140), Labordia lorenciana (Kauai; Wood et al. 2007, p.
198), Ochrosia haleakalae (Maui; Oppenheimer 2015, in litt.),
Phyllostegia helleri (Kauai; HBMP 2010), P. stachyoides (Molokai, Maui,
and Hawaii Island; PEPP 2012, p. 133; PEPP 2013, pp. 158-159; PEPP
2014, pp. 140-142), Pritchardia bakeri (Oahu; Hodel 2012, pp. 42, 73),
Ranunculus mauiensis (Kauai, Oahu, Molokai, Maui, and Hawaii Island;
HBMP 2010), Sanicula sandwicensis (Maui and Hawaii Island; PEPP 2012,
p. 148), Santalum involutum (Kauai; Harbaugh et al. 2010, pp. 835-836),
Schiedea diffusa ssp. diffusa (Molokai, Maui; HBMP 2010), S. pubescens
(Molokai, Lanai, and Maui; Wood 2005d, in litt.; HBMP 2010), Sicyos
macrophyllus (Maui and Hawaii Island; Pratt 2008, in litt.), Solanum
nelsonii (NWHI, Niihau, Molokai, Maui, and Hawaii Island; PEPP 2012, p.
156; PEPP 2014, p. 167), and Wikstroemia skottsbergiana (Kauai;
Mitchell et al. 2005, in litt.)) and to the band-rumped storm-petrel
(Lehua, Kauai, Maui, Kahoolawe, Lanai, and Hawaii Island; Pyle and Pyle
2009, in litt.).
Rat Impacts on Plants: Rats affect native plants by eating fleshy
fruits, seeds, flowers, stems, leaves, roots, and other plant parts
(Atkinson and Atkinson 2000, p. 23), and by stripping bark and cutting
small branches (twig cutting) in search of moisture and nutrients, with
detrimental impacts to
[[Page 67839]]
plants' vigor and regeneration (Abe and Umeno 2011, pp. 27-39; Nelson
2012, pp. 1-4, 8-9). Studies in New Zealand have demonstrated that
differential regeneration as a consequence of rat predation alters
species composition of forested areas (Cuddihy and Stone 1990, pp. 68-
69). Rats have caused declines or even the total elimination of island
plant species (Campbell and Atkinson 1999 in Atkinson and Atkinson
2000, p. 24). In the Hawaiian Islands, rats may consume as much at 90
percent of the seeds produced by some native plants, and in some cases
prevent regeneration of forest species completely (Cuddihy and Stone
1990, pp. 68-69). Hawaiian plants with fleshy fruit, such as Cyanea and
Pritchardia, are particularly susceptible to rat predation (Cuddihy and
Stone 1990, pp. 67-69). Predation of seeds by rats poses a serious and
ongoing threat to all the Hawaiian Pritchardia palms, including P.
bakeri, because rats are able to consume every seed in a fruiting
stalk, preventing successful reproduction (Hodel 2012, pp. 42, 73).
Fossil pollen records indicate that Pritchardia palms were once among
the dominant species of coastal, lowland, and interior forests of
Hawaii (Burney et al. 2001, pp. 630-631; Chapin et al. 2007, p. 21);
today, complete coverage by all age classes of Pritchardia occurs only
on small islets currently unoccupied by rats (Athens 2009, p. 1498).
Because rats occur in seven of the ecosystems in which these species
occur, the results from these studies, in addition to direct
observations by field biologists, suggest that predation by rats can
directly damage or destroy native plants.
Rat Impacts on the Band-Rumped Storm-Petrel: Introduced predators
are the most serious threat facing the band-rumped storm-petrel. Rats
occur on all the main Hawaiian Islands, and populations are also high
on Lehua; however, attempts to control rats on Lehua are ongoing
(Parkes and Fisher 2011, 48 pp.). Ground-, crevice-, and burrow-nesting
seabirds, as well as their eggs and young, are highly susceptible to
predation by rats; storm-petrels are the most susceptible of seabirds
to rat predation and have experienced population-level impacts and
extirpation as a result (Simons 1984, p. 1073; Jones et al. 2008, pp.
20-21). Evidence from the islands of Hawaii and Maui show that the
Hawaiian petrel, a much larger seabird that nests in some of the same
areas as the band-rumped storm-petrel, suffers huge losses to
introduced predators (Johnston 1992, in litt.; Hodges and Nagata 2001,
pp. 308-310; Hu et al. 2001, p. 234). The effects of introduced
predators on the breeding success of the band-rumped storm-petrel are
probably similar to the documented effects on the breeding success of
Hawaiian petrels because these birds are similarly vulnerable.
Population modeling showed that consistent predation of Hawaiian
petrels, where reproductive success was reduced to 35 percent and adult
survival was 80 percent, could drive a population to extinction in 20
to 30 years (Simons 1984, pp. 1071-1073). Rat bones were collected from
a band-rumped storm-petrel nest on a sheer cliff on Kauai, and two live
rats were observed moving along small rock ledges in the same area
(Wood et al. 2002, p. 8), demonstrating that even remote and otherwise
inaccessible nest sites are not safe from these predators. Because rats
are present in all three ecosystems in which the band-rumped storm-
petrel occurs (coastal, dry cliff, and wet cliff), predation by rats
likely results in decreases in the numbers and populations of the band-
rumped storm-petrel. We do not anticipate a reduction of this threat in
the near future.
Barn Owl Impacts on the Band-Rumped Storm-Petrel
Two species of owls, the native pueo (Asio flammeus sandwichensis)
and the introduced barn owl (Tyto alba), are known to prey on native
Hawaiian birds. For example, between 1996 and 1998, 10 percent of nest
failures of the puaiohi (small Kauai thrush, Myadestes palmeri), an
endangered forest bird, on Kauai were attributed to owls (Snetsinger et
al. 1994, p. 47; Snetsinger et al. 2005, pp. 72, 79). The band-rumped
storm petrel only comes to land after dark, and likely avoids predation
by the pueo, which hunts in daylight (Hawaii DOFAW 2005). The nonnative
barn owl, however, is a nocturnal hunter and may prey on the storm-
petrel. Barn owls were introduced to Kauai, Oahu, Molokai, and Hawaii
Island between 1958 and 1963, to control rats and mice in sugar cane
plantations, and now they occur on all of the main islands (USFWS 2013,
p. 9). Barn owls are well-known predators of storm-petrels and other
seabirds on islands (LeCorre and Jouventin 1997, p. 215; Velarde et al.
2007, p. 286; Guerra et al. 2014, p. 182; Ringler et al. 2015, p. 79).
Direct impacts of barn owls on band-rumped storm-petrels in Hawaii are
not well documented, but evidence and numerous anecdotal reports exist
of barn owls preying on seabirds in the main Hawaiian islands,
including the threatened Newell's shearwater and endangered Hawaiian
petrel, and including on Kauai and Lehua, where band-rumped storm-
petrels are known to nest (summarized in USFWS 2013, pp. 11-12).
Because barn owls occur throughout the range of the band-rumped storm-
petrel in Hawaii, they are likely to be predators of these seabirds.
Cat Impacts on the Band-Rumped Storm-Petrel
Cats (Felis catus) were introduced to Hawaii in the early 1800s,
and are present on all the main Hawaiian Islands (Tomich 1986, p. 101).
Cats are notorious for their predation on birds (Tomich 1986, p. 102).
Native mammalian carnivores are absent from oceanic islands because of
their low dispersal ability, but once introduced, are significant
predators on seabird colonies and terrestrial birds that have no innate
defenses against predation by these animals (Scott et al. 1986, p. 363;
Ainley et al. 1997, p. 24; Ziegler 2002, p. 243; Hess and Banko 2006,
in litt.; Nogales et al. 2013, p. 804). Cats may have contributed to
the extinction of the Hawaiian rail (Porzana sandwichensis) (Stone 1985
in Stone and Scott 1985, p. 266). Although cats are more common at
lower elevations, there are populations in areas completely isolated
from human presence, including montane forests and alpine areas of Maui
and Hawaii Island (Lindsey et al. in Pratt et al. 2009, p. 277; Scott
et al. 1986, p. 363). Examination of the stomach contents of feral cats
at Hakalau Forest NWR (Hawaii Island) found native and introduced birds
to be the most common prey item (Banko et al. 2004, p. 162). Cats are
believed to prey on roosting or incubating adult band-rumped storm-
petrels and young, as evidenced by carcasses found in Hawaii Volcanoes
National Park depredated by cats (Hu, pers. comm. in Slotterback 2002,
in litt.; Hess et al. 2008, pp. 11, 14). Predation by cats is well
known for the endangered Hawaiian petrel, which has some accessible and
well-studied nesting areas; this species shares life-history and
evolutionary traits with the band-rumped storm-petrel that make both
vulnerable to nonnative mammalian predators. We expect the band-rumped
storm-petrel to experience impacts of cat predation similar to those
documented in the Hawaiian petrel. On Mauna Loa (Hawaii Island), feral
cats were major predators of Hawaiian petrels (Hu et al. 2001, p. 234),
and on Haleakala (Maui), almost half of the known mortalities of
Hawaiian petrels between 1964 and 1996 were attributed to cats
(Natividad Hodges and Nagata 2001, p. 312; Hu et al. 2001, p. 234).
Population modeling of the Hawaiian petrel indicated that the petrel
population would be unable to
[[Page 67840]]
withstand any level of predation for long, and even with seemingly low
levels of predation, the petrel population would be reduced by half in
fewer than 30 years (Simons 1984, p. 1073). The band-rumped storm
petrel is smaller in size than the petrel, but also nests in burrows
and rock-crevices, lacks co-evolved predator avoidance behavior, and
has a lengthy incubation and fledgling period, making this species
highly vulnerable to predation by introduced mammals. Because feral
cats occur in all four ecosystems in which the band-rumped storm petrel
occurs, they are likely to be significant predators of these birds.
Mongoose Impacts on the Band-Rumped Storm-Petrel
The small Indian mongoose (Herpestes auropunctatus) was introduced
to Hawaii in 1883, to control rodents in sugar cane plantations (Tomich
1986, pp. 95-96). This species quickly became widespread on Oahu,
Molokai, Maui, and Hawaii Island, from sea level to elevations as high
as 7,000 ft (2,130 m) (Tomich 1986, pp. 93-94). Mongooses have been
sighted, and two captured, on Kauai, but it is still uncertain if the
species is established there or how large populations might be (Kauai
Invasive Species Committee 2013, in litt.; The Garden Island 2012, in
litt.; Hess et al. in Pratt et al. 2009, p. 429). Mongooses are
omnivorous, are known to prey on Hawaiian birds and their eggs, and are
considered a likely factor in the decline of the endangered Hawaiian
goose (nene, Branta sandvicensis) (Tomich 1986, p. 97). They are known
or suspected predators on other Hawaiian birds, including the Hawaiian
crow (alala, Corvus hawaiiensis), Hawaiian duck (koloa, Anas
wyvilliana), Hawaiian coot (alae keokeo, Fulica alai), Hawaiian stilt
(aeo, Himantopus mexicanus knudseni), Hawaiian gallinule (ula,
Gallinula chloropus sandvicensis), Hawaiian petrel, and Newell's
shearwater. Bird extinctions in other areas are attributed to
mongooses, such as the loss of the barred-wing rail (Nesoclopeus
poecilopterus) in Fiji, and the Jamaica petrel (Pterodroma caribbaea)
(Hays and Conant 2007, p. 6). Birds extirpated from islands occupied by
mongooses retain their populations on islands known to be mongoose-free
(Hays and Conant 2007, p. 7). In Hawaii, mongooses occur in habitat
types where they are not found within their natural range, and they
have no predators and few communicable diseases or parasites. Because
mongooses occur in all four ecosystems in which the band-rumped storm
petrel occurs, they are likely to be significant predators of the band-
rumped storm-petrel.
Nonnative Fish Impacts on the Orangeblack Hawaiian Damselfly
Predation by nonnative fishes is a threat to the orangeblack
Hawaiian damselfly. Similar to the aquatic insects, Hawaii has a low
diversity of freshwater fishes, with only five native species in two
families (gobies (Gobiidae) and sleepers (Eleotridae)) that occur on
all the main islands (Devick 1991, p. 196). Information on these five
species indicates that the Hawaiian damselflies probably experienced
limited natural predation pressure from these native fishes (Kido 1997,
p. 493; Englund 1999, p. 236). Conversely, fish predation has been an
important factor in the evolution of behavior in damselfly naiads in
continental systems (Johnson 1991, p. 13). Some species of damselflies,
including the native Hawaiian species, are not adapted to coexist with
some fish species, and are found only in bodies of water without fish
(Henrikson 1988, pp. 179-180; McPeek 1990a, pp. 92-93). The naiads of
these species tend to occupy more exposed positions and engage in
conspicuous foraging behavior that makes them susceptible to predation
by fishes (Macan 1977, p. 47; McPeek 1990b, p. 1722). The introduction
of nonnative fishes has been implicated in the extirpation of a species
related to the orangeblack Hawaiian damselfly, the endangered Pacific
Hawaiian damselfly (Megalagrion pacificum), from Oahu, Kauai, and
Lanai, and from many streams on the remaining islands where it occurs
(Moore and Gagne 1982, pp. 1-4). More than 70 species of fish have been
introduced into Hawaiian freshwater habitats (Devick 1991, p. 189;
Englund and Eldredge in Staples and Cowie 2001, p. 32; Englund 2004, in
litt., p.27). The impact of fish introductions prior to 1900 cannot be
assessed because this predates the initial collection of damselflies in
Hawaii (Perkins 1913, p. clxxvi). In 1905, two species, the mosquito
fish (Gambusia affinis) and the sailfin molly (Poecilia latipinna),
were introduced for biological control of mosquitoes (Van Dine 1907,
pp. 6-9). In 1922, three additional species were established for
mosquito control, the green swordtail (Xiphophorus helleri), the
moonfish (Xiphophorus maculatus), and the guppy (Poecilia reticulata).
By 1935, the orangeblack Hawaiian damselfly was found only in waters
without introduced fishes (Williams 1936, p. 289; Zimmerman 1948b, p.
341; Polhemus 1993b, p. 591; Englund 1998, p. 235). Beginning about
1980, a large number of new fish introductions began in Hawaii,
originating primarily from the aquarium fish trade (Devick 1991, p.
189). This recent wave of fish introductions on Oahu corresponded with
the drastic decline and range reduction of other Hawaiian damselfly
species: The endangered oceanic Hawaiian damselfly (Megalagrion
oceanicum), the endangered crimson Hawaiian damselfly (M. leptodemas),
and the endangered blackline Hawaiian damselfly (M. nigrohamatum
nigrolineatum). Currently, these damselflies are found only in
drainages or higher parts of stream systems where nonnative fish are
not yet established (Englund and Polhemus 1994, pp. 8-9; Englund 2004,
in litt., p. 27). In summary, Hawaiian damselflies evolved with few, if
any, predatory fishes and the lack of defensive behavior makes the
orangeblack Hawaiian damselfly particularly vulnerable to, and are
threatened by, predation by nonnative fish.
Nonnative Fish Impacts on the Anchialine Pool Shrimp
In Hawaii, the introduction of nonnative fishes into anchialine
pools and the ensuing predation by nonnative fishes is considered the
greatest threat to native shrimp within anchialine pool systems
(Bailey-Brock and Brock 1993, p. 354). These impacts are discussed
further under Factor E. Other Natural or Manmade Factors Affecting
Their Continued Existence, below.
Bullfrog Impacts on the Orangeblack Hawaiian Damselfly
Native to the eastern United States and the Great Plains region,
the bullfrog (Rana catesbeiana, Lithobates catesbeiana), was first
introduced to Hawaii in 1899, to help control insects, and has become
established on all the main Hawaiian Islands (Bryan 1931, pp. 62-63;
Bury and Whelan 1985, p. 1; Lever 2003, p. 203). The bullfrog is
flexible in both habitat and food requirements (McKeown 1996, pp. 24-
27; Bury and Whelan 1984, pp. 3-7; Lever 2003, pp. 203-204), and can
utilize any water source within a temperature range of 60 to 75 degrees
Fahrenheit ([deg]F) (16 to 24 degrees Celsius ([deg]C)) (DesertUSA
2008). Englund et al. (2007, pp. 215, 219) found a strong correlation
between the presence of bullfrogs and the absence of Hawaiian
damselflies in their study of streams on all the main Hawaiian Islands.
Because bullfrogs are omnivorous feeders and occur in the same habitat
as the orangeblack Hawaiian damselfly, we consider predation by
bullfrogs a threat to the orangeblack Hawaiian damselfly.
[[Page 67841]]
Introduced Invertebrates
Slugs
Herbivory by nonnative slugs is a threat to 10 of the 39 plant
species (Cyanea kauaulaensis (Maui); Deparia kaalaana (Maui),
Dryopteris glabra var. pusilla (Kauai), Hypolepis hawaiiensis var.
mauiensis (Maui), Ochrosia haleakalae (Maui, Hawaii Island),
Phyllostegia brevidens (Maui), P. stachyoides (Molokai, Maui),
Ranunculus mauiensis (Kauai, Maui), Schiedea diffusa ssp. diffusa
(Maui), and S. pubescens (Molokai, Maui)) through mechanical damage,
destruction of plant parts, and mortality (see Table 2) (Joe 2006, p.
10; HBMP 2010; PEPP 2011, pp. 149, 170; PEPP 2012, pp. 71-72, 117-118,
133, 144-145, 153; PEPP 2013, pp. 54, 67, 91, 125-126, 158-159, 177-
178, 185; Oppenheimer and Bustamente 2014, p. 106; PEPP 2014, pp. 73,
112-114, 136, 141-142, 154-156, 159, 162-163). Slugs are known to
damage individuals of Cyanea and Cyrtandra species in the wild (Wood
2001, in litt.; Sailer and Kier 2002, in litt.; PEPP 2007, p. 38; PEPP
2008, pp. 23, 29, 52-53, 57). Information in the U.S. Army's 2005
``Status Report for the Makua Implementation Plan'' indicates that
herbivory by slugs can be a threat to all species of Cyanea, and can
result in up to 80 percent seedling mortality (U.S. Army Garrison 2005,
p. 3-51). Slug damage has also been reported on other Hawaiian plants
including Argyroxiphium grayanum (greensword), Alsinidendron sp.,
Hibiscus sp., Schiedea kaalae (maolioli), Solanum sandwicense (popolo
aiakeakua), and Urera sp. (Gagne 1983, pp. 190-191; Sailer 2006, pers.
comm. in Joe 2006, pp. 28-34). Joe and Daehler (2008, p. 252) found
that native Hawaiian plants are more vulnerable to slug damage than
nonnative plants. In particular, they found that individuals of the
endangered plants Cyanea superba and Schiedea obovata had 50 percent
higher mortality when exposed to slugs as compared to individuals that
were within exclosures without slugs. Because slugs are reported in
five ecosystems (lowland mesic, lowland wet, montane wet, montane
mesic, and wet cliff) on all the main Hawaiian Islands, the data from
the studies cited above, in addition to direct observations by field
biologists, indicate that slugs can directly damage or destroy native
plants.
Black Twig Borers
The black twig borer (Xylosandrus compactus) is known to infest a
wide variety of common plant taxa, including rare native plant species
(Davis 1970, p. 39; Extension Entomology and US-CTAHR Integrated Pest
Management Program 2006, p. 1). This insect pest burrows into branches,
introduces a pathogenic fungus as food for its larvae, and lays its
eggs (Davis 1970, p. 39). Twigs, branches, and entire plants can be
damaged or killed from an infestation (Extension Entomology and UH-
CTAHR Integrated Pest Management Program 2006, in litt.). On the
Hawaiian Islands, the black twig borer has many hosts, disperses
easily, and is probably present at most elevations up to 2,500 ft (762
m) (Howarth 1985, pp. 152-153). The black twig borer is reported as a
threat to Labordia lorenciana and Nothocestrum latifolium (Ching-Harbin
2015, in litt.; Kishida 2015, in litt.).
Backswimmers
Backswimmers are aquatic true bugs (Heteroptera) in the family
Notonectidae, so called because they swim upside down. Backswimmers are
voracious predators and frequently feed on prey much larger than
themselves, such as tadpoles, small fish, and other aquatic
invertebrates including damselfly naiads (Borror et al. 1989, p. 296;
Zalom 1978, p. 617). Backswimmers (several species) were introduced in
recent times. Buenoa pallipes (NCN) has been recorded from Hawaii
Island, Oahu, Maui, and Kauai (Zimmerman 1948, pp. 232-233; Larsen
1996, p. 40). This species is found in streams and can be abundant in
lowland ponds and reservoirs. It feeds on any suitably sized insect,
including damselfly naiads (Zalom 1978, p. 617). Two additional species
of backswimmers have become established in Hawaii, Anisops kuroiwae
(NCN) on Maui and Lanai, and Notonecta indica (NCN) on Hawaii Island,
Oahu, and Maui (Larsen 1996, pp. 39-40). Predation by backswimmers is a
threat to the orangeblack Hawaiian damselfly (Haines 2015, in litt.).
Ants
At least 47 species of ants are known to be established in the
Hawaiian Islands (Hawaii Ants 2008, 11 pp.). No native ant species
occur in Hawaii, and the native yellow-faced bee species in Hawaii
evolved in the absence of predation pressure from ants. Ants are known
to prey upon Hawaiian yellow-faced bee (Hylaeus) species, with
observations of drastic reductions in yellow-faced bee populations in
ant-infested areas (Medeiros et al. 1986, pp. 45-46; Reimer 1994, p.
17; Stone and Loope 1987, p. 251; Cole et al. 1992, pp. 1313, 1317,
1320). The presence of ants in nearly all of the low-elevation habitat
sites currently and historically occupied by yellow-faced bee species
may preclude these species' recovery in some of these areas (Reimer
1994, pp. 17-18; Daly and Magnacca 2003, pp. 9-10). Although the
primary impact of ants on Hawaii's native invertebrate fauna is via
predation, they also compete for nectar (Reimer 1994, p. 17; Howarth
1985, p. 155; Hopper et al. 1996, p. 9; Holway et al. 2002, pp. 188,
209; Daly and Magnacca 2003, p. 9; Lach 2008, p. 155) and nest sites
(Krushelnycky et al. 2005, pp. 6-7). Some ant species may affect
yellow-faced bee species indirectly as well, by consuming seeds of
native host plants, thereby reducing the plants' recruitment and
fecundity (Bond and Slingsby 1984, p. 1031). The threat of ant
predation on the yellow-faced bees is amplified by the fact that most
ant species have winged reproductive adults and can quickly expand
their range by establishing new colonies in suitable habitat (Staples
and Cowie 2001, p. 55). In addition, these attributes allow some ants
to destroy otherwise geographically isolated populations of native
arthropods (Nafus 1993, pp. 19, 22-23). Several studies suggest a
serious ecosystem-level effect of invasive ants on plant pollination
(Krushelnycky et al. 2005, p. 9; Lach 2008, p. 155). Where ranges
overlap, ants compete with native pollinators such as yellow-faced bees
and preclude them from pollinating native plants (Howarth 1985, p.
157), potentially leading to a decrease in availability of the bees'
native plant food sources. Lach (2008, p. 155) found that yellow-faced
bees that regularly consume pollen from flowers of Metrosideros
polymorpha (ohia) were entirely absent from trees with flowers visited
by the ant Pheidole megacephala.
The four most aggressive ant species in Hawaii are the big-headed
ant (Pheidole megacephala), the yellow crazy ant (Anoplolepis
gracilipes), the tropical fire ant (Solenopsis geminata), and S.
papuana (NCN). The big-headed ant is native to central Africa and was
first reported in Hawaii in 1879 (Krushelnycky et al. 2005, p. 24).
This species occurs from coastal to mesic habitat up to 4,000 ft (1,220
m) in elevation. With few exceptions, native insects have been
eliminated in habitats where the big-headed ant is present (Perkins
1913, p. xxxix; Gagne 1979, p. 81; Gillespie and Reimer 1993, p. 22).
Native habitat of the yellow crazy ant is not known, but it is
speculated the species originated in West Africa (MacGown 2015, in
litt.). It occurs in low to mid elevations (less than 2,000
[[Page 67842]]
ft (600 m)) in rocky areas of moderate annual rainfall (less than 100
in (250 cm)) (Reimer et al. 1990, p. 42). Although surveys have not
been conducted to ascertain this species' presence in each of the known
habitats occupied by the seven yellow-faced bees, we know that the
yellow crazy ant occurs adjacent to some of the identified populations
sites based upon observations of their expanding range and their
preference for coastal and dry forest habitat (as indicated where the
species is most commonly collected) (Antweb 2015, in litt.; Magnacca
and King 2013, pp. 13-14). Direct observations indicate that Hawaiian
arthropods are susceptible to predation by this ant species. Gillespie
and Reimer (1993, pp. 21, 26) and Hardy (1979, pp. 37-38) documented
the complete elimination of native spiders from mesic and dry forests
after they were invaded by the big-headed ant and the yellow crazy ant.
Lester and Tavite (2004, p. 291) found that the yellow crazy ant in the
atolls of Tokelau (Central Polynesia) form very high densities in a
relatively short period of time with locally serious consequences for
invertebrate diversity. Densities of 3,600 individuals collected in
pitfall traps within a 24-hour period were observed, as well as
predation on invertebrates ranging from crabs to other ant species.
Results from these and other studies (Reimer et al. 1990, p. 47)
indicate that yellow crazy ants have the potential as predators to
profoundly affect endemic insect fauna in areas they occupy. We believe
that the yellow crazy ant is a threat to populations of the Hawaiian
yellow-faced bees in areas within their range.
Solenopsis papuana, native to the Pacific region but not to Hawaii,
is the only abundant, aggressive ant that has invaded intact mesic and
wet forest, as well as coastal and lowland dry ecosystems. First
detected in 1967, this species occurs from sea level to over 3,600 ft
(1,100 m) on all of the main Hawaiian Islands, and is still expanding
its range (Reimer et al. 1990, p. 42; Reimer 1993, p. 14). Studies have
been conducted that suggest a negative effect of this ant species on
indigenous invertebrates (Gillespie and Reimer 1993, p. 21). Although
surveys have not been conducted to ascertain the presence of S. papuana
in each of the known ecosystems occupied by the seven yellow-faced
bees, because of the expanding range of this introduced ant species,
and its widespread occurrence in coastal to wet habitats, it is a
possible threat to all known populations of the seven yellow-faced
bees.
Solenopsis geminata is also considered a significant threat to
native invertebrates in Hawaii (Wong and Wong 1988, p. 171). Found in
drier areas of all the main Hawaiian Islands, it displaced Pheidole
megacephala as the dominant ant in some localities more than 20 years
ago (Wong and Wong 1988, p. 175). Known to be a voracious predator,
this ant species was documented to significantly increase native fruit
fly mortality in field studies in Hawaii (Wong and Wong 1988, p. 175).
Solenopsis geminata is included among the eight species ranked as
having the highest potential risk to New Zealand species in a detailed
pest risk assessment for the country (GISD 2011, in litt.), and is
included as one of the five ant species listed among the ``100 of the
World's Worst Invaders'' (Manaaki Landcare Research 2015, in litt.). In
addition to predation, S. geminata workers tend honeydew-producing
members of the Homoptera suborder, especially mealybugs, which can
affect plants directly and indirectly through the spread of disease
(Manaaki Landcare Research 2015, in litt.). Although surveys have not
been conducted to ascertain the presence of S. geminata in each of the
known seven yellow-faced bees' habitat sites, because of its expanding
range and widespread presence, S. geminata is a threat to all known
populations of the seven yellow-faced bees.
Although we have no direct information that correlates the decrease
in populations of the seven yellow-faced bees in this final rule due to
the establishment of nonnative ants, predation of and competition with
other yellow-faced bee species by ants has been documented, resulting
in clear reductions in or absence of populations (Magnacca and King
2013, p. 24). We expect similar predation impacts to the seven yellow-
faced bees to continue as a result of the widespread presence of ants
throughout the Hawaiian Islands, their highly efficient and non-
specific predatory behavior, and their ability to quickly disperse and
establish new colonies. Therefore, we conclude that predation by
nonnative ants represents a serious threat to the continued existence
of the seven yellow-faced bees, now and into the future.
Wasps
Predation by the western yellow jacket wasp (Vespula pensylvanica)
is a serious and ongoing threat to the seven yellow-faced bees (Gambino
et al. 1987, p. 170; Wilson et al. 2009, pp. 1-5). The western yellow
jacket is a social wasp species native to mainland North America. It
was first reported on Oahu in the 1930s (Sherley 2000, p. 121), and an
aggressive race became established in 1977 (Gambino et al. 1987, p.
170). In temperate climates, the western yellow jacket wasp has an
annual life cycle, but in Hawaii's tropical climate, colonies of this
species persist year round, allowing growth of large populations
(Gambino et al. 1987, p. 170) and thus a greater impact on prey
populations. Most colonies occur between 2,000 and 3,500 ft (600 and
1050 m) in elevation (Gambino et al. 1990, p. 1088), although they can
also occur at sea level. The western yellow jacket wasp is known to be
an aggressive, generalist predator and has been documented preying upon
Hawaiian yellow-faced bee species (Gambino et al. 1987, p. 170; Wilson
et al. 2009, p. 2). It has been suggested that the western yellow
jacket wasp may compete for nectar with native Hawaiian invertebrates,
but we have no information to suggest this represents a threat to the
seven yellow-faced bees. Predation by the western yellow jacket wasp is
a significant threat to the seven yellow-faced bee species because of
the wasps' presence in habitat combined with the small number of
occurrences and small population sizes of the Hawaiian yellow-faced
bees.
Summary of Factor C
We are unaware of any information that indicates that disease is a
threat to the 39 plant species. We are also unaware of any information
that indicates that disease is a threat to the band-rumped storm-
petrel, the orangeblack Hawaiian damselfly, or the anchialine pool
shrimp, Procaris hawaiana. It has been suggested that transmission of
disease from alien invertebrates by mutual flower visitation is a
threat to the seven yellow-faced bees (Hylaeus spp.), but we currently
have no evidence that this is occurring.
We consider predation and herbivory by one or more of the nonnative
animal species (pigs, goats, axis deer, black-tailed deer, sheep,
mouflon, cattle, rats, barn owls, cats, mongooses, fish, slugs, ants,
black twig borers, and wasps) to pose an ongoing threat to 35 of the 39
plant species and to all 10 animal species throughout their ranges for
the following reasons:
(1) Observations and reports have documented that pigs, goats, axis
deer, black-tailed deer, sheep, mouflon, and cattle browse 27 of the 39
plant species, in addition to other studies demonstrating the negative
impacts of ungulate browsing on native plant species of the islands. If
the numbers and range of blackbuck antelope increase, their browsing
will be a threat to native plants that occur on Molokai,
[[Page 67843]]
including host plants for the yellow-faced bees.
(2) Nonnative rats and slugs (either singly or combined) cause
mechanical damage to plants and destruction of plant parts (branches,
flowers, fruits, and seeds), and are considered a threat to 22 of the
39 plant species.
(3) Rats also prey upon adults, juveniles, and eggs of the band-
rumped storm-petrel, and are linked with the dramatic decline of many
closely related bird species. Because rats are found in all of the
ecosystems in which the band-rumped storm-petrel occurs, we consider
predation by rats to be a serious and ongoing threat.
(4) Barn owls and cats have established populations in the wild on
all the main islands, and mongooses have established populations on all
the main islands except for Kauai. All of these nonnative animals are
known to prey on ground- and burrow-nesting seabirds. Predation by
these animals is a serious and ongoing threat to the band-rumped storm-
petrel.
(5) The absence of Hawaiian damselflies (including the orangeblack
Hawaiian damselfly) from ponds, pools, and other aquatic habitat on the
main Hawaiian Islands is strongly correlated with the presence of
predatory nonnative fish; numerous observations and reports suggest
nonnative predatory fishes eliminate native damselflies from these
habitats. Accordingly, predation by nonnative fishes is a serious and
ongoing threat to the orangeblack Hawaiian damselfly. Predation by
bullfrogs, backswimmers, and Jackson's chameleons, and competition with
caddisflies are threats to the orangeblack Hawaiian damselfly.
(6) Once introduced to anchialine pools, nonnative fish, through
predation and competition for food sources, directly affect anchialine
pool shrimp, including Procaris hawaiana, and also disrupt anchialine
pool ecology.
(7) Damage and destruction by the black twig borer is a threat to
two plant species, Labordia lorenciana and Nothocestrum latifolium.
(8) Predation by nonnative ants and wasps poses a threat to all
seven yellow-faced bees.
These threats are serious and ongoing, act in concert with other
threats to the species, and are expected to continue or increase in
magnitude and intensity into the future without effective management
actions to control or eradicate them. The effects of the combined
threats suggest the need for immediate implementation of recovery and
conservation methods.
Factor D. The Inadequacy of Existing Regulatory Mechanisms
Overview
Currently, no existing Federal, State, or local laws, treaties, or
regulations specifically conserve or protect 48 of the 49 species
(except the band-rumped storm-petrel by the Migratory Bird Treaty Act
(MBTA; 16 U.S.C. 703-712)), or adequately address the threats to any of
the 49 species (see Table 2). There are a few small programs and
organizations that conduct vegetation monitoring and nonnative species
and predator control, but these activities are nonregulatory, and
neither continuation of these conservation efforts nor funding for them
is guaranteed.
Federal laws pertaining to the 49 species addressed here include
Executive Order (E.O.) 13112, the MBTA, the Lacey Act (16 U.S.C. 3371-
3378; 18 U.S.C. 42-43), the Federal Noxious Weed List (7 CFR 360.200),
and the Convention on International Trade in Endangered Species of Wild
Fauna and Flora (CITES). The U.S. Department of Agriculture (USDA)
inspects propagative and restricted plant materials and animals, and
implements ``Special Local Needs'' rules for pesticide use, but only on
a species-by-species basis. The Department of Homeland Security-Customs
and Border Protection (CBP) is responsible for inspecting commercial,
private, and military vessels and aircraft, and related cargo and
passengers arriving from foreign locations. However, CBP focuses on
quarantine issues involving non-propagative plant materials; wooden
packing materials, timber, and products; internationally regulated
commercial species under CITES; and federally listed noxious plants,
seeds, soils, and pests of concern to the continental United States,
such as pests to mainland U.S. forests and agriculture.
Hawaii State law regarding natural resource protections include
those under Hawaii revised statutes (HRS): Plant and nondomestic animal
quarantine and microorganism import (HRS 11-3-150A) and noxious weed
control (HRS 11-3-152); flood control (HRS 12-2), water and land
development (HRS 12-174), and State water code (HRS 12-2-174D);
wildlife (general wildlife, hunting, game birds, game mammals, and wild
birds and other wildlife) (HRS 12-4-183D); aquatic resources and
wildlife-alien aquatic organisms (HRS 12-5-187A); general and
miscellaneous, invasive species council (HRS 12-6-194); conservation of
aquatic life, wildlife, and land plants (HRS 12-6-195D); and Natural
Area Reserves (NARs) (HRS 12-6-195). These laws are interpreted and
implemented under Hawaii administrative rules (HAR). Applicable HARs
include: Noxious weed rules (HAR 4-6-68); plant and nondomestic animal
quarantine, microorganism import rules (HAR 4-6-ch 71A, 71C), and plant
intrastate rules (HAR4-6-72); rules regulating game mammal hunting (HAR
13-5-2-ch 123; indigenous wildlife, endangered and threatened wildlife,
and introduced wild birds (HAR 13-5-2-ch 124); protection of instream
uses of water (HAR 13-7-ch 169), and NARs system (HAR 13-9-ch 208-210).
Private and local programs that provide protections, and that help
to implement Federal and State environmental regulations, laws, and
rules for one or more of the 49 species, include the Hawaii Invasive
Species Committee (HISC), the Coordinating Group on Alien Pest Species
(CGAPS), and the Hawaii Association of Watershed Partnerships (HAWP).
In addition, the Plant Extinction Protection Program (PEPP) was created
to protect Hawaii's rare plant species in need of immediate
conservation efforts, by monitoring, propagating, outplanting, and
providing some protection from threats.
We discuss Federal and State regulatory mechanisms, along with
agencies and groups authorized to implement them, and the coordination
between them, below.
Federal Regulatory Mechanisms
On February 3, 1999, Executive Order (E.O.) 13112 was signed
establishing the National Invasive Species Council (NISC). This E.O.
requires that a Council of Departments dealing with invasive species be
created to prevent the introduction of invasive species; provide for
their control; and minimize the economic, ecological, and human health
impacts that invasive species cause. Invasive species include aquatic
plant and animal species, terrestrial plants and animal species, and
plant and animal pathogens. This E.O. was reviewed in 2005 (NISC 2005).
NISC uses a cooperative approach to enhance the Federal Government's
response to the threat of invasive species, and emphasizes prevention,
early detection and rapid response, and sharing of information. See our
discussion below concerning the Hawaii Invasive Species Committee
(HISC) regarding the effectiveness of this law.
The MBTA is the domestic law that implements the United States'
commitment to four international conventions (with Canada, Japan,
Mexico, and Russia) for the protection
[[Page 67844]]
of shared migratory bird resources. The MBTA regulates most aspects of
take, possession, transport, sale, purchase, barter, export, and import
of migratory birds and prohibits the killing, capturing, and collecting
of individuals, eggs, and nests, unless such action is authorized by
permit. While the MBTA does prohibit actions that directly kill a
covered species, unlike the Endangered Species Act (Act), it does not
prohibit habitat modification that indirectly kills or injures a
covered species, affords no habitat protection when the birds are not
present, and provides only very limited mechanisms for addressing
chronic threats to covered species, such as nonnative predators.
The Lacey Act authorizes the Secretary of the Interior to list as
``injurious'' any wildlife deemed to be harmful to human beings, to the
interests of agriculture, horticulture, forestry, or to wildlife or the
wildlife resources of the United States. The Service inspects arriving
wildlife products, and enforces the injurious wildlife provisions of
the Lacey Act. Among other provisions, the Lacey Act prohibits
importation of injurious mammals, birds, fish, amphibians and reptiles
listed in the Lacey Act or which are declared by the Secretary of the
Interior through regulation to be injurious to human beings,
agriculture, horticulture, forestry or wildlife; however, these
prohibitions do not apply to plants and organisms other than those
listed or designated by regulations as injurious wildlife (USFWS 2016,
in litt.). The current list of animals considered as ``injurious
wildlife'' is provided at 50 CFR part 16. The list includes fruit bats,
mongoose, European rabbits and hares, wild dogs, rats or mice, raccoon
dogs, brushtail possum (the species introduced to New Zealand),
starlings, house sparrows, mynas, dioch, Java sparrows, red whiskered
bulbuls, walking catfish, mitten crabs, zebra mussels, fish in the
snakehead family, four species of carp, salmonids, brown tree snakes,
and pythons (USFWS 2012, 50 CFR part 16). The Lacey Act requires
declarations of importation only for formal entries (i.e., commercial
shipments), but not for informal entries (i.e., personal shipments)
(USDA-APHIS 2015, in litt.). Additionally, a species may still be
imported or transported across State lines while it is being considered
for addition to the list of ``injurious wildlife'' (Fowler et al. 2007,
pp. 357-358). Mongoose, rabbits, rats, mice, house sparrows, mynas,
Java sparrows, and red whiskered bulbuls are already established in
Hawaii, are difficult and costly to control, or are not controlled at
all. None of the aquatic species on the injurious species list is
present in Hawaii.
The continued spread of injurious species indicates the limited
effectiveness of the Lacey Act in preventing introductions of such
species to the State (Fowler et al. 2007, p. 357). As an example of
continued introduction of nonnative species in Hawaii, opossums
(Didelphis virginiana) have been found in shipping containers on Oahu
in 2005, 2011, and most recently in 2015 (Star Advertiser 2015b, in
litt). This species is not included on the Lacey Act's list of
injurious wildlife. Opossums are omnivorous scavengers, consuming a
wide variety of food items including insects, small vertebrates, bird
eggs, slugs and snails, and fruits and berries (Oregon Department of
Fish and Wildlife 2015, in litt.; Clermont College 2015, in litt.). If
opossums were to establish wild populations in Hawaii, their predation
on ground-nesting seabirds, insects, and snails could negatively affect
the band-rumped storm petrel, the orangeblack Hawaiian damselfly, one
or more of the 39 plants, and endangered snail species.
The Department of Agriculture-Animal and Plant Health Inspection
Service-Plant Protection and Quarantine (USDA-APHIS-PPQ) inspects
propagative plant material, provides identification services for
arriving plants and animals, conducts pest risk assessments, and other
related matters, but focuses on pests of wide concern across the United
States (HDOA 2009, in litt.). The USDA-APHIS-PPQ's Restricted Plants
List restricts the import of a limited number of noxious weeds. If not
specifically prohibited, current Federal regulations allow plants to be
imported from international ports with some restrictions. The Federal
Noxious Weed List (see 7 CFR 360.200; USDA 2012) includes more than 100
of the many globally known invasive plants, 21 of which are already
established in Hawaii. Plants on the list do not require a weed risk
assessment prior to importation from international ports.
A local organization (under the Institute of Pacific Islands
Forestry-USFS), Pacific Island Ecosystems at Risk (PIER) has compiled a
complete list of those plant species that are a threat to ecosystems in
the Pacific Islands, and those that are potentially invasive and are
present in the Pacific Region, along with a weed-risk assessment for
most of them (https://www.hear.org/pier/, last updated May 15, 2013).
There are over 1,000 plant species on the PIER list, and, in our
proposed rule (80 FR 58820, September 30, 2015; see pp. 58869-58881),
we discuss 114 of these invasive plants that currently have the
greatest impacts on the 49 species. In addition, the USDA-APHIS-PPQ is
in the process of finalizing rules to include a weed risk assessment
for plants newly imported to Hawaii (and that may not yet appear on the
PIER list).
Water extraction is a threat to the plant species (Cyclosorus
boydiae), the orangeblack Hawaiian damselfly, and the anchialine pool
shrimp Procaris hawaiana. The U.S. Army Corps of Engineers (COE) has
regulatory jurisdiction under section 404 of the Clean Water Act (33
U.S.C. 1251 et seq.) for activities that would result in a discharge of
dredged or fill material into waters of the United States; however, in
issuing permits for such activities, the COE does not typically
establish minimal instream flow standards (IFS) as a matter of policy
(U.S. Army 1985, RGL 85-6).
State Regulatory Mechanisms
The Hawaii Endangered Species law (HRS 195D) prohibits take,
possession, sale, transport or commerce in designated species. This
includes aquatic as well as terrestrial animal species, and terrestrial
plants (not freshwater or marine plants). This State law also
recognizes as endangered or threatened those species determined to be
endangered or threatened pursuant to the Act. This Hawaii law states
that a threatened species (under the Act) or an indigenous species may
be determined to be an endangered species under State law. Protection
of these species is under the authority of Hawaii's Department of Land
and Natural Resources, and under administrative rule (HAR 13-5-2-Ch
124). Although this State law can address threats such as habitat
modification, light attraction, and line collision through HCPs that
address the effects of individual projects or programs, it does not
address the pervasive threats to the 49 species posed by nonnative
predators and feral ungulates.
The importation of nondomestic animals, including aquatic species
and microorganisms, is regulated by a permit system (HAR 4-71) managed
through the Hawaii Department of Agriculture (HDOA). In addition,
transport of plants and plant parts between Hawaiian Islands is managed
through the HDOA (HAR 4-6-72), but only for those species that have
already been determined to be pest species. The objective of these
administrative rules is to implement the requirements of HRS 11-3-150A.
The list of nondomestic
[[Page 67845]]
animals (HAR 4-71) is defined by providing a list of those animals
considered domestic: dog, cat, horse, ass (burro or donkey), cattle and
beefalo, sheep, goat, swine, pot-bellied pig, alpaca, llama, rabbit,
chicken, turkeys, pigeons, ducks, geese, and their hybrids. Examples of
regulated pests are listed at HAR 4-72, including nonnative insects,
slugs, insects, plants, and plant viruses that can damage or harm
commercial crops. The HDOA's Board of Agriculture maintains lists of
nondomestic animals that are prohibited from entry, animals without
entry restrictions, or those that require a permit for import and
possession. The HDOA requires a permit to import animals, and
conditionally approves entry for individual possession, businesses
(e.g., pets and resale trade, retail sales, and food consumption), or
institutions. However, habitat destruction and modification, and
predation, by feral domestic animals (such as goat and cats,
respectively) are two primary threats to the 49 species not addressed
by the HDOA prohibitions and permitting process.
The State of Hawaii allows importation of most plant taxa, with
limited exceptions, if shipped from domestic ports (HLRB 2002; USDA-
APHIS-PPQ 2010; CGAPS 2009). Hawaii's plant import rules (HAR 4-70)
regulate the importation of 13 plant taxa of economic interest,
including pineapple, sugarcane, palms, and pines. Certain horticultural
crops (e.g., orchids) may require import permits and have pre-entry
requirements that include treatment or quarantine or both either prior
to or following entry into the State.
Critical biosecurity gaps include inadequate staffing, facilities,
and equipment for Federal and State inspectors devoted to invasive
species interdiction (HLRB 2002; USDA-APHIS-PPQ 2010; CGAPS 2009). In
recognition of these gaps, a State law has been passed that allows the
HDOA to collect fees for quarantine inspection of freight entering
Hawaii (Act 36 (2011) HRS 150A-5.3). Legislation enacted in 2011 (H.B.
1568) requires commercial harbors and airports to provide biosecurity
and inspection facilities to facilitate the movement of cargo through
ports. This bill is a significant step toward optimizing biosecurity
capacity in the State, but only time will determine its effectiveness
(Act 2011 (11)). We believe there is a need for all civilian and
military port and airport operations and construction to make
biosecurity concerns a core objective.
As an example, the threat of introduction of nonnative species is
evidenced by the 2013 discovery of presence of the nonnative coconut
rhinoceros beetle (CRB, Oryctes rhinoceros), which quickly spread from
its known point of introduction across the island of Oahu in a few
months (HISC 2014, in litt. + maps; HDOA 2014, in litt.). The CRB is
considered one of the most damaging insects to coconut and African oil
palm trees in southern and southeast Asia, as well as the western
Pacific Islands, and could devastate populations of native and
nonnative palm trees in Hawaii (Giblin-Davis 2001 in HISC 2014, in
litt.). A rapid response team headed by HDOA (with USDA, University of
Hawaii, U.S. Navy, and other partners) has set up pheromone traps
island-wide, and capture and range delineation efforts are ongoing,
along with funding for support services to control the CRB (HISC 2014,
in litt.). However, existing regulatory mechanisms did not prevent the
introduction of this pest species into Hawaii. These regulatory
mechanisms, such as HAR 71A and HAR 71C (regarding release of nonnative
species) and H.B. 1568 (pertaining to State law to enforce biosecurity
measures), therefore, are inadequate to prevent introduction of
nonnative species. Efforts to control the CRB continue, but whether
those efforts will be effective is unknown at this time.
Hawaii's noxious weed law was enacted to prevent the introduction
and transport of noxious weeds or their seeds or vegetative
reproductive parts into any area that is reasonably free of those
noxious weeds (HRS 11-3-152), and it states that the Hawaii Department
of Agriculture shall take necessary measures to restrict the
introduction and establishment of specific noxious weeds in such areas.
Hawaii administrative rule (HAR 4-6-68) further defines the term
``noxious weed'' and the criteria for designation of plants as such and
criteria for designation of a noxious weed ``free area.'' The list of
noxious weeds, compiled in 1992, consists of 79 plant species, 49 of
which were not yet established in Hawaii. Since that time, 20 species
on the list have become established in Hawaii: Bocconia frutescens
(plume poppy), Cereus uruguayanus (spiny tree cactus), Chromolaena
odorata (siamweed), Cortaderia jubata (Andean pampas grass), Cytisus
scoparius (Scotch broom), Hyptis suaveolens (wild spikenard), Malachra
alceifolia (malachra), Melastoma spp. (melastoma; two species now
established, M. candidum and M. sanguineum), Miconia spp. (miconia; M.
calvescens now on four islands), Passiflora pulchella (wingleaf
passionfruit), Piper aduncum (spiked pepper), Prosopis juliflora
(algarroba), Pueraria phaseoloides (tropical kudzu), Rubus sieboldii
(Molucca raspberry), Senecio madagascariensis (fireweed), Solanum
elaeagnifolium (silverleaf nightshade), Solanum robustum (shrubby
nightshade), Solanum torvum (turkeyberry), and Spartium junceum
(Spanish broom). Thus, despite State legislation and regulations
addressing invasive and noxious species, their entry into the State
continues.
The State manages the use of surface and ground water resources
through the Commission on Water Resource Management (CWRM), as mandated
by the State Water Code (HRS 174, HAR 13-168-196). The State considers
all natural flowing surface water (streams, springs, seeps) as State
property (HRS 174C), and the DLNR has management responsibility for the
aquatic organisms in these waters (HRS Annotated 1988, Title 12 1992
Cumulative Supplement). In Hawaii, instream flow is regulated by
establishing standards on a stream-by-stream basis. The standards
currently in effect represent flow conditions in 1987 (status quo), the
year the administrative rules (State Water Code) were adopted (HRS
174C-71, HAR title 13, ch 169-44-49). Because of the complexity of
establishing instream flow standards (IFS) for 376 perennial streams,
the Commission retains interim IFS at status quo levels as set in 1987
(CWRM 2009, in litt.; CRWM 2014, in litt). In the Waiahole Ditch
Combined Contested Hearing on Oahu (1991-2006), the Hawaii Supreme
Court determined that status quo interim IFS were not adequate, and
required the Commission to reassess the IFS for Waiahole Ditch and
other streams Statewide (Cast No. CCH-OA95-1; Maui Now.com, in litt.).
The Commission has been gathering information to fulfill this
requirement since 2006, but no IFS recommendations have been made to
date (CWRM 2008, p. 3-153; CRWM 2014, in litt.).
In addition, in the Hawaii Stream Assessment Report (HDLNR 1990;
prepared in coordination with the National Park Service (NPS)), the
Commission identified high-quality rivers and streams (and portions
thereof) that may be placed within a Wild and Scenic River System. This
report ranked 70 out of 176 analyzed rivers and streams as outstanding
high-quality habitat, and recommended that streams meeting certain
criteria be protected from further development (HDLNR 1990, pp. xxi-
xxiv). However, there is no mechanism within the State's Water Code to
designate and set aside these
[[Page 67846]]
streams, or to identify and protect stream habitat. Accordingly,
damselfly populations (including the orangeblack Hawaiian damselfly)
are at risk of continued loss of habitat.
Hawaii's DLNR Division of Aquatic Resources (DAR) is responsible
for conserving, protecting, and enhancing the State's renewable
resources of aquatic life and habitat (HDAR 2015, in litt.; DLNR-DAR
2003, p. 3-13). The release of live nonnative fish or other nonnative
aquatic life into any waters of the State is prohibited (HRS 187A-6.5),
and DAR has the authority to seize, confiscate, or destroy as a public
nuisance any of these prohibited species (HRS 187A-2; HRS 187A-6.5).
However, the DAR recognizes that nonnative species continue to enter
the State and move between islands (DLNR-DAR 2003, p. 2-12).
There are no existing regulatory mechanisms that specifically
protect Hawaii's anchialine pools (habitat for the anchialine pool
shrimp Procaris hawaiana and the orangeblack Hawaiian damselfly);
however 2 anchialine pools on Maui and 12 anchialine pool on Hawaii
Island are located within State NARs. State NARs were created to
preserve and protect samples of Hawaii's ecosystems and geological
formations, and are monitored. Designation as a State NAR prohibits the
removal of any native organism and the disturbance of pools (HAR 13-
209-4). Though signs are posted at NARs to notify the public that
anchialine pools are off-limits to bathers, off-road vehicle use around
the pools, and other activities, the anchialine pools are in remote
areas and the State does not have sufficient funding to effectively
enforce those restrictions.
Nonnative ungulates pose threats of habitat destruction and
modification and predation (herbivory) to 37 of the 39 plants species,
and of habitat destruction and modification to 9 of the 10 animals in
this rule (see Table 2). The State provides opportunities to the public
to hunt game mammals (ungulates including feral pigs and goats, axis
deer, black-tailed deer, and mouflon, sheep and mouflon-sheep hybrids)
on 91 State-designated public hunting areas (within 45 units) on all
the main Hawaiian Islands except Kahoolawe and Niihau (HAR-DLNR 2010,
13-123; HDLNR 2009b, pp. 25-30). On Niihau, public hunting
opportunities are managed by a private business (Niihau Safaris Inc.
2015, in litt.). The State's management objectives for game mammals
range from maximizing public hunting opportunities (i.e., ``sustained
yield'') in some areas to removal by State staff or their designees
from other areas (HAR-DLNR 2010, p. 12-123; HDLNR 2009b, pp. 25-30).
Thirty of the 39 plant species, the band-rumped storm-petrel, the
orangeblack Hawaiian damselfly, and three yellow-faced bees have
populations in areas where habitat is used for game enhancement and
game populations are maintained at levels for public hunting (Holmes
and Joyce 2009, 4 pp.; HAR-DLNR 2010, p. 12-123; HBMP 2010). Public
hunting areas are defined, but not fenced, and game mammals have
unrestricted access to most areas across the landscape, regardless of
underlying land-use designation. While fences are sometimes built to
protect areas from game mammals, the current number and locations of
fences are not adequate to prevent habitat destruction and modification
for 46 of the 49 species. One additional State regulation (HRS 12-183D)
was enacted recently to prevent intra-island transport of axis deer
only. There are no other State regulations than those described above
that address protection of the species and their habitat from feral
ungulates.
Under statutory authorities provided by HRS title 12, subtitle 4,
183D Wildlife, the DLNR maintains HAR ch 124 (2014), which defines
``injurious wildlife'' as ``any species or subspecies of animal except
game birds and game mammals which is known to be harmful to
agriculture, aquaculture, indigenous wildlife or plants, or constitute
a nuisance or health hazard and is listed in the exhibit entitled
Exhibit 5, Chapter 13-124, List of Species of Injurious Wildlife in
Hawaii.'' Under HAR 13-124-3(d), ``no person shall, or attempt to: (1)
Release injurious wildlife into the wild; (2) Transport them to island
or locations within the State where they are not already established
and living in a wild state; and (3) Export any such species or the dead
body or parts thereof, from the State. Permits for these actions may be
considered on a case-by-case basis.'' This law was enacted after an
incident in 2012 of interisland transport of axis deer (for hunting
purposes) to Hawaii Island, which was without axis deer previously.
Local Mechanisms
Local biologists and botanists recognize the urgent need to address
the importation of nonnative, invasive species, and are working to
implement actions required; however, their funding is not guaranteed.
We discuss the four primary groups below.
In 1995, the Coordinating Group on Alien Pest Species (CGAPS), a
partnership of managers from Federal, State, County, and private
agencies and organizations involved in invasive species work in Hawaii,
was formed in an effort to coordinate policy and funding decisions,
improve communication, increase collaboration, and promote public
awareness (CGAPS 2009). This group facilitated the formation of the
Hawaii Invasive Species Council (HISC), which was created by
gubernatorial executive order in 2002, to coordinate local initiatives
for the prevention of introduction and for control of invasive species
by providing policy-level direction and planning for the State
departments responsible for invasive species issues (CGAPS 2009). In
2003, the Governor signed into law Act 85, which conveys statutory
authority to the HISC to continue to coordinate approaches among the
various State and Federal agencies, and international and local
initiatives, for the prevention and control of invasive species (HDLNR
2003, p. 3-15; HISC 2009, in litt.; HRS 194-2). Some of the recent
priorities for the HISC include interagency efforts to control
nonnative species such as the plants Miconia calvescens (miconia) and
Cortaderia spp. (pampas grass), coqui frogs (Eleutherodactylus coqui),
the CRB, and ants (HISC 2009, 2013, and 2015, in litt.; OISC 2015, in
litt.; https://dlnr.hawaii.gov/hisc). Budget cuts beginning in 2009
restricted State funding support of HISC, resulting in a serious
setback of conservation efforts (HISC 2009; HISC 2015).
The Hawaii Association of Watershed Partnerships comprises 11
separate partnerships on six Hawaiian Islands. These partnerships are
voluntary alliances of public and private landowners, ``committed to
the common value of protecting forested watersheds for water recharge,
conservation, and other ecosystem services through collaborative
management'' (https://hawp.org/partnerships). Funding for the
partnerships is provided through a variety of State and Federal
sources, public and private grants, and in-kind services provided by
the partners and volunteers. However, budget cuts of 40 to 60 percent
have occurred since 2009, with serious impacts to the positive
contributions of these groups to implementing the laws and rules that
can protect and control threats to one or more of the 49 species.
Another group was established to coordinate State and Federal
agency efforts in the protection of rare endemic plant species in the
State and Guam and the Commonwealth of the Northern Mariana Islands
(CNMI), Hawaii's Plant Extinction Prevention Program (PEPP). This
program identifies and supports the ``rarest of the rare'' plant
species in
[[Page 67847]]
need of immediate conservation efforts. The goal of PEPP is to prevent
the extinction of plant species that have fewer than 50 individuals
remaining in the wild.
These four partnerships, CGAPS, HISC, HAWP, and PEPP, are stop-gap
measures that attempt to address issues that are not resolved by
individual State and Federal agencies. The capacity of State and
Federal agencies and their nongovernmental partners in Hawaii to
provide sufficient inspection services, enforce regulations, and
mitigate or monitor the effects of nonnative species is limited due to
the large number of taxa currently causing damage (CGAPS 2009). Many
invasive, nonnative species established in Hawaii currently have
limited but expanding ranges, and they cause considerable concern.
Resources available to reduce the spread of these species and counter
their negative effects are limited. Control efforts are focused on a
few invasive species that cause significant economic or environmental
damage to commercial crops and public and private lands. Comprehensive
control of an array of nonnative species and management to reduce
disturbance regimes that favor them remain limited in scope. If current
levels of funding and regulatory support for control of nonnative
species are maintained, the Service expects existing programs to
continue to exclude, or, on a very limited basis, control these species
only in the highest priority areas. Threats from established nonnative
species are ongoing and are expected to continue into the future.
As an example of current and future challenges for biosecurity in
Hawaii, a strain of the plant rust Puccinia psidii (ohia rust) was
first noticed affecting stands of the nonnative rose apple (Syzygium
jambos) and the native Metrosideros (ohia) seedlings (both in the plant
family Myrtaceae) in nurseries in 2005. Metrosideros spp. are a
dominant component of native forest in Hawaii, providing watershed
protection and habitat for native wildlife. The Hawaii Board of
Agriculture recommended a quarantine rule be passed against the
introduction of all new strains of ohia rust (through transmission on
Myrtaceae species used in the horticulture trade), to prevent
destruction of ohia forests and the risk to agriculture and
horticulture industries (Environment Hawaii 2015, pp. 1,8-9). However,
the rule remains in draft form and under review (HDOA 2015, in litt.),
accessed August 1, 2016). An example of the failure of biosecurity in
Hawaii and the speed with which a new invader can cause widespread
destruction is the introduction of the gall wasp Quadrastichus
erythrinae. This highly destructive wasp was detected in Taiwan in
2003. Despite evidence of its rapid advance across the Pacific Basin
with concomitant loss of populations of native and ornamental trees in
the genus Erythrina, this wasp arrived and naturalized in Hawaii in
2005 (Gramling 2005, p. 1). The wasp dispersed throughout the main
Hawaiian Islands within weeks, and as a result, the endemic wiliwili,
Erythrina sandwicensis, was quickly devastated (Rubinoff et al. 2010,
p. 24).
On the basis of the information provided above, existing State and
Federal regulatory mechanisms are not preventing the introduction of
nonnative species and pathogens into Hawaii via interstate and
international pathways, or via intrastate movement of nonnative species
between islands and watersheds. Nor do these mechanisms address the
current threats posed to the 49 species by established nonnative
species. Therefore, State and Federal regulatory mechanisms do not
adequately protect the 49 species, or their habitats, from the threat
of new introductions of nonnative species or the continued expansion of
nonnative species populations on and between islands and watersheds.
The impacts from these threats are ongoing and are expected to continue
into the future.
Summary of Factor D
Existing State and Federal regulatory mechanisms are not preventing
the introduction into Hawaii of nonnative species or controlling the
spread of nonnative species between islands and watersheds, or
establishing or maintaining instream flow standards. Water extraction
is a threat to one plant, Cyclosorus boydiae, to the orangeblack
Hawaiian damselfly, and the anchialine pool shrimp (Factor A). Habitat-
altering ungulates and nonnative plants (Factor A) pose major ongoing
threats to all 49 species addressed in this rule. Thirty-five of the 39
plants and all 10 animals experience the threat of predation or
herbivory by nonnative animals (Factor C). The seven yellow-faced bees
and the orangeblack Hawaiian damselfly experience competition with
nonnative insect species (Factor E). The intentional or inadvertent
introduction of nonnative species and their spread within Hawaii, and
the damage caused by existing populations of nonnative species,
continues despite existing regulatory mechanisms designed to address
this threat (in all its manifestations described above) to all 49
species. No existing regulatory mechanisms effectively address
maintenance of instream flow, springs, seeps, and anchialine pools or
address the threats of water extraction and stream modification for the
anchialine pool shrimp and orangeblack Hawaiian damselfly. All of these
threats are ongoing and are expected to continue into the future;
therefore, we conclude the existing regulatory mechanisms are
inadequate to reduce or eliminate these threats to the 49 species.
Factor E. Other Natural or Manmade Factors Affecting Their Continued
Existence
Other factors that pose a threat to some or all of the 49 species
include artificial lighting and structures, ingestion of marine debris
and plastics, dumping of trash and the introduction of nonnative fish
into anchialine pools, recreational use of and sedimentation of
anchialine pools, low numbers of individuals and populations,
hybridization, lack of or declining regeneration, competition with
nonnative invertebrates, and loss of host plants. Each threat is
discussed in detail below, along with identification of which species
are affected by these threats. The impacts of climate change to these
species and their ecosystems have the potential to exacerbate all of
the threats described below.
Artificial Lighting and Structures Effects on the Band-Rumped Storm-
Petrel
Artificial lights are a well-documented threat to night-flying
seabirds such as petrels, shearwaters, and storm-petrels (Croxall et
al. 2012, p. 28). A significant impact to the band-rumped storm-petrel
results from the effects of artificial (night) lighting on fledglings
and, to a lesser degree, on adults. Lighting of roadways, resorts,
ballparks, residences, and other development, as well as on cruise
ships out at sea, both attracts and confuses night-flying storm-petrels
and other seabirds (Harrison et al. 1990, p. 49; Reed et al. 1985, p.
377; Telfer et al. 1987, pp. 412-413; Banko et al. 1991, p. 651).
Storm-petrels use the night sky to navigate and possibly to search for
bioluminescent marine prey (Telfer et al. 1987, p. 412). Artificial
lights can attract night-flying seabirds and result in ``fallout''
(birds becoming grounded) when birds become confused and exhaust
themselves circling around lights or collide with buildings,
powerlines, or other structures. Once grounded, these birds are at risk
of predation or being run over by cars (Reed et al. 1985, p. 377;
Telfer et al. 1987, p. 410). Vulnerability to artificial lighting
varies among species and age classes and is influenced by season, lunar
phase, and weather conditions.
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Young birds are more likely than adults to become disoriented by
manmade light sources (Montevecchi 2006, pp. 101-102). Over a 12-year
period (1978 to 1990), Harrison et al. (1990, p. 49) reported that 15
band-rumped storm-petrels, 13 of which were fledglings, were recovered
on Kauai as a result of fallout. Between 1991 and 2008, another 21
band-rumped storm-petrels were collected on Kauai (Holmes and Joyce
2009, p. 2). Currently, fallout due to light pollution is recorded
almost annually on Kauai (Kauai Island Utility Cooperative 2015, in
litt.). In addition, band-rumped storm-petrels may be attracted to
lights at sea and collide with boats; this source of injury and
mortality has been documented for other storm-petrel species (e.g.,
Black 2005, p. 67). The actual extent of such loss and its overall
impact on the band-rumped storm-petrel population in Hawaii is not
known because scavengers often prevent the detection or recovery of the
dead or injured birds, and the scattered and remote nesting areas of
this species preclude demographic monitoring to quantify the impacts of
this source of mortality. However, given the probable small total
number of band-rumped storm-petrels nesting in Hawaii and the threats
they face from nonnative predators such as rats and cats, any
additional mortalities are likely to have negative impacts on the
population.
A related threat to seabirds in Hawaii, including the band-rumped
storm-petrel, is collision with structures such as communication towers
and utility lines (Cooper and Day 1998, pp. 16-18; Podolsky et al.
1998, pp. 23-33). Several seabird species that nest in the Hawaiian
Islands, including the Newell's Townsend's shearwater (federally listed
as threatened), the Hawaiian petrel (federally listed as endangered),
and the band-rumped storm-petrel, regularly commute between inland nest
sites and the ocean. These birds commute at night, when manmade
obstacles such as communication towers and utility lines are difficult
to see. They strike these unseen obstacles, and often die or are
injured as a result. An early study estimated that 340 Newell's
Townsend's shearwater fledglings die annually on the eastern and
southern shores of Kauai as a result of collisions (Podolsky et al.
1998, p. 30); however, current analyses for all seabirds on Kauai
indicate the number of collisions with utility lines is much higher,
over 2,000 strikes per year (using site-specific strike rates), but
numbers of birds that hit utility lines is site-dependent (Travers et
al. 2014, pp. 19, 29-37; Service 2015, in litt., Slide 21). Absent
preventative measures, the impact to the band-rumped storm-petrel from
artificial lighting and collisions with structures is expected to
increase as the human population grows and development continues on the
Hawaiian Islands.
Other Human Effects on the Band-Rumped Storm-Petrel
Other factors that may negatively affect the band-rumped storm-
petrel include commercial fisheries interactions and alteration of prey
base upon which the band-rumped storm-petrel depends. Commercial
fisheries are known to adversely affect certain species of seabirds
(Furness 2003, pp. 33-35). Seabirds are caught in fishing gear and
suffer mortality by drowning. Seabirds also come into contact with and
consume deep-water fish to which they would not normally have access,
and can become contaminated by high levels of heavy metals in these
fish (Furness 2003, p. 34). Commercial fisheries also cause depletion
of small pelagic schooling fish, a significant food source for seabirds
(Furness 2003, p. 34). The potential effects of these activities have
not been assessed for the band-rumped storm-petrel; however, storm-
petrels have been observed to attend fishing vessels (e.g., Yorio and
Caille 1999, p. 21; Yeh et al. 2013, p. 146), and the effects of
fishery interactions on this species are likely to be similar to those
documented for other seabird species in the same order
(Procellariiformes or tubenoses; albatrosses and petrels). In addition,
plastics and other debris in the open ocean can be ingested
accidentally by band-rumped storm-petrels and pose a threat to this
species (Ryan 1989, p. 629). Although a study by Moser and Lee (1992,
p. 85) found no evidence of plastic ingestion by band-rumped storm-
petrels, the sample size was very small (4 individuals) and inadequate
to conclusively determine whether this species suffers from ingestion
of plastics. Other species of storm-petrels have been documented to
ingest plastics (Bond and Lavers 2013, p. 3; Ryan 2015, p. 20; Wilcox
et al. 2015, p. 3), and band-rumped storm-petrels are likely to do so
also. Many closely related seabirds do suffer ill effects from
ingestion of plastics, including physical damage to the digestive
tract, effects of toxins carried on the plastics, and resulting
mortality (Ryan 1989, pp. 623-629; Tanaka et al. 2013, pp. 2-3).
Effects of Recreational Use, and Dumping of Trash and Nonnative Fish
Into Anchialine Pools
On Hawaii Island, it is estimated that up to 90 percent of the
anchialine pools have been destroyed or altered by human activities
(Brock 2004, p. i). The more recent human modification of anchialine
pools includes bulldozing and filling of pools (Bailey-Brock and Brock
1993, p. 354). Trampling damage from use of anchialine pools for
swimming and bathing has been documented (Brock 2004, pp. 13-17).
Historically, pools were sometimes modified with stone walls and steps
by Hawaiians who used them for bathing. There are no documented
negative impacts to pond biota as a result of this activity; however,
introduction of soaps and shampoos is of concern (Brock 2004, p. 15).
The depressional features of anchialine pools make them susceptible
to dumping. Refuse found in degraded pools and pools that have been
filled with rubble have been dated to about 100 years old, and the
practice of dumping trash into pools continues today (Brock 2004, p.
15). For example, Lua O Palahemo (Hawaii Island) is located
approximately 560 ft (170 m) from a sandy beach frequented by visitors
who fish and swim. There are multiple dirt roads that surround the pool
making it highly accessible. Plastic bags, paper, fishing line, water
bottles, soda cans, radios, barbed wire, and a bicycle have been
documented within the pool (Kensley and Williams 1986, pp. 417-418;
Bozanic 2004, p. 1; Wada 2010, in litt.). Introduction of trash
involving chemical contamination into anchialine pools, as has been
observed elsewhere on Hawaii Island (Brock 2004, pp. 15-16),
drastically affects water quality and results in local extirpation of
anchialine pool shrimp species.
Anchialine pool habitats can gradually disappear when wind-blown
materials accumulate through a process known as senescence (Maciolek
and Brock 1974, p. 3; Brock 2004, pp. 11, 35-36). Conditions promoting
rapid senescence include an increased amount of sediment deposition,
good exposure to light, shallowness, and a weak connection with the
water table, resulting in sediment and detritus accumulating within the
pool instead of being flushed away with tidal exchanges and ground
water flow (Maciolek and Brock 1974, p. 3; Brock 2004, pp. 11, 35-36).
Sedimentation degrades the health of Hawaiian anchialine pool systems
in which the anchialine pool
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shrimp, Procaris hawaiana, and the orangeblack Hawaiian damselfly
occur.
In general, the accidental or intentional introduction and spread
of nonnative fishes (bait and aquarium fish) is considered the greatest
threat to anchialine pools in Hawaii (Brock 2004, p. 16). Maciolek
(1983, p. 612) found that the abundance of shrimp in a given population
is indirectly related to predation by fish. Lua O Palahemo is
vulnerable to the intentional dumping of nonnative bait and aquarium
fishes because the area is accessible to vehicles and human traffic,
although due to its remote location, it is not monitored regularly by
State agency staff. The release of mosquito fish and tilapia into the
Waikoloa Anchialine Pond Preserve (WAAPA) at Waikoloa, North Kona,
Hawaii, resulted in the infestation of all ponds within an
approximately 3-ha (8-ac) area, which represented about two-thirds of
the WAAPA. Within 6 months, all native hypogeal (subterranean) shrimp
species disappeared (Brock 2004, p. iii). Nonnative fish drive
anchialine species out of the lighted, higher productivity portion of
the pools, into the surrounding water table bed rock, subsequently
leading to the decimation of the benthic community structure of the
pool (Brock 2004, p. iii). In addition, nonnative fish prey on and
exclude native hypogeal shrimp that are usually a dominant and
essential faunal component of anchialine pool ecosystems (Brock 2004,
p. 16; Bailey-Brock and Brock 1993, pp. 338-355). The loss of the
shrimp changes ecological succession by reducing herbivory of
macroalgae, allowing an overgrowth and change of pool flora. This
overgrowth changes the system from clear, well-flushed basins to a
system characterized by heavy sedimentation and poor water exchange,
which increases the rate of pool senescence (Brock 2004, p. 16).
Nonnative fishes, unlike native fishes, are able to complete their life
cycles within anchialine pool habitats, and remain a permanent
detrimental presence in all pools in which they are introduced (Brock
2004, p. 16). In Hawaii, the most frequently introduced fishes are
those in the Poeciliidae family (freshwater fish which bear live young)
and include mosquito fish, various mollies (Poecilia spp.), and
tilapia, that prey on and exclude the herbivorous aquatic animals upon
which Procaris hawaiana feeds. More than 90 percent of the 600 to 700
anchialine habitats in the State of Hawaii were degraded between 1974
and 2004, due to the introduction of nonnative fishes (Brock 2004, p.
24). According to Brock (2012, pers. comm.), sometime in the 1980s,
nonnative fishes were introduced into Lua O Palahemo. It is our
understanding that the fish were subsequently removed by illegal use of
a fish poison (EPA 2007. pp. 22-23; Finlayson et al. 2010, p. 2), and
to our knowledge the pool is currently free of nonnative fish; however,
nonnative fish could be introduced into the pool at any time.
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