Endangered and Threatened Wildlife and Plants; 90-Day Finding on Five Petitions to List Seven Species of Hawaiian Yellow-faced Bees as Endangered, 34077-34088 [2010-14430]
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[Docket No. FWS-R1-ES-2010–0012]
[MO 92210-0-0008-B2]
Endangered and Threatened Wildlife
and Plants; 90-Day Finding on Five
Petitions to List Seven Species of
Hawaiian Yellow-faced Bees as
Endangered
AGENCY: Fish and Wildlife Service,
Interior.
ACTION: Notice of 90–day petition
finding and initiation of status review.
SUMMARY: We, the U.S. Fish and
Wildlife Service (Service), announce a
90–day finding on five petitions to list
seven species of Hawaiian yellow-faced
bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana) as
endangered and designate critical
habitat under the Endangered Species
Act of 1973, as amended (Act).
We find that the petitions present
substantial scientific or commercial
information indicating that listing these
seven species of Hawaiian yellow-faced
bees may be warranted. Therefore, with
the publication of this notice we are
initiating a status review of these
species and will issue 12–month
findings on our determination as to
whether the petitioned actions are
warranted. To ensure that the status
reviews are comprehensive, we are
soliciting scientific and commercial data
and other information regarding these
species. We will make a determination
on critical habitat for these species if,
and when, we initiate a listing action.
DATES: To allow us adequate time to
conduct this review, we request that
information you submit be received by
us on or before August 16, 2010. Please
note that if you are using the Federal
eRulemaking Portal (see ‘‘ADDRESSES’’
section, below), the deadline for
submitting an electronic comment is
Eastern Standard Time on this date.
We will post all information received
on https://www.regulations.gov. This
generally means that we will post any
personal information you provide us
(see the Request for Information section
below for more details).
After August 16, 2010, you must
submit information directly to the Field
Office (see FOR FURTHER INFORMATION
CONTACT section below). Please note that
we might not be able to address or
incorporate information that we receive
after the above requested date.
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ADDRESSES: You may submit
information by one of the following
methods:
• Federal eRulemaking Portal: https://
www.regulations.gov. In the box that
reads ‘‘Enter Keyword or ID,’’ enter the
Docket number for this finding, which
is FWS-R1-ES-2010-0012. Check the box
that reads ‘‘Open for Comment/
Submission,’’ and then click the Search
button. You should then see an icon that
reads ‘‘Submit a Comment.’’ Please
ensure that you have found the correct
rulemaking before submitting your
comment.
• U.S. mail or hand-delivery: Public
Comments Processing, Attn: FWS-R1ES-2010-0012; Division of Policy and
Directives Management; U.S. Fish and
Wildlife Service; 4401 N. Fairfax Drive,
Suite 222; Arlington, VA 22203.
FOR FURTHER INFORMATION CONTACT:
Loyal Mehrhoff, Field Supervisor,
Pacific Islands Fish and Wildlife Office,
300 Ala Moana Boulevard, Room 3-122,
Honolulu, HI 96850; by telephone (808–
792–9400); or by facsimile (808–792–
9581). Persons who use a
telecommunications device for the deaf
(TTD) may call the Federal Information
Relay Service (FIRS) at 800–877–8339.
SUPPLEMENTARY INFORMATION:
Request for Information
When we make a finding that a
petition presents substantial
information indicating that listing a
species may be warranted, we are
required to promptly review the status
of the species (status review). For the
status review to be complete and based
on the best available scientific and
commercial information, we request
information on the seven species of
Hawaiian yellow-faced bees (H.
anthracinus, H. assimulans, H. facilis,
H. hilaris, H. kuakea, H. longiceps, and
H. mana) from governmental agencies,
Native American Tribes, the scientific
community, industry, or any other
interested parties. We seek information
on:
(1) The species’ biology, range, and
population trends, including:
(a) Habitat requirements for feeding,
breeding, and sheltering;
(b) Genetics and taxonomy;
(c) Historical and current range
including distribution patterns;
(d) Historical and current population
levels, and current and projected trends;
and
(e) Past and ongoing conservation
measures for the species, its habitat or
both.
(2) Information about the seven
Hawaiian yellow-faced bees relevant to
the factors that are the basis for making
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a listing determination for a species
under section 4(a) of the Act (16 U.S.C.
1531 et seq.), which are:
(a) The present or threatened
destruction, modification, or
curtailment of the species’ 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.
(3) Whether insect collecting presents
a threat to any of the seven Hawaiian
yellow-faced bee species.
(4) The potential cumulative effects of
these factors that may threaten or
endanger the seven Hawaiian yellowfaced bee species.
(5) Management programs for the
conservation of the seven Hawaiian
yellow-faced bee species.
(6) The potential effects of climate
change on the seven Hawaiian yellowfaced bee species and their habitat.
If, after the status reviews, we
determine that listing any of the seven
Hawaiian yellow-faced bees is
warranted, we will propose critical
habitat (see definition in section 3(5)(A)
of the Act) under section 4 of the Act,
to the maximum extent prudent and
determinable at the time we propose to
list the species. Therefore, with regard
to areas within the geographical range
currently occupied by these species, we
also request data and information on
what may constitute physical or
biological features essential to the
conservation of these species; where
these features are currently found; and
whether any of these features may
require special management
considerations or protection. In
addition, we request data and
information regarding whether there are
areas outside the geographical area
occupied by these species that are
essential to the conservation of these
seven species. Please provide specific
comments and information as to what,
if any, critical habitat you think we
should propose for designation if these
species are proposed for listing, and
why such habitat meets the
requirements of section 4 of the Act.
Please include sufficient information
with your submission (such as scientific
journal articles or other publications) to
allow us to verify any scientific or
commercial information you include.
Please note that submissions merely
stating support for or opposition to the
action under consideration without
providing supporting information,
although noted, will not be considered
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in making a determination, as section
4(b)(1)(A) of the Act (16 U.S.C. 1531 et
seq.) directs that determinations as to
whether any species is a threatened or
endangered species must be made
‘‘solely on the basis of the best scientific
and commercial data available.’’
You may submit your information
concerning this status review by one of
the methods listed in the ADDRESSES
section. If you submit information via
https://www.regulations.gov, your entire
submission—including any personal
identifying information—will be posted
on the website. If your submission is
made via a hardcopy that includes
personal identifying information, you
may request at the top of your document
that we withhold this personal
identifying information from public
review. However, we cannot guarantee
that we will be able to do so. We will
post all hardcopy submissions on https://
www.regulations.gov.
Information and materials we receive,
as well as supporting documentation we
used in preparing this finding, is
available for you to review at https://
www.regulations.gov, or you may make
an appointment during normal business
hours at the Pacific Islands Fish and
Wildlife Office (see FOR FURTHER
INFORMATION CONTACT).
Background
Section 4(b)(3)(A) of the Act (16
U.S.C. 1533(b)(3)(A)) requires that we
make a finding on whether a petition to
list, delist, or reclassify a species
presents substantial scientific or
commercial information to indicate that
the petitioned action may be warranted.
We are to base this finding on
information provided in the petition,
supporting information submitted with
the petition, and information otherwise
available in our files. To the maximum
extent practicable, we are to make this
finding within 90 days of our receipt of
the petition and publish our notice of
this finding promptly in the Federal
Register.
Our standard for substantial scientific
or commercial information within the
Code of Federal Regulations (CFR) with
regard to a 90–day petition finding is
‘‘that amount of information that would
lead a reasonable person to believe that
the measure proposed in the petition
may be warranted’’ (50 CFR 424.14(b)).
If we find that substantial scientific or
commercial information was presented,
we are required to promptly conduct a
species status review, which we
subsequently summarize in our 12–
month finding.
We received five petitions dated
March 23, 2009, from Scott Hoffman
Black, Executive Director of the Xerces
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Society (hereafter, ‘‘the petitioner’’). The
petitioner requested that we list seven
species of Hawaiian yellow-faced bees
as endangered and that critical habitat
be designated under the Act. The
petitions were clearly identified as
petitions and included the requisite
identification of addresses and
telephone numbers, but did not include
the signature of the petitioner, as
stipulated in 50 CFR 424.14(a).
Nevertheless, we recognize the five
documents as petitions. Each petition
contained information regarding the
species’ taxonomy and ecology,
historical and current distribution,
present status, and potential causes of
decline and current and potential
imminent threats.
On May 8, 2009, we sent a letter to the
petitioner acknowledging our receipt of
the petitions and explaining that we
reviewed the information presented in
the petition and determined that issuing
an emergency regulation temporarily
listing the species under section 4(b)(7)
of the Act was not warranted at that
time. This notice constitutes our 90–day
finding for the petitions to list seven
species of Hawaiian yellow-faced bees
(Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana).
Species Information
Taxonomy and Description of Hylaeus
The seven species of bees described in
the petitions belong to the genus
Hylaeus. Hylaeus is a large, globally
distributed genus comprised of over 500
species worldwide. In the Hawaiian
Islands, the genus Hylaeus is
widespread and very diverse, with 60
native species, including 20 that are
endemic to single islands (Magnacca
2007, p. 174). All 60 Hawaiian species
are in the subgenus Nesoprosopis
(Magnacca and Danforth 2006, p. 393).
The Hylaeus genus belongs to the
Colletidae family of bees, also known as
plasterer bees due to their habit of lining
their nests with salival secretions.
The species of Hylaeus are commonly
known as yellow-faced bees or masked
bees for their yellow-to-white facial
markings. All of the Hylaeus species
roughly resemble small wasps in
appearance, due to their slender bodies
and their seeming lack of setae (sensory
hairs). However, Hylaeus bees have
plumose (branched) hairs on the body
that are longest on the sides of the
thorax. To a discerning eye, it is these
plumose setae that readily distinguish
them from wasps (Michener 2000, p.
55).
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Life History of Hylaeus
Nests of Hylaeus species are usually
constructed opportunistically within
dead twigs or plant stems, or other
similarly small natural cavities under
bark or rocks. This is unlike the nests of
many other bee species, which are
purposefully excavated or constructed
underground. Like other Hylaeus,
Hawaiian Hylaeus also lack strong
mandibles and other adaptations for
digging and often use nest burrows
abandoned by other insect species (Daly
and Magnacca 2003, p. 9). The adult
male and female bees feed upon flower
nectar for nourishment. Many species,
including the Hawaiian Hylaeus, lack an
external structure for carrying pollen,
called a scopa, and instead internally
transport collected pollen, often mixed
with nectar, within their crop (stomach).
Hawaiian Hylaeus species are grouped
within two categories: Ground-nesting
species that require relatively dry
conditions and wood-nesting species
which are found within wetter areas
(Zimmerman 1972, p. 533; Daly and
Magnacca 2003, p. 11).
The female Hylaeus bee lays eggs in
brood cells that she constructs in the
nest and lines with a self-secreted
cellophane-like material. Prior to sealing
the nest, the female provides her young
with a mass of semiliquid nectar and
pollen that is left alongside her eggs.
Upon hatching, the grub-like larvae eat
the provisions left for them, pupate, and
eventually emerge as adults (Michener
2000, p. 24).
The role of bees as pollinators
maintaining communities of native flora
in a diversity of habitats is widely
recognized (Cane and Tepedino 2001, p.
1; Kremen et al. 2007, pp. 302, 307;
National Research Council 2007, p. 13).
Recent studies of visitation records of
Hawaiian Hylaeus bees to native flowers
(Daly and Magnacca 2003, p. 11) and
pollination studies of native plants
(Sakai et al. 1995, pp. 2524–2528; Cox
and Elmqvist 2000, p. 1,238; Sahli et al.
2008, p. 1) have demonstrated that
Hawaiian Hylaeus species almost
exclusively visit native plants to collect
nectar and pollen and, in the process,
pollinate these plants. Hylaeus bees are
very rarely found visiting nonnative
plants for nectar and pollen (Magnacca
2007, pp. 186, 188), and are almost
completely absent from habitats
dominated by nonnative plant species
(Daly and Magnacca 2003, p. 11). Sahli
et al. (2008, p. 1) quantified pollinator
visitation rates to all of the flowering
plant species in communities on a
Hawaiian lava flow dating from 1855 to
understand how pollination webs and
the integration of native and alien
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species changes with elevation. In that
study, eight flowering plants were
observed at six sites, which ranged in
elevation from 880 to 2400 meters (m)
(2,887 to 7874 feet (ft)). The study also
found that the proportion of native
pollinators changed along the elevation
gradient; at least 40 to 50 percent of
visits were from alien pollinators at low
elevation, as opposed to 4 to 20 percent
of visits by alien pollinators at mid to
high elevations. Hylaeus bees were less
abundant at lower elevations, and there
were lower visitation rates of any
pollinators to native plants at lower
elevations, which suggests that Hylaeus
may not be easily replaceable by
nonnative pollinators (Sahli et al., 2008,
p. 1). Because Hylaeus species are likely
critical pollinators of one or more native
Hawaiian plant species, it is believed
that their decline or eventual extinction
may negatively impact dependent native
plant species (Hopper et al. 1996, p. 8;
Cox and Elmqvist 2000, p. 1238).
Taxonomy and Description of Each
Petitioned Hylaeus Species
Unless clearly stated that the
information is from our files, all
information, statements, and references
cited regarding the taxonomy,
descriptions, life history, and range and
distribution are based on information
submitted in the petitions.
Hylaeus anthracinus
Taxonomy
Hylaeus anthracinus was first
described as Prosopis anthracina by
Smith in 1873 (Daly and Magnacca
2003, p. 55), and transferred to
Nesoprosopis 20 years later (Perkins
1899, pp. 75), and then Nesoprosopis
was reduced to a subgenus of Hylaeus
in 1923 (Meade-Waldo 1923, p. 1).
Although the distinctness of this species
is unquestioned, recent genetic evidence
(Magnacca and Brown, submitted)
suggests that H. anthracinus may be
composed of three cryptic (not
recognized) species which represent the
populations on Hawaii; Maui and
Kahoolawe; and Molokai and Oahu.
Description
Hylaeus anthracinus is a mediumsized black bee with clear to smoky
wings and black legs. The male has a
single large yellow spot on his face,
while below the antennal sockets the
face is yellow. The female is entirely
black and can be distinguished by the
black hairs on the end of the abdomen
and an unusual mandible that has three
teeth, a characteristic that is shared only
with H. flavifrons, a closely related
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species on Kauai (Daly and Magnacca
2003, p. 53).
Life History
The diet of the larval stage of Hylaeus
anthracinus is unknown, although the
larvae are presumed to feed on stores of
pollen and nectar collected and
deposited in the nest by the adult
female. Likewise, the nesting habits of
H. anthracinus are not known, but the
species is thought to nest within the
stems of coastal shrubs (Magnacca
2005a, p. 2).
Hylaeus anthracinus adults have been
observed visiting the flowers of
Sesbania tomentosa (oahi), Scaevola
sericea (naupaka kahakai), Sida fallax
(ilima), Argemone glauca (pua kala),
Chamaesyce celastroides (akoko),
Chamaesyce degeneri (akoko),
Heliotropium anomalum (hinahina),
and Myoporum sandwicense (naio).
This species has also been collected
from inside the fruit capsule of Kadua
coriacea (kioele) (Magnacca 2005a, p. 2).
Hylaeus anthracinus has also been
observed visiting Tournefortia argentea
(tree heliotrope), a tree native to tropical
Asia, Madagascar, tropical Australia,
and Polynesia, for nectar and pollen
(Wagner et al. 1999, p. 398; Daly and
Magnacca 2003, p. 55; Magnacca 2007a,
p. 181). The species was first collected
on Oahu in 1864-1865, and is
naturalized and documented from all of
the main islands except Kahoolawe
(Wagner et al. p. 398). It is described as
introduced by Magnacca (2007, p. 181).
Hylaeus anthracinus commonly occurs
alongside other Hylaeus species,
including H. longiceps and H. flavipes.
Range and Distribution
Hylaeus anthracinus was historically
known from numerous coastal strand
and lowland dry forest locations up to
2,000 feet (ft) (610 meters (m)) in
elevation on the islands of Hawaii,
Lanai, Maui, Molokai, and Oahu.
According to the petition, between 1997
and 2008, surveys for Hawaiian Hylaeus
were conducted at 43 sites throughout
the Hawaiian Islands that were either
historic collecting localities for H.
anthracinus, or potentially suitable
habitat for this species. Hylaeus
anthracinus was observed at 14 of the
43 survey sites, but had disappeared
from each of the 9 historically occupied
sites that were surveyed (petition p. 7).
Several of the historical collection sites,
such as Honolulu and Waikiki on Oahu,
and Kealakekua Bay on Hawaii, no
longer contain Hylaeus habitat, which
has been replaced by urban
development or is dominated by
nonnative vegetation (Liebherr and
Polhemus 1997, pp. 346–347; Daly and
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Magnacca 2003, p. 55; Magnacca 2007,
pp. 186–188).
Hylaeus anthracinus is currently
restricted to small populations in a few
small patches of coastal and lowland
dry habitat (Magnacca 2005a, p. 2); one
location on Kahoolawe; five locations
on the island of Hawaii, two locations
on Maui, three locations on Molokai,
and two locations on Oahu (Xerces
2009a, pp. 9-10). The petition does not
define the context applied to the term
‘‘small,’’ and we have no additional
information in our files. Accordingly,
we are presenting the population
information as characterized by the
petitioner. The 2004 H. anthracinus
collection on the island of Hawaii
occurred in montane dry forest
(Magnacca 2005a, p. 2). Although it was
previously unknown from the island of
Kahoolawe, H. anthracinus was
observed at one location on the island
in 2002 (Daly and Magnacca 2003, p.
55). According to the petition, it is
believed to be extirpated from Lanai
(Daly and Magnacca 2003, p. 55).
Hylaeus assimulans
Taxonomy
Hylaeus assimulans was first
described as Nesoprosopis assimulans
(Perkins 1899, pp. 75, 101–102), and
then Nesoprosopis was reduced to a
subgenus of Hylaeus in 1923 (MeadeWaldo 1923, p. 1). The species was most
recently described as Hylaeus
assimulans by Daly and Magnacca in
2003 (pp. 55–56).
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Description
Hylaeus assimulans is distinguished
by its large size relative to other coastal
Hylaeus species and slightly smoky to
smoky-colored wings. The male is black
with yellow face marks, with an almost
entirely yellow clypeus (lower face
region) with additional marks on the
sides that narrow dorsally (towards the
top). The male also has brown
appressed (flattened) hairs on the tip of
the abdomen. The female is entirely
black, large-bodied, and has no distinct
punctuation on the abdomen (Daly and
Magnacca 2003, p. 56).
Life History
The diet of the larval stage of Hylaeus
assimulans is unknown, although the
larvae are presumed to feed on stores of
pollen and nectar collected and
deposited in the nest by the female
adult (Xerces 2009b, p. 11). Likewise,
the nesting habits of H. assimulans are
not known, but the species is thought to
nest underground, as do other closely
related species (Magnacca 2005b, p. 2).
Hylaeus assimulans adults have been
observed visiting the flowers of
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Lipochaeta lobata (nehe) and Sida fallax
(ilima), this species’ likely primary host
plant (Xerces 2009b, p. 10). Hylaeus
assimulans appears to be closely
associated with plants in the genus
Sida, and the petitioner suggests this
yellow-faced bee species may be more
common where this plant is abundant
(Daly and Magnacca 2003, pp. 58, 217;
Magnacca 2007, p. 183). The petition
contains information indicating that in
recent collections, H. assimulans seems
to be more common in dry forest at
relatively higher elevations, and is less
often found in coastal strand habitat.
The petition states that the frequency of
H. assimulans observations in higher,
dry forest may be related to the
abundance of Sida in the understory
(Magnacca 2005b, p. 2). The petitioner
also states that it is likely that H.
assimulans visits several other native
plants, including Acacia koa (koa),
Metrosideros polymorpha (ohia),
Styphelia tameiameiae (pukiawe), and
species of Scaevola (naupaka) and
Chamaesyce (akoko), which are
frequented by other Hylaeus species as
well.
Range and Distribution
Historically, Hylaeus assimulans was
known from numerous coastal strand
and lowland dry locations up to 2,000
ft (610 m) in elevation on the islands of
Lanai, Maui, and Oahu. Although there
are no collections from Molokai, the
petition states that H. assimulans also
occurred there because all other species
of Hylaeus known from Maui, Lanai,
and Oahu also occurred on Molokai
(Xerces 2009b, p. 6). Between 1997 and
2008, surveys for Hawaiian Hylaeus
were conducted in 25 sites on Oahu,
Maui, Kahoolawe, Lanai, and Molokai.
Hylaeus assimulans was absent from six
of its historic localities on Oahu, Maui,
and Lanai (Xerces 2009b, pp. 6-7).
Hylaeus assimulans was not observed at
19 other sites with potentially suitable
habitat on Oahu, Maui, Lanai, and
Molokai, including several sites from
which other native Hylaeus species
have been recently collected (Daly and
Magnacca 2003, p. 56; Xerces 2009b, p.
7).
Currently, Hylaeus assimulans is
found in a few small patches of coastal
and lowland dry habitat at two locations
on Lanai, two locations on Maui, and
one location on Kahoolawe (Daly and
Magnacca 2003, p. 58; Magnacca 2005,
p. 2). According to the petition, this
species has likely been extirpated from
Oahu since it was absent from the
island’s best extant coastal strand
habitat at Kaena Point (Kaena Point
Natural Area Reserve (NAR)) (Magnacca
2005, p. 2).
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Hylaeus facilis
Taxonomy
According to the petitioner, Hylaeus
facilis is a member of the H. difficilis
species group, and is closely related to
H. chlorostictus and H. simplex.
Hylaeus facilis was first described as
Prosopis facilis by Smith in 1879 (Daly
and Magnacca, p. 80), based on a
specimen erroneously reported from
Maui. According to Blackburn and
Cameron (1886 and 1887), the species’
type locality was Pauoa Valley on Oahu
(Daly and Magnacca 2003, p. 80). The
species was later transferred to the
genus Nesoprosopis (Perkins 1899, pp.
75, 77). Nesoprosopis was subsequently
reduced to a subgenus of Hylaeus
(Meade-Waldo 1923, p. 1). The species
was most recently recognized by Daly
and Magnacca (2003, p. 80) as Hylaeus
facilis.
Description
Hylaeus facilis is a medium-sized bee
with smoky colored wings. The male
has an oval yellow mark on its face that
covers the entire clypeus (lower face
region), and a narrow stripe beside the
eyes, but is otherwise unmarked. The
large, externally visible gonoforceps
(paired lateral outer parts of the male
genitalia) distinguish H. facilis from the
closely related H. simplex (Daly and
Magnacca 2003, p. 83). The female is
entirely black, and indistinguishable
from females of H. difficilis and H.
simplex (Daly and Magnacca 2003, pp.
81–82).
Life History
The diet of the larval stage of Hylaeus
facilis is unknown, although the larvae
are presumed to feed on stores of pollen
and nectar collected and deposited in
the nest by the adult female. The nesting
habits of H. facilis have not been
observed, but the species is thought to
nest underground as do the closely
related species H. chlorostictus and H.
simplex (Daly and Magnacca 2003, p.
83; Magnacca 2005c, p. 2).
According to the petition, the native
host plants of adult Hylaeus facilis are
unknown, but it is likely that this
species visits several plants that other
Hylaeus species are known to frequent,
including Acacia koa (koa),
Metrosideros polymorpha (ohia),
Styphelia tameiameiae (pukiawe),
Scaevola spp. (naupaka), and
Chamaesyce spp. (akoko). Hylaeus
facilis has also been observed visiting
the nonnative Tourneforia argentea (tree
heliotrope) for nectar and pollen
(Magnacca 2007, p. 181).
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Range and Distribution
Description
Hylaeus facilis was historically
known from Lanai, Maui, Molokai, and
Oahu, in dry shrubland to wet forest,
from coastal to montane habitat up to
3,281 ft (1,000 m) in elevation (Gagne
and Cuddihy 1999, p. 93; Daly and
Magnacca 2003, pp. 81, 83). Perkins
(1899, p. 77) remarked that H. facilis
was among the most common and
widespread Hylaeus species on Oahu
and all of Maui Nui (Lanai, Molokai,
and Maui) (Magnacca 2007, p. 183). The
petitioner contends that although the
species was widely collected within a
diverse range of habitats, it probably
prefers dry to mesic forest and
shrubland (Magnacca 2005c, p. 2),
which are increasingly rare and patchily
distributed habitats (Smith 1985, pp.
227–233; Juvik and Juvik 1998, p. 124;
Wagner et al. 1999, pp. 66-67, 75;
Magnacca 2005c, p. 2).
The petition states that Hylaeus facilis
has almost entirely disappeared from
most of its historical range (Daly and
Magnacca 2003, p. 7; Magnacca 2007, p.
183), and the abundance of specimens
in the collections at the Bishop Museum
in Honolulu demonstrates the historic
prevalence of this species in a diverse
array of habitats and elevations
(Magnacca 2007, p. 183). Between 1998
and 2006, 39 sites on Oahu, Maui,
Lanai, and Molokai were surveyed; H.
facilis was absent from each of the 13
historical localities that were revisited
(Magnacca 2007, p. 183). Hylaeus facilis
was not observed at 26 other sites with
potentially suitable habitat, including
many sites from which other native
Hylaeus species have been recently
collected (Daly and Magnacca 2003, pp.
7, 81–82; Magnacca 2007, p. 183).
Currently, Hylaeus facilis is only
known from three sites, one each on the
islands of Maui, Molokai, and Oahu
(Daly and Magnacca 2003, pp. 81–82;
Magnacca 2005c, p. 2). According to the
petitioner, this species is likely
extirpated from Lanai (Xerces 2009c, p.
7).
Hylaeus hilaris is distinguished by its
large size (male wing length is 0.185
inches (in) (4.7 millimeters (mm))
relative to other coastal Hylaeus species.
The wings of this species are slightly
smoky to smoky colored, and it is the
most colorful of the Hawaiian Hyaleus
species. The face of the male is almost
entirely yellow, with yellow markings
on the legs and thorax, and the
metasoma (middle portion of the
abdomen) are usually predominantly
red. Females are drably colored, with
various brownish markings. As with
other cleptoparasitic (see ‘‘Life History’’
below) species, H. hilaris lacks the
specialized pollen-sweeping hairs of the
front legs (Daly and Magnacca 2003, pp.
9, 106). It is also one of only two
Hawaiian Hyaleus species to possess
apical (at the end or tip of a structure)
bands of fine white hairs on the
segments of the metasoma.
Hylaeus hilaris
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Taxonomy
Hylaeus hilaris was first described as
Prosopis hilaris by Smith in 1879 (Daly
and Magnacca 2003, pp. 103–104),
transferred to the genus Nesoprosopis
20 years later (Perkins 1899, pp. 75),
and then Nesoprosopis was reduced to
a subgenus of Hylaeus in 1923 (MeadeWaldo 1923, p. 1). In 2003, Daly and
Magnacca described the species as
Hylaeus hilaris (Daly and Magnacca
2003, pp. 103–104).
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Life History
Most adult Hawaiian Hylaeus species
consume nectar for energy; however,
Hylaeus hilaris has yet to be observed
actually feeding from flowers. Hylaeus
hilaris and the four species related to it
(H. hostilis, H. inquilina, H.
sphecodoides, and H. volatilis) are
known as cleptoparasites or cuckoo
bees. The mated female does not
construct a nest or collect pollen, but
instead enters the nest of another
species and lays an egg in a partially
provisioned cell. Upon emerging, the
cleptoparasitic larva kills the host egg
and consumes the provisions, pupates,
and eventually emerges as an adult. As
a result of this lifestyle shift, H. hilaris
bees have lost the pollen-collecting
hairs that other species possess on the
front legs. Cleptoparasitism is actually
quite common among bees:
approximately 25 percent of known bee
species have evolved to become
cleptoparasites. Among the world’s
bees, other than the Hawaiian Hylaeus
group, no cleptoparasites are known
from the family Colletidae (Daly and
Magnacca 2003, p. 9). The larvae of H.
hilaris and their diet are unknown
(Magnacca 2005d, p. 2); however, the
species is known to lay its eggs within
the nests of H. anthracinus, H.
assimulans, and H. longiceps (Perkins
1913, p. lxxxi). Although the species has
never been observed at flowers, H.
hilaris adults presumably consume
nectar as a food source (Xerces 2009 d,
p. 9). Hylaeus hilaris depends on a
number of related Hylaeus host species
for its parasitic larvae, and its
population size is inherently much
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smaller than its host species (Xerces
2009d, p. 9).
Range and Distribution
Hylaeus hilaris was historically
known from coastal strand habitat on
the islands of Lanai, Maui, and Molokai.
The petitioner states that it is believed
to have occurred along much of the
coast of these islands since its primary
hosts, H. anthracinus, H. assimulans,
and H. longiceps, likely extended
throughout this habitat. According to
the petition, nearly all of the coastal
strand habitat has been either developed
or degraded, and is no longer suitable
for H. hilaris (Liebherr and Polhemus
1997, pp. 346–347; Magnacca 2007, pp.
186–188). Hylaeus hilaris was absent
from three of its historical population
sites revisited by researchers between
1998 and 2006. It was also not observed
at 10 additional sites with potentially
suitable habitat where other native
Hylaeus species have been recently
collected (Daly and Magnacca 2003, pp.
103, 106).
The petitioner states that this species
has been collected only twice in the last
70 years, but acknowledges a gap of
about 70 years between major collecting
efforts (Xerces 2009d, p. 6). Hylaeus
hilaris has recently been collected on
two occasions; once in 1989 and again
in 1999. The species was absent from
each of its historical localities that were
revisited between 1998 and 2006
(Xerces 2009d, p. 6). Currently, the only
known population of H. hilaris is
located on The Nature Conservancy’s
Moomomi Preserve on Molokai (Daly
and Magnacca 2003, pp. 103, 106;
Magnacca 2005d, p. 2). According to the
petition, it is no longer extant on Lanai
(Xerces 2009d, p. 6).
Hylaeus kuakea
Taxonomy and Description
Hylaeus kuakea was first described by
Daly and Magnacca (2003, pp. 1, 125–
127) from specimens collected in 1997
in the Waianae Mountains on Oahu.
Hylaeus kuakea is a small, black bee
with slightly smoky colored wings. This
species does not fit into any of the welldefined Hylaeus species groups. Its
facial marks are similar to those of the
H. difficilis group and to H. anthracinus,
but it can be distinguished by its
unusual ivory facial marking covering
the clypeus (the lower face region).
Hylaeus kuakea also resembles H.
anthracinus, but has a denser, more
distinct arrangement of setae (sensory
hairs) on the head and generally
narrower marks next to the compound
eyes (Daly and Magnacca 2003, p. 125;
Magnacca 2005e, p. 2). Only two adult
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Description
male specimens have been collected;
females have yet to be collected or
observed.
Life History
The diet of the larval stage of Hylaeus
kuakea is unknown, although the larvae
are presumed to feed on stores of pollen
and nectar collected and deposited in
the nest by the adult female (Xerces
2009e, p. 7). The nesting habits of H.
kuakea have not been observed, but the
species is believed to be related to other
wood-nesting Hawaiian Hylaeus species
(Magnacca and Danforth 2006, p. 403).
According to information in the
petition, the native host plants of the
adult Hylaeus kuakea are unknown, but
it is likely that this species visits several
plants that other Hylaeus species are
known to frequent, including Acacia
koa, Metrosideros polymorpha,
Styphelia tameiameiae, Scaevola spp.,
and Chamaesyce spp. (Magnacca 2005e,
p. 2).
Range and Distribution
Hylaeus kuakea is only known from
two collections made in Moho Gulch
Ridge, at the northern end of Honouliuli
Preserve, at an elevation of about 1,900
ft (579 m) in the Waianae Mountains on
Oahu. Hylaeus kuakea is found in
lowland mesic forest, which is
increasingly rare and patchily
distributed on Oahu (Smith 1985, pp.
227–233; Juvik and Juvik 1998, p. 124;
Wagner et al. 1999, pp. 66–67, 75).
According to the petitioner, although
there is potentially suitable lowland
mesic habitat in Honouliuli Preserve, no
other individuals of Hylaeus kuakea
were found in surveys subsequent to the
type collection in 1997 (Magnacca 2007,
p. 184). In addition, Perkins did not
collect this species in surveys in the
Honouliuli Preserve vicinity or in
nearby areas in 1899, 1910, and 1911
(Xerces 2009e, p. 6). The petitioner
therefore concludes that the extreme
rarity of this species, its absence from
nearby sites, and the fact that it was not
discovered until very recently suggest
that very few populations remain
(Magnacca 2005e, p. 2).
Hylaeus longiceps
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Taxonomy
Hylaeus longiceps was first described
in 1899 as Nesoprosopis longiceps
(Perkins 1899, pp. 75, 98), and then
Nesoprosopis was reduced to a
subgenus of Hylaeus in 1923 (MeadeWaldo 1923, p. 1). Daly and Magnacca
(2003, pp. 133–134) most recently
described the species as Hylaeus
longiceps.
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Hylaeus longiceps is a small to
medium-sized, black bee with clear to
slightly smoky colored wings. Its
distinguishing characteristics are its
long head and the facial marks of the
male. The lower face of the male is
marked with a yellow band that extends
at the sides of the face in a broad stripe
above the antennal sockets. The area
above the clypeus (lower face region) is
very long and narrow, and the scape
(the first antennal segment) is noticeably
twice as long as it is wide. The female
is entirely black and unmarked (Daly
and Magnacca 2003, p. 133).
Life History
The diet of the larval stage of Hylaeus
longiceps is unknown, although the
larvae are presumed to feed on stores of
pollen and nectar collected and
deposited in the nest by the female
adult (Xerces 2009a, p. 15). The nesting
habits of H. longiceps are unknown, but
the species is thought to nest
underground, as in other closely related
species (Magnacca 2005f, p. 2).
Hylaeus longiceps adults have been
observed visiting the flowers of a wide
variety of plants, including Scaevola
coriacea (dwarf naupaka), Sida fallax,
Scaevola spp. (naupaka kahakai),
Sesbania tomentosa (ohai), Myoposum
sandwicense (naio), Santalum
ellipticum (iliahialoe, coast
sandalwood), Chamaesyce degeneri
(akoko), and Vitex rotundifolia
(pohinahina) (Xerces 2009a, p. 14). The
petitioner reports that it is likely that H.
longiceps visits several plant species
that other Hylaeus species are known to
frequently visit, including Scaevola
spp., Chamaesyce spp., Tournefortia
argentea, Jacquemontia ovata (pau o
hiiaka), and Sida fallax (Magnacca
2005f, p. 2).
Range and Distribution
Hylaeus longiceps is historically
known from numerous coastal strand
and lowland dry shrubland locations up
to 2,000 ft (610 m) in elevation on the
islands of Lanai, Maui, Molokai, and
Oahu. The species is primarily known
from coastal habitat, but is infrequently
collected in dry shrubland. Hylaeus
longiceps is rarely observed in higher
elevation dry forests. Perkins (1899, p.
98) noted that H. longiceps was locally
abundant, and probably occurred
historically throughout much of the
leeward and lowland areas on Maui Nui
(Maui, Molokai, Lanai, and Kahoolawe)
and Oahu, since its host plants, Sida
fallax, Chamaesyce spp., Scaevola spp.,
and Jaquemontia ovata, occurred
throughout these areas (Magnacca 2005f,
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p. 2). The petitioner states that most of
the habitat in these areas has been either
developed or degraded and is no longer
suitable for H. longiceps (Liebherr and
Polhemus 1997, pp. 346–347; Magnacca
2007, pp. 186–188).
Hylaeus longiceps is now restricted to
small populations in small patches of
coastal and lowland habitat on Lanai,
Maui, Molokai, and Oahu (Magnacca
2005f, p. 2). Twenty-five sites that were
either historic collecting localities for H.
longiceps or contained potentially
suitable habitat for this species were
surveyed between 1997 and 2008.
Hylaeus longiceps was observed at only
six of the surveyed sites: three sites on
Lanai, one site on Maui, one site on
Molokai, and one site on Oahu. Only
one historic location, Waieu Dune on
Maui, still supports a population of H.
longiceps (Daly and Magnacca 2003, p.
135).
Hylaeus mana
Taxonomy and Description
Hylaeus mana was first described by
Daly and Magnacca (2003, pp. 135–136)
from four specimens collected in 2002
on the leeward side of the Koolau
Mountains on Oahu. This species is an
extremely small, gracile (gracefully
slender) black bee with yellow markings
on the face. The smallest of all Hawaiian
Hylaeus species, H. mana is a member
of the dumetorum species group. The
face of the male is largely yellow below
the antennae, extending dorsally in a
narrowing stripe. The female’s face has
three yellow lines, one against each eye,
and a transverse stripe at the apex of the
clypeus (lower face region). The
female’s other markings are the same as
the male’s (Daly and Magnacca 2003, p.
135). Hylaeus mana can be
distinguished from H. mimicus and H.
specularis, with whom its range
overlaps, by its extremely small size, the
shape of the male’s genitalia, the
female’s extensive facial marks, and a
transverse rather than longitudinal
clypeal marking (Daly and Magnacca
2003, p. 138).
Life History
The diet of the larval stage of Hylaeus
mana is unknown, although the larvae
are presumed to feed on stores of pollen
and nectar collected and deposited in
the nest by the adult female (Xerces
2009e, p. 7). The nesting habits of H.
mana are not well known, but it is
assumed the species is closely related to
other wood-nesting Hawaiian Hylaeus
species (Magnacca and Danforth 2006,
p. 403).
Adult specimens of Hylaeus mana
were collected while they visited
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flowers of Santalum freycinetianum var.
freycinetianum (iliahi, sandalwood), a
native Hawaiian plant found only on
Oahu and Molokai (Wagner et al. 1999,
p. 1221). The petitioner asserts that it is
likely that H. mana visits several other
native plant species, including Acacia
koa, Metrosideros polymorpha,
Styphelia tameiameiae, Scaevola spp.,
and Chamaesyce spp. (Magnacca 2005g,
p. 2).
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Range and Distribution
Hylaeus mana is only known from
lowland mesic forest located along the
Manana Trail in the Koolau Mountains
on Oahu, at an elevation of about 1,400
ft (427 m). Few Hylaeus bees have been
found in this type of koa-dominated,
lowland mesic forest on Oahu (Daly and
Magnacca 2003, p. 138). This type of
forest is increasingly rare and patchily
distributed on Oahu (Smith 1985, pp.
227–233; Juvik and Juvik 1998, p. 124;
Wagner et al. 1999, pp. 66–67, 75).
According to the petition, because the
first collection of Hylaeus mana was
made in 2002, the historic range and
current distribution, other than the
collection on Manana Trail, are
unknown at this time (Magnacca 2005g,
p. 2). This species was not found in
surveys of potentially suitable habitat in
the same general area by Perkins in
1899, 1910, and 1911 (Xerces 2009e, p.
6). The petitioner therefore concludes
that the extreme rarity of this species, its
absence from nearby sites, and the fact
that it was not discovered until very
recently suggest that very few
populations remain (Magnacca 2005g, p.
2).
We accept the characterization of the
seven species of Hawaiian yellow-faced
bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana) as
described in the information provided
by the petitioner.
Factors Affecting the Species
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
CFR 424, set forth 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; or (E) other natural or
manmade factors affecting its continued
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existence. In making this 90–day
finding, we evaluated whether
information on threats to the seven
Hawaiian Hylaeus bee species presented
in the petitions and available in our files
at the time of the review of the petitions
constitute substantial scientific or
commercial information such that
listing the species may be warranted.
Our evaluation of this information is
discussed below. Unless clearly stated
that the information is from our files, all
threats described below and their effects
on the seven Hawaiian Hylaeus bee
species are based on information
submitted in the petitions. Any
references provided in support of
particular statements related to potential
threats are the petitioner’s references,
unless specifically identified otherwise.
A. Present or Threatened Destruction,
Modification, or Curtailment of the
Species’ Habitat or Range
According to the petitions,
degradation and loss of coastal and
lowland habitat used by Hylaeus bees
on all of the main Hawaiian Islands is
the primary threat to these seven species
(Cuddihy and Stone 1990, pp. 60–61;
Daly and Magnacca 2003, pp. 55, 173).
Coastal and lowland habitats have been
severely altered and degraded, partly
because of past and present land
management practices, including
agriculture, grazing, and urban
development; the deliberate and
accidental introductions of nonnative
animals and plants; and recreational
activities. In addition, the petitions
present information indicating that fire
is a potential threat to the habitat of
these seven species in some locations.
Habitat Destruction and Modification by
Urbanization and Land Use Conversion
Increased access to coastal areas, and
resulting habitat disturbance, has been
facilitated by coastal development and
roadbuilding (Cuddihy and Stone 1990,
pp. 94–95). As described in the
petitions, five species (Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, and H. longiceps) were once
widespread and common in coastal
strand habitat (Perkins 1912, p. 688).
These five Hylaeus species are now
absent from all of Perkins’ coastal
collection localities. Hylaeus facilis has
recently been collected in coastal
habitat at Kuololimu Point, and H.
hilaris has recently been collected in
coastal habitat at Moomomi Preserve,
Molokai (Xerces 2009c, p. 9).
The petitioner states that lowland dry
forests and shrublands have been
heavily impacted by urbanization and
conversion to agriculture or pasture
throughout the Hawaiian Islands, with
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the estimated loss of more than 90
percent of dry forests and shrublands
(Bruegmann 1996, p. 26; Juvik and Juvik
1998, p. 124). Four species (Hylaeus
anthracinus, H. assimulans, H. facilis,
and H. longiceps) were once widespread
and found within lowland dry habitat
on several islands, including Hawaii,
Lanai, Maui, Molokai, and Oahu, but are
largely absent from their historical
population sites on these islands.
Mesic forest, once abundant and
considered the most diverse of all
Hawaiian forest types (Rock 1913, p. 9),
is now very rare, with much of it
converted to pasture, or military or
agricultural use, or lost to urbanization
(Cuddihy and Stone 1990, p. 61;
Magnacca 2007, p. 187). Fire has also
negatively impacted this habitat type, as
is discussed below. Hylaeus facilis was
historically the most wide ranging of the
seven bee species in terms of the variety
of habitats from which it was recorded,
which included mesic forest on Lanai,
Maui, Molokai, and Oahu. This species
is now restricted to single locations on
the islands of Molokai and Oahu.
The petitioner identified the loss of
coastal, dry lowland, and montane wet
forest habitat on Oahu, Lanai, Maui, and
Molokai as a contributing factor to the
decline of H. facilis, but acknowledges
that ‘‘although recorded from several
sites currently considered to be wet
forest, it is possible that H. facilis would
not normally inhabit this [habitat] in a
natural state.’’ The petitioner attributes
the current observation of this species at
sites now known to be wetter than they
were during the early Perkins’ collecting
period to the more open understory
vegetation (Perkins 1899, p. 76). It is
conceivable that the loss of mesic forest
habitat used by H. kuakea is due to
urbanization and land use conversion,
although the petitioner presents no
information in this regard, nor do we
have information in our files regarding
the historical locations of these two
species, both of which were only
recently collected (H. kuakea in 1997;
H. mana in 2002).
Habitat Destruction and Modification by
Nonnative Plants
The petitioner states that the spread of
nonnative plant species is one of the
primary causes of decline, and a current
threat to the existing populations of
each of the seven Hylaeus bee species,
because they depend closely on native
vegetation for nectar and pollen, and the
bees are almost entirely absent from
habitat dominated by invasive,
nonnative vegetation (Sakai et al. 2002,
pp. 276, 291; Daly and Magnacca 2003,
p. 11; Liebherr 2005, p. 186). According
to information available in our files and
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presented by the petitioner, the native
flora within a majority of lowland
habitat on the Hawaiian Islands below
1,969 ft (600 m) is being replaced by
aggressive, nonnative plant species
(Cuddihy and Stone 1990, pp. 73–74;
Wagner et al. 1999, p. 52). The
petitioner states that many native plant
species that are replaced by nonnative
plants were once foraging resources for
numerous Hylaeus species (Cox and
Elmqvist 2000, p. 1238; Daly and
Magnacca 2003, p. 11; USFWS 1999, pp.
145, 163, 171, 180; USFWS 2008, pp. 7,
9). Six of the seven Hylaeus bee species
(Hylaeus anthracinus, H. assimulans, H.
facilis, H. kuakea, H. longiceps, and H.
mana) are most often found in dry and
mesic forest and shrubland habitat (Daly
and Magnacca 2003, p. 11), and the
greatest proportion of endangered or atrisk Hawaiian plant species are also
limited to these same habitats; 25
percent of Hawaiian listed plant species
are from dry forest and shrubland alone
(Sakai et al. 2002, pp. 276, 291, 292).
The petitioner asserts that lowland dry
and mesic forests now support lessdiverse Hylaeus communities because
many native plants used for foraging are
extirpated from these habitats
(Magnacca 2007, pp. 186–187).
The petitioner states that besides
Scaevola sericea (naupaka kahakai),
native vegetation is lacking along most
of the coastline of the major Hawaiian
Islands, and that Hylaeus bees cannot
survive on this plant alone (Magnacca
2007, p. 187). The petitioner also states
that native coastal vegetation in many
areas, such as Moomomi Preserve on
Molokai, which currently is the only
known location for Hylaeus hilaris, is
threatened by Prosopis pallida (kiawe),
an invasive, nonnative, deciduous
thorny tree (Xerces 2009a, p. 25; 2009b,
p. 17; 2009c, p. 21; 2009d, p. 11).
According to the petitions, many of
the native plants that serve as foraging
resources for the adults of the seven
Hylaeus bee species are declining due to
a lack of pollinators (Daly and Magnacca
2003, p. 11; USFWS 2008, pp. 7, 9) and
are found only in very small
populations (USFWS 1999, pp. 145,
163, 171, 180; Cox and Elmqvist 2000,
p. 1238). The petitioner points out, for
example, that H. longiceps and H.
anthracinus are known to forage on the
federally endangered plant Sesbania
tomentosa (ohai). Both H. longiceps and
H. anthracinus also visit Chamaesyce
celastroides var. kaenana (akoko), a
federally endangered plant endemic to
coastal dry shrubland on Oahu (Daly
and Magnacca 2003, pp. 55, 74).
Hylaeus longiceps is also known to
forage on the endangered Scaevola
coriacea (dwarf naupaka) (USFWS 1999,
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p. 145; Daly and Magnacca 2003, pp. 55,
135). In addition, H. anthracinus has
been collected from inside the fruit
capsule of Kadua coriacea (kioele), a
federally endangered dry forest plant,
known from fewer than 300 individuals
on the island of Hawaii (USFWS 2008,
p. 5; Christian Torres, USFWS, pers.
comm. 2009).
Habitat Destruction and Modification by
Nonnative Ungulates
The petitioner claims that the decline
of native plant communities has likely
had a negative impact on Hawaii’s
Hylaeus bee species (Cuddihy and Stone
1990, pp. 59–66, 88–94, 73–76; USFWS
2006, p. 2684). The presence of
nonnative mammals, such as feral pigs
(Sus scrofa), cattle (Bos taurus), goats
(Capra hircus), and axis deer (Axis
axis), is considered one of the primary
factors underlying the alteration and
degradation of native vegetation and
habitat in the Hawaiian Islands (Stone
1985, pp. 262–263; Cuddihy and Stone
1990, pp. 60–66; 73 FR 73801). Beyond
the direct effects of trampling and
consuming native plants, nonnative
ungulates contribute significantly to
increased erosion, and their behavior
(i.e., rooting, moving across large
expanses) facilitates the spread and
establishment of competing, invasive,
nonnative plant species (Xerces 2009a,
p. 26; 2009b, p. 18; 2009c, pp. 21–22,
2009d, pp. 12–13, 2009e, p. 10). Several
endangered coastal and lowland plant
species that are threatened by the
browsing, trampling, and digging
activities of nonnative ungulates are
confirmed foraging sources for Hylaeus
species and, therefore, are likely
foraging sources for these seven Hylaeus
species (USFWS 1999, pp. 145, 163,
171, 180; Daly and Magnacca 2003, pp.
11, 13).
Habitat Destruction and Modification by
Fire
The petitions state that fire can
dramatically alter the species
composition of plant communities in
coastal and lowland habitats (Hughes et
al. 1991, p. 743; Blackmore and
Vitousek 2000, p. 625), and thus
potentially impact Hylaeus populations.
The petitioner also suggests that
ordnance-induced fires on the Army’s
Pohakuloa Training Area on the island
of Hawaii may threaten the dry forest
habitat of Hylaeus anthracinus. Fires
were uncommon in the Hawaiian
Islands until the arrival of humans
about 2,000 years ago (Smith and
Tunison 1992, pp. 394–395). Native
habitat in the Hawaiian Islands has been
increasingly colonized by fire-adapted
invasive plant species that take the
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place of, and permanently replace,
native plant species (Cuddihy and
Stone, pp. 88–94; Smith and Tunison
1992, pp. 394–395; D’Antonio et al.
2000, pp. 73–74). This process has been
facilitated by nonnative ungulates,
which alter the floral composition of
native habitats, making conditions more
conducive to fire. This impact occurs
because of the browsing and trampling
of native vegetation, and the spreading
of seeds of nonnative, fire-adapted plant
species such as Melinis minutiflora
(molasses grass) and Schizachyrium
condensatum (tufted beardgrass)
(D’Antonio et al. 2000, pp. 73–74).
Habitat Destruction and Modification by
Recreational Activities
The petitions state that some of the
best habitat areas for Hylaeus species
are also popular recreational sites,
particularly those areas located within
coastal strand habitat (Xerces 2009a, p.
27; 2009c, p. 22). Human impacts at
recreational sites may include removal
or trampling of vegetation on or near
trails and the compaction of vegetation
by off-road vehicles (Xerces 2009a, p.
27; 2009c, p. 22). In particular, the
petitioner claims that Hylaeus facilis
habitat may be threatened by
recreational activities, such as hunting
and hiking on the Poamoho Trail on
Oahu (Hawaii Department of Land and
Natural Resources 2000, p. 15; Xerces
2009c, p. 22). According to the
petitions, some of the best remaining
habitat for H. anthracinus and H.
longiceps includes Kaena Point (on
Oahu), Kona Coast State Park,
Makalawena, Mokuauia, and South
Point (on the island of Hawaii), areas
that are popular recreational sites with
largely unregulated access (Xerces
2009a, p. 27).
Habitat Destruction and Modification by
Climate Change
The petitioner asserts that a changing
climate may cause shifts in the range of
Hylaeus host plant species, which can
be especially detrimental to dependent
pollinators like these seven species
when combined with habitat loss
(National Research Council 2007, p.
102). Most bees have difficulty crossing
large geographical barriers (Michener
2000, p. 103), and successive
generations of solitary species like
Hylaeus tend to nest in the same area
year after year. The petitioner points out
that the seven Hylaeus bee species are
restricted to habitat patches where
native host plant species are present,
and argues that they are not likely to
disperse far to find new habitat (Xerces
2009a, p. 30; 2009b, p. 21; 2009c, p. 25;
2009d, p. 14; 2009e, p. 13). Thus, the
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ecology of these seven species,
combined with the patchy distribution
of their remaining habitat, may hinder
their dispersal if relocation becomes
necessary due to climate-influenced
changes in distribution of host plant
species (Magnacca 2007, pp. 173, 181–
183, 188) and cause the extirpation of
remaining populations of Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, H. kuakea, H. longiceps, and
H. mana.
The petitioner states that climate
change may also have a deleterious
effect upon the seven Hylaeus bee
species due to climate-induced changes
in rainfall patterns, since these species
prefer relatively dry habitats, some of
which lack groundwater sources. The
petitioner presents a concern that a
predicted rise in sea level in the
Hawaiian Islands (Baker et al. 2006, p.
1) might threaten coastal strand
populations of the seven Hylaeus bee
species. The petitions cite one study
that predicted that sea level rise in the
Northwestern Hawaiian Islands will
cause a median projected loss of land of
3 to 65 percent with a 19-in (48-cm) sea
level rise, and a maximum projected
loss of land of 5 to 75 percent with a 35in (88-cm) sea level rise (Baker et al.
2006, p. 1). Although none of the seven
Hylaeus bees occurs on the
Northwestern Hawaiian Islands, the
petitioner concludes that sea level rise
will also impact the populations of the
five species of Hawaiian yellow-faced
bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H.
longiceps) inhabiting coastal sites on the
main Hawaiian Islands.
Summary of Factor A
In summary, we find that the
information provided in the petitions
presents substantial scientific or
commercial information indicating that
the petitioned actions may be warranted
due to the present or threatened
destruction, modification, or
curtailment of the species’ habitat or
range. The petitioner has provided no
information, and we have no
information in our files to substantiate
the claim that there will be climateinduced changes in rainfall patterns in
the areas where the seven species occur,
or that relatively dry habitats will be
negatively impacted. The petitions did
identify numerous potential factors that
may be affecting Hylaeus anthracinus,
H. assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana,
including habitat loss and degradation
due to urbanization and land
conversion; replacement of native host
plants by nonnative plants caused by
the browsing, trampling, and rooting
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activities of nonnative ungulates, which
facilitates the establishment of
nonnative plants in disturbed areas;
conversion by fire of native plant
communities to plant communities
dominated by nonnative, fire-adapted
plants; and the removal or trampling of
native vegetation by people and
compaction of native vegetation by offroad vehicles in popular recreational
areas, particularly in coastal strand
habitat. Information in our files also
indicates these factors may present a
threat to the seven species of Hylaeus.
We, therefore, conclude the petitions
present substantial information to
indicate that the present or threatened
destruction or modification of habitat
may present a threat to H. anthracinus,
H. assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana.
B. Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
According to the petitioner, Hylaeus
facilis, H. hilaris, H. kuakea, and H.
mana, each with two or fewer known
populations, are especially vulnerable to
overcollection because the collection of
even a few individuals could
significantly reduce the production of
offspring (Xerces 2009c, p. 23; 2009d, p.
13; 2009e, p. 11). However, the petitions
also acknowledge that because of the
high fecundity of individual insects, the
collection of insects does not pose a
threat to their populations (Xerces
2009c, p. 23; 2009d, p. 11; 2009e, p. 11),
which introduces an element of
uncertainty concerning this claim.
Insect collecting is a valuable
component of research, including
taxonomic work, and is often necessary
for documenting the existence of
populations and population trends. The
petitioner has not presented information
with which we can evaluate whether the
overcollection of Hylaeus facilis, H.
hilaris, H. kuakea, or H. mana may
present a threat to any of these species,
or determine whether this activity has
resulted in population declines. In this
regard, neither the petitions, nor
information available in our files,
presents information that would
indicate overcollection may present a
significant threat to H. anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, or H. mana.
C. Disease or predation
Neither the petitions nor information
in our files presents information that
would indicate disease is a current
threat to Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, or H. mana.
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Predation by Nonnative Ants
Ants are known to prey upon Hylaeus
species (Medeiros et al. 1986, pp. 45–46;
Reimer 1994, p. 17), thereby directly
eliminating them from specific areas.
The petitions state that ants are not a
natural component of Hawaii’s
arthropod fauna, and the native species
of the islands evolved in the absence of
predation pressure from ants. They also
state that ants can be particularly
destructive predators because of their
high densities, recruitment behavior,
aggressiveness, and broad range of diet
(Reimer 1993, pp. 17–18). The petitions
also state that the threat of ant predation
on the seven Hylaeus bee species is
amplified by the fact that most ant
species have winged reproductive
adults (Borror et al. 1989, p. 738) and
can quickly establish new colonies in
suitable habitats (Staples and Cowie
2001, p. 55). In addition, the petitions
state that these attributes allow some
ants to destroy otherwise geographically
isolated populations of native
arthropods (Nafus 1993, pp. 19, 22–23).
According to the petitions, at least 47
species of ants are known to be
established in the Hawaiian Islands
(Hawaii Ants 2008, pp. 1–11). Native
insect fauna, likely including Hylaeus
bees (Zimmerman 1948, p. 173; Reimer
et al. 1990, pp. 40–43; HEAR database
2005, pp. 1–2), have been severely
impacted by at least four particularly
aggressive ant species: The big-headed
ant (Pheidole megacephala), the longlegged ant (also known as the yellow
crazy ant) (Anoplolepis gracilipes),
Solenopsis papuana (no common
name), and Solenopsis geminata (no
common name). The petitions state that
numerous other species of ants are
recognized as threats to Hawaii’s native
invertebrates, and an unknown number
of new species of ants are established
every few years (Staples and Cowie
2001, p. 53). The petitions state that due
to their preference for drier habitat sites,
ants are more likely to occur in high
densities in the dry and mesic habitat
currently occupied by the seven bees
(Xerces 2009a, p. 28; 2009b, p. 19;
2009c, p. 24; 2009d, pp. 13–14; 2009e,
pp. 11–12).
The petitions state that the longlegged ant appeared in Hawaii in 1952;
now occurs on Kauai, Oahu, Maui, and
Hawaii (Reimer et al. 1990, p. 42); and
inhabits low-to-mid-elevation (less than
2,000-ft (600-m)) rocky areas of
moderate rainfall (less than 100 in (250
cm) annually) (Reimer et al. 1990, p.
42). The petitioner also states that direct
observations indicate that Hawaiian
arthropods are susceptible to predation
by this species; Gillespie and Reimer
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(1993, p. 21) and Hardy (1979, p. 34)
documented the impacts to native
insects within the Kipahulu area on
Maui after this area was invaded by the
long-legged ant. The petitioner
concludes that although only cursory
observations exist, long-legged ants are
thought to be a threat to populations of
the seven Hylaeus bees within dry to
mesic areas within their elevation range
(Reimer et al. 1990, p. 42).
Solenopsis papuana is the only
abundant, aggressive ant that has
invaded intact mesic to wet forest from
sea level to over 2,000 ft (600 m) on all
of the main Hawaiian Islands, and is
still expanding its range (Reimer 1993,
p. 14). The petitions state that because
of this species’ expanding range, it may
threaten populations of Hylaeus facilis
in mesic areas up to 2,000 ft (600 m) in
elevation (Reimer 1993, p. 14).
The petitions state that the presence
of ants in nearly all of the low-elevation
habitat sites historically and currently
occupied by the seven Hylaeus bee
species may increase the uncertainty of
Hylaeus recovery within these areas.
Hylaeus populations are known to be
drastically reduced in ant-infested areas
(Medeiros et al. 1986, pp. 45–46; Stone
and Loope 1987, p. 251; Cole et al. 1992,
pp. 1313, 1317, 1320; Reimer 1994, p.
17). Although ant species’ primary
impact on the native invertebrate fauna
is via predation (Reimer 1994, p. 17),
they also compete for nectar (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
impact Hylaeus species indirectly as
well, by predating on seeds of native
plants (Bond and Slingsby 1984, p.
1031). The petitioner suggests that the
greatest ecosystem-level effect of
invasive ants has been on pollination.
Additionally, where ranges overlap, ants
compete with native pollinators such as
Hylaeus species and preclude them
from pollinating native plants. For
example, the big-headed ant is known to
actively rob nectar from flowers without
pollinating them (Howarth 1985, p.
157). Lach (2008, p. 155) found that
Hylaeus species that regularly collect
pollen from flowers of Metrosideros
polymorpha were entirely absent from
trees that had their flowers exposed to
big-headed ant foraging.
The Hylaeus egg, larvae, and pupal
stages are more vulnerable to attack by
ants than the mobile adult bee (Daly and
Magnacca 2003, p. 10). Invasive ants
have severely impacted ground-nesting
Hylaeus species in particular (Cole et al.
1992, pp. 1317, 1320; Medeiros et al.
1986, pp. 45–46), because their nests are
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easily accessible, and in or near the
ground. Since Hylaeus anthracinus, H.
facilis, H. hilaris, and H. longiceps are
related to other ground-nesting Hylaeus
species, they may also be susceptible to
ant predation (Magnacca 2005g, p. 2).
The rarity or disappearance of native
Hylaeus species, including the seven
petitioned Hawaiian yellow-faced bee
species, from historically documented
localities over the past 100 years is
likely due to a variety of factors. There
is no information that conclusively
correlates the decrease in Hylaeus
observations with the establishment of
nonnative ants in low-to-montane and
dry-to-wet habitats on the Hawaiian
Islands; however, their collective
presence suggests that nonnative ants
may have played a role in the decline
of some populations of the seven
Hylaeus bee species evaluated in this
finding.
Predation by Nonnative Western
Yellowjacket Wasps
The petitioner suggests that Vespula
pensylvanica (the western yellowjacket
wasp) is a potentially serious threat to
the seven Hylaeus bees. This assertion
is supported by literature available in
our files (Gambino et al. 1987, p. 170;
Wilson et al. 2009, pp. 1–5). The
western yellowjacket wasp is a social
wasp species native to the mainland of
North America. It was first reported
from 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 yellowjacket wasp has an
annual life cycle, but in Hawaii’s
tropical climate, colonies of this species
persist through a second year, allowing
them to have larger numbers of
individuals (Gambino et al. 1987, p.
170) and thus a greater impact on prey
populations. Most colonies are found
between 1,969 and 3,445 ft (600 and
1,050 m) in elevation (Gambino et al.
1990, p. 1,088), although they can also
occur at sea level. The western
yellowjacket wasp is known to be an
aggressive, generalist predator (Gambino
et al. 1987, p. 170), and has been
documented preying upon Hawaiian
Hylaeus species (Wilson et al. 2009, p.
2). The petitioner argues that predation
by the western yellowjacket wasp is a
potentially significant threat to Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, H. kuakea, H. longiceps, and
H. mana because of their small
population sizes. This may present a
particular threat to H. facilis, H. hilaris,
H. kuakea, and H. mana because each
species has two or fewer populations.
The petitions also suggest that the
western yellowjacket wasp may
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compete for nectar with Hylaeus
species, but the petitions provide no
information indicating that competition
for nectar is a threat.
Predation by Nonnative Parasitoid
Wasps
The petitions state that native and
nonnative parasitoid wasps parasitize
some Oahu Hylaeus species and may
pose a threat to H. kuakea and H. mana
(Xerces 2009e, p. 12). The petitions also
state that Hylaeus larvae are known to
be attacked by parasitoid wasps within
the Encyrtidae and Eupelmidae families,
although it is unconfirmed whether
parasitoid wasps utilize H. kuakea and
H. mana as nutritional hosts for their
larvae (Xerces 2009e, p. 12). However,
the petitions did not provide any
evidence, and we have nothing in our
files, to support these claims.
Summary of Factor C
Overall, the petitions provided
substantial scientific or commercial
information indicating that the
petitioned actions may be warranted
due to disease or predation. Neither the
petitions, nor information available in
our files, present data that would
indicate that predation by parasitoid
wasps presents a threat to any of the
Hylaeus species addressed in this
finding. Although the petitions suggest
that the western yellowjacket wasp may
compete for nectar with Hylaeus
species, no information was presented
that would allow us to evaluate whether
this presents a significant threat to any
of the petitioned species. However,
observations and reports have
documented that ants are particularly
destructive predators because of their
high densities, broad range of diet, and
ability to establish new colonies in
otherwise geographically isolated
locations, because the reproductive
adults are able to fly (Xerces 2009a, pp.
27–28; 2009b, pp. 19–20; 2009c, p. 23;
2009d, pp. 13–14, 2009e, p. 11). In
addition, the western yellowjacket wasp
has been documented to prey upon
Hawaiian Hylaeus species (Xerces
2009a, p. 29; 2009b, p 20; 2009c, p. 24;
2009d, pp. 14–15, 2009e, pp. 12–13).
Accordingly, we conclude the petitions
present substantial information
indicating that Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana may
be threatened because of predation by
nonnative ants and the nonnative
western yellowjacket wasp.
D. The Inadequacy of Existing
Regulatory Mechanisms
The petitioner stated that there are no
existing Federal, State, or local laws,
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treaties, or regulations that specifically
conserve or protect habitat for the seven
Hylaeus bee species. The petitioner does
acknowledge that some historic and
current collection localities are
protected from development by Federal
or State agencies; one of two known
populations of H. facilis occurs at
Kalaupapa National Historical Park on
Molokai, and three species (H.
anthracinus, H. hilaris, and H.
longiceps) occur at the State’s Kaena
Point Natural Area Reserve (NAR) on
Oahu, Kanaio NAR on Maui, or The
Nature Conservancy’s Moomomi
Preserve on Molokai. The petitioner
asserts that conservation of the seven
Hylaeus bees will likely require active
management of their known population
sites, involving exclusion and removal
of feral ungulates, control and removal
of nonnative plant and insect species,
and the restoration of native vegetation
(Magnacca 2007, p. 185). The petitions
state that existing regulatory
mechanisms are inadequate to provide
the necessary active management
needed to protect the seven Hylaeus
species (Xerces 2000a, p. 29; 2000b, p.
20; 2000c, p 24; 2000d, p. 15, 2000e, p.
13). However, there was no specific
information provided in the petitions
about existing regulatory mechanisms
that could protect these species. We are
also not aware of any regulatory
mechanisms that address the seven
Hylaeus species.
The petitioners claim that there are no
protections provided by existing State or
Federal regulations to effectively
address potential threats to the seven
species of Hawaiian yellow-faced bees
(Xerces 2000a, p. 29; 2000b, p. 20;
2000c, p 24; 2000d, p. 15; 2000e, p. 13).
However, the petitioners did not
provide any additional information
about existing regulatory mechanisms
that could protect these species, and we
have nothing in our files that describes
any regulatory mechanisms that address
the seven Hylaeus species. While
information presented by the petitioner
indicates that threats to the petitioned
species may be posed by habitat
destruction and degradation by
nonnative ungulates and nonnative
plants and through predation by
nonnative insects, none of these threats
are posed by an inadequacy of
regulatory mechanisms. We, therefore,
find that the petitions do not present
substantial information indicating that
the inadequacy of existing regulatory
mechanisms may present a threat to
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, or H. mana. However, we
will further evaluate the adequacy of
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existing regulatory mechanisms for
protecting the seven species of
Hawaiian yellow-faced bees and their
habitats during our status review.
E. Other Natural or Manmade Factors
Affecting the Species’ Continued
Existence
Small Number of Populations and
Individuals
Species that are endemic to single
islands or known from few, widely
dispersed locations are inherently more
vulnerable to extinction than
widespread species because of the
higher risks from genetic bottlenecks,
random demographic fluctuations,
climate change, and localized
catastrophes such as hurricanes,
landslides, and drought (Lande 1988, p.
1455; Mangel and Tier 1994, p. 607;
Pimm et al. 1988, p. 757). These
problems can be further magnified when
populations are few and restricted to a
limited geographic area, and the number
of individuals is very small. Populations
with these characteristics face an
increased likelihood of stochastic
extinction due to changes in
demography, the environment, genetics,
or other factors, in a process described
as an extinction vortex (Gilpin and
´
Soule 1986, pp. 24–25). Small, isolated
populations often exhibit a reduced
level of genetic variability or genetic
depression due to inbreeding, which
diminishes the species’ capacity to
adapt and respond to environmental
changes, thereby lessening the
probability of long-term persistence
´
(Frankham 2003, pp. S22–S29; Soule
1986, pp. 31–34). The negative impacts
associated with small population size
and vulnerability to random
demographic fluctuations or natural
catastrophes can be further magnified by
synergistic interactions with other
threats.
The petitioner states that all of the
petitioned Hylaeus bee species are rare,
have very small populations, and are
likely more vulnerable to habitat change
and stochastic events due to low genetic
variability (Daly and Magnacca 2003, p.
3; Magnacca 2007, p. 173, Petition p.
13). Literature cited by the petitioner
states that about three-quarters of the
species described from the Hawaiian
Islands by Perkins (1899, 1910, 1911)
have been collected recently. Some are
still as rare or as abundant as he
observed, yet others, formerly abundant,
have not been collected recently (Daly
and Magnacca 2003, p. 3). Five species
have not been collected recently from
one or more islands from which they are
historically known, 7 are restricted to
endangered habitat, 10 are considered to
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be very rare and potentially endangered,
and 10 have not been collected recently
and could be extinct (Magnacca 2007, p.
3). The petitioner asserts that Hylaeus
facilis, H. hilaris, H. kuakea, and H.
mana have not been recently observed
at some historical collection sites, and
that each of these species now has two
or fewer known populations, which
could increase the risk of extinction due
to stochastic events such as hurricanes,
landslides, large wildfires, or prolonged
drought (Jones et al. 1984, p. 209; Smith
and Tunison 1992, p. 398; Petition p.
13). Since H. hilaris is cleptoparasitic
and restricted to one known remaining
population, it is at particularly high risk
of extinction because of the rarity of its
hosts and the fact that it is the most
habitat-specific of all Hawaiian bees
(Daly and Magnacca 2003). The
recurrence intervals for stochastic
events of this nature have not been
explicitly defined, which introduces
some uncertainty regarding potential
effects to the petitioned species. The
fact that a species is potentially
vulnerable to stochastic processes does
not necessarily mean that it is
reasonably likely to experience, or have
its status affected by, a given stochastic
process within timescales that are
meaningful under the Act.
While we recognize the inherent
species risks of small population size
and small numbers of individuals, we
currently lack information needed to
assess this potential threat to the status
of the petitioned species. We will
investigate issues related to Hylaeus
population size and species
susceptibility to catastrophic stochastic
events during the status review in order
to better address this concern in the 12–
month finding.
Competition with Nonnative Insects
There are 15 known species of
nonnative bees in Hawaii (Snelling
2003, p. 342), including two nonnative
Hylaeus species (Magnacca 2007, p.
188). According to the petitioner, most
nonnative bees inhabit areas dominated
by nonnative vegetation and do not
compete with native Hawaiian bees
(Daly and Magnacca 2003, p. 13). 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 tree line (Howarth 1985, p.
156). The petitioner reports that
European honey bees have been
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observed foraging on Hylaeus host
plants such as Scaevola spp. and
Sesbania tomentosa. However, the
petitioner does not present information
indicating that Hawaiian Hylaeus
populations have declined because of
competition with European honey bees
for nectar and pollen (Magnacca 2007,
p. 188). The petitioner asserts that
populations of the European honey bee
are not as vulnerable to predation by
nonnative ant species as are Hylaeus
bees (see Factor C above). The petitioner
refers to a study by Lach (2008, p. 155),
who observed that although Hylaeus
bees that regularly collect pollen from
the flowers of Metrosideros polymorpha
trees were entirely absent from trees
whose flowers had been visited by the
big-headed ant, visits by the European
honey bee were not affected by bigheaded ant presence.
As described by the petitioner, other
nonnative bees found in areas of native
vegetation include Ceratina species
(carpenter bees), Hylaeus albonitens
(Australian colletid bees), and
Lasioglossum impavidum (no common
name) (Magnacca 2007, p. 188). The
petitioner suggests that these nonnative
bees may impact native Hylaeus bees
such as H. facilis through competition
for pollen, based on their similar size
and flower preferences. However, the
petitioner acknowledges that the impact
of these species on native Hylaeus bees
has not been studied (Magnacca 2007, p.
188). The petitioner also suggests that
parasitoid wasps may compete for
nectar with native Hylaeus species (Daly
and Magnacca 2003, p. 10), but did not
present supporting information in this
regard. No information on the potential
threat to the species from parasitoid
wasps is available in our files.
Summary of Factor E
In summary, the petitions provided
substantial scientific or commercial
information indicating that the
petitioned actions may be warranted
due to other factors affecting the
species’ continued existence. The
petitioner did not present information,
nor is information available in our files,
indicating that competition from
parasitoid wasps or other nonnative
bees, such as Ceratina species, Hylaeus
albonitens, and Lasioglossum
impavidum, presents a threat to the
petitioned species. However, the
petitions do present information
indicating that Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana may
be threatened because of their very
small populations and low genetic
variability, which may make them
vulnerable to habitat change and
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stochastic events such as droughts. Each
of the petitions characterizes the
population status of the petitioned
species as ‘‘small and isolated’’ or
‘‘extremely rare, very small
populations,’’ and we do not have any
contrary information in our files. The
petitioner also presents information
indicating that competition with the
European honey bee may present a
threat to the seven Hylaeus bee species.
We, therefore, conclude that the petition
presents substantial scientific
information indicating that other natural
or manmade factors affecting the
species’ continued existence may
threaten Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana.
These factors include the species’ small
numbers of populations and individuals
and competition with nonnative
European honey bees.
Finding
We have reviewed the petitions,
supporting information provided by the
petitioner, and information in our files,
and we evaluated that information to
determine whether the sources cited
support the claims made in the
petitions. On the basis of our evaluation
of the petition under section 4(b)(3)(A)
of the Act, we have determined that the
petition presents substantial scientific
or commercial information indicating
that listing the seven Hylaeus bees as
threatened or endangered may be
warranted. This finding is based on
information that indicates these species’
continued existence may be affected by
destruction or modification of their
coastal strand and lowland forest and
shrubland habitat from urbanization and
land conversion, nonnative plants,
nonnative ungulates, fire, recreational
activities (Factor A); predation by
nonnative ants and the western
yellowjacket wasp (Factor C);
inadequate protection from threats by
existing regulatory mechanisms (Factor
D); and other natural or manmade
factors such as small population size,
and competition with the European
honey bee (Factor E). The petitioner
does not present substantial information
that these seven Hylaeus bees are
threatened by overcollection (Factor B)
currently or in the future.
Because we have found that the
petition presents substantial
information indicating that listing the
seven Hylaeus bee species may be
warranted, we are initiating status
reviews to determine whether listing
these seven species under the Act is
warranted. At the conclusion of the
status reviews we will issue 12–month
findings, in accordance with section
PO 00000
Frm 00027
Fmt 4702
Sfmt 4702
4(b)(3)(B) of the Act, as to whether or
not the Service believes a proposal to
list Hylaeus anthracinus, H. assimulans,
H. facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana is warranted.
The ‘‘substantial information’’
standard for a 90–day finding differs
from the Act’s ‘‘best scientific and
commercial data’’ standard that applies
to a status review to determine whether
a petitioned action is warranted. A 90–
day finding does not constitute a status
review under the Act. In a 12–month
finding, we will determine whether a
petitioned action is warranted after we
have completed a thorough status
review of the species, which is
conducted following a substantial 90–
day finding. Because the Act’s standards
for 90–day and 12–month findings are
different, as described above, a
substantial 90–day finding does not
mean that the 12–month finding will
determine that listing is warranted.
References Cited
A complete list of all references cited
herein is available on the Internet at
https://www.regulations.govand upon
request from the Pacific Islands Fish
and Wildlife Office (see FOR FURTHER
INFORMATION CONTACT).
Author
The primary authors of this notice are
the staff of the Pacific Islands Fish and
Wildlife Office (see FOR FURTHER
INFORMATION CONTACT).
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: June 3, 2010.
Daniel M. Ashe,
Acting Director, U.S. Fish and Wildlife
Service.
[FR Doc. 2010–14430 Filed 6–15–10; 8:45 am]
BILLING CODE 4310–55–S
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
50 CFR Part 665
[Docket No. 0907211157–0224–02]
RIN 0648–AX76
Fisheries in the Western Pacific;
Community Development Program
Process
AGENCY: National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
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[Federal Register Volume 75, Number 115 (Wednesday, June 16, 2010)]
[Proposed Rules]
[Pages 34077-34088]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2010-14430]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R1-ES-2010-0012]
[MO 92210-0-0008-B2]
Endangered and Threatened Wildlife and Plants; 90-Day Finding on
Five Petitions to List Seven Species of Hawaiian Yellow-faced Bees as
Endangered
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 90-day petition finding and initiation of status
review.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), announce a
90-day finding on five petitions to list seven species of Hawaiian
yellow-faced bees (Hylaeus anthracinus, H. assimulans, H. facilis, H.
hilaris, H. kuakea, H. longiceps, and H. mana) as endangered and
designate critical habitat under the Endangered Species Act of 1973, as
amended (Act).
We find that the petitions present substantial scientific or
commercial information indicating that listing these seven species of
Hawaiian yellow-faced bees may be warranted. Therefore, with the
publication of this notice we are initiating a status review of these
species and will issue 12-month findings on our determination as to
whether the petitioned actions are warranted. To ensure that the status
reviews are comprehensive, we are soliciting scientific and commercial
data and other information regarding these species. We will make a
determination on critical habitat for these species if, and when, we
initiate a listing action.
DATES: To allow us adequate time to conduct this review, we request
that information you submit be received by us on or before August 16,
2010. Please note that if you are using the Federal eRulemaking Portal
(see ``ADDRESSES'' section, below), the deadline for submitting an
electronic comment is Eastern Standard Time on this date.
We will post all information received on https://www.regulations.gov. This generally means that we will post any
personal information you provide us (see the Request for Information
section below for more details).
After August 16, 2010, you must submit information directly to the
Field Office (see FOR FURTHER INFORMATION CONTACT section below).
Please note that we might not be able to address or incorporate
information that we receive after the above requested date.
ADDRESSES: You may submit information by one of the following methods:
Federal eRulemaking Portal: https://www.regulations.gov. In
the box that reads ``Enter Keyword or ID,'' enter the Docket number for
this finding, which is FWS-R1-ES-2010-0012. Check the box that reads
``Open for Comment/Submission,'' and then click the Search button. You
should then see an icon that reads ``Submit a Comment.'' Please ensure
that you have found the correct rulemaking before submitting your
comment.
U.S. mail or hand-delivery: Public Comments Processing,
Attn: FWS-R1-ES-2010-0012; Division of Policy and Directives
Management; U.S. Fish and Wildlife Service; 4401 N. Fairfax Drive,
Suite 222; Arlington, VA 22203.
FOR FURTHER INFORMATION CONTACT: Loyal Mehrhoff, Field Supervisor,
Pacific Islands Fish and Wildlife Office, 300 Ala Moana Boulevard, Room
3-122, Honolulu, HI 96850; by telephone (808-792-9400); or by facsimile
(808-792-9581). Persons who use a telecommunications device for the
deaf (TTD) may call the Federal Information Relay Service (FIRS) at
800-877-8339.
SUPPLEMENTARY INFORMATION:
Request for Information
When we make a finding that a petition presents substantial
information indicating that listing a species may be warranted, we are
required to promptly review the status of the species (status review).
For the status review to be complete and based on the best available
scientific and commercial information, we request information on the
seven species of Hawaiian yellow-faced bees (H. anthracinus, H.
assimulans, H. facilis, H. hilaris, H. kuakea, H. longiceps, and H.
mana) from governmental agencies, Native American Tribes, the
scientific community, industry, or any other interested parties. We
seek information on:
(1) The species' biology, range, and population trends, including:
(a) Habitat requirements for feeding, breeding, and sheltering;
(b) Genetics and taxonomy;
(c) Historical and current range including distribution patterns;
(d) Historical and current population levels, and current and
projected trends; and
(e) Past and ongoing conservation measures for the species, its
habitat or both.
(2) Information about the seven Hawaiian yellow-faced bees relevant
to the factors that are the basis for making
[[Page 34078]]
a listing determination for a species under section 4(a) of the Act (16
U.S.C. 1531 et seq.), which are:
(a) The present or threatened destruction, modification, or
curtailment of the species' 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.
(3) Whether insect collecting presents a threat to any of the seven
Hawaiian yellow-faced bee species.
(4) The potential cumulative effects of these factors that may
threaten or endanger the seven Hawaiian yellow-faced bee species.
(5) Management programs for the conservation of the seven Hawaiian
yellow-faced bee species.
(6) The potential effects of climate change on the seven Hawaiian
yellow-faced bee species and their habitat.
If, after the status reviews, we determine that listing any of the
seven Hawaiian yellow-faced bees is warranted, we will propose critical
habitat (see definition in section 3(5)(A) of the Act) under section 4
of the Act, to the maximum extent prudent and determinable at the time
we propose to list the species. Therefore, with regard to areas within
the geographical range currently occupied by these species, we also
request data and information on what may constitute physical or
biological features essential to the conservation of these species;
where these features are currently found; and whether any of these
features may require special management considerations or protection.
In addition, we request data and information regarding whether there
are areas outside the geographical area occupied by these species that
are essential to the conservation of these seven species. Please
provide specific comments and information as to what, if any, critical
habitat you think we should propose for designation if these species
are proposed for listing, and why such habitat meets the requirements
of section 4 of the Act. Please include sufficient information with
your submission (such as scientific journal articles or other
publications) to allow us to verify any scientific or commercial
information you include.
Please note that submissions merely stating support for or
opposition to the action under consideration without providing
supporting information, although noted, will not be considered in
making a determination, as section 4(b)(1)(A) of the Act (16 U.S.C.
1531 et seq.) directs that determinations as to whether any species is
a threatened or endangered species must be made ``solely on the basis
of the best scientific and commercial data available.''
You may submit your information concerning this status review by
one of the methods listed in the ADDRESSES section. If you submit
information via https://www.regulations.gov, your entire submission--
including any personal identifying information--will be posted on the
website. If your submission is made via a hardcopy that includes
personal identifying information, you may request at the top of your
document that we withhold this personal identifying information from
public review. However, we cannot guarantee that we will be able to do
so. We will post all hardcopy submissions on https://www.regulations.gov.
Information and materials we receive, as well as supporting
documentation we used in preparing this finding, is available for you
to review at https://www.regulations.gov, or you may make an appointment
during normal business hours at the Pacific Islands Fish and Wildlife
Office (see FOR FURTHER INFORMATION CONTACT).
Background
Section 4(b)(3)(A) of the Act (16 U.S.C. 1533(b)(3)(A)) requires
that we make a finding on whether a petition to list, delist, or
reclassify a species presents substantial scientific or commercial
information to indicate that the petitioned action may be warranted. We
are to base this finding on information provided in the petition,
supporting information submitted with the petition, and information
otherwise available in our files. To the maximum extent practicable, we
are to make this finding within 90 days of our receipt of the petition
and publish our notice of this finding promptly in the Federal
Register.
Our standard for substantial scientific or commercial information
within the Code of Federal Regulations (CFR) with regard to a 90-day
petition finding is ``that amount of information that would lead a
reasonable person to believe that the measure proposed in the petition
may be warranted'' (50 CFR 424.14(b)). If we find that substantial
scientific or commercial information was presented, we are required to
promptly conduct a species status review, which we subsequently
summarize in our 12-month finding.
We received five petitions dated March 23, 2009, from Scott Hoffman
Black, Executive Director of the Xerces Society (hereafter, ``the
petitioner''). The petitioner requested that we list seven species of
Hawaiian yellow-faced bees as endangered and that critical habitat be
designated under the Act. The petitions were clearly identified as
petitions and included the requisite identification of addresses and
telephone numbers, but did not include the signature of the petitioner,
as stipulated in 50 CFR 424.14(a). Nevertheless, we recognize the five
documents as petitions. Each petition contained information regarding
the species' taxonomy and ecology, historical and current distribution,
present status, and potential causes of decline and current and
potential imminent threats.
On May 8, 2009, we sent a letter to the petitioner acknowledging
our receipt of the petitions and explaining that we reviewed the
information presented in the petition and determined that issuing an
emergency regulation temporarily listing the species under section
4(b)(7) of the Act was not warranted at that time. This notice
constitutes our 90-day finding for the petitions to list seven species
of Hawaiian yellow-faced bees (Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H. longiceps, and H. mana).
Species Information
Taxonomy and Description of Hylaeus
The seven species of bees described in the petitions belong to the
genus Hylaeus. Hylaeus is a large, globally distributed genus comprised
of over 500 species worldwide. In the Hawaiian Islands, the genus
Hylaeus is widespread and very diverse, with 60 native species,
including 20 that are endemic to single islands (Magnacca 2007, p.
174). All 60 Hawaiian species are in the subgenus Nesoprosopis
(Magnacca and Danforth 2006, p. 393). The Hylaeus genus belongs to the
Colletidae family of bees, also known as plasterer bees due to their
habit of lining their nests with salival secretions.
The species of Hylaeus are commonly known as yellow-faced bees or
masked bees for their yellow-to-white facial markings. All of the
Hylaeus species roughly resemble small wasps in appearance, due to
their slender bodies and their seeming lack of setae (sensory hairs).
However, Hylaeus bees have plumose (branched) hairs on the body that
are longest on the sides of the thorax. To a discerning eye, it is
these plumose setae that readily distinguish them from wasps (Michener
2000, p. 55).
[[Page 34079]]
Life History of Hylaeus
Nests of Hylaeus species are usually constructed opportunistically
within dead twigs or plant stems, or other similarly small natural
cavities under bark or rocks. This is unlike the nests of many other
bee species, which are purposefully excavated or constructed
underground. Like other Hylaeus, Hawaiian Hylaeus also lack strong
mandibles and other adaptations for digging and often use nest burrows
abandoned by other insect species (Daly and Magnacca 2003, p. 9). The
adult male and female bees feed upon flower nectar for nourishment.
Many species, including the Hawaiian Hylaeus, lack an external
structure for carrying pollen, called a scopa, and instead internally
transport collected pollen, often mixed with nectar, within their crop
(stomach). Hawaiian Hylaeus species are grouped within two categories:
Ground-nesting species that require relatively dry conditions and wood-
nesting species which are found within wetter areas (Zimmerman 1972, p.
533; Daly and Magnacca 2003, p. 11).
The female Hylaeus bee lays eggs in brood cells that she constructs
in the nest and lines with a self-secreted cellophane-like material.
Prior to sealing the nest, the female provides her young with a mass of
semiliquid nectar and pollen that is left alongside her eggs. Upon
hatching, the grub-like larvae eat the provisions left for them,
pupate, and eventually emerge as adults (Michener 2000, p. 24).
The role of bees as pollinators maintaining communities of native
flora in a diversity of habitats is widely recognized (Cane and
Tepedino 2001, p. 1; Kremen et al. 2007, pp. 302, 307; National
Research Council 2007, p. 13). Recent studies of visitation records of
Hawaiian Hylaeus bees to native flowers (Daly and Magnacca 2003, p. 11)
and pollination studies of native plants (Sakai et al. 1995, pp. 2524-
2528; Cox and Elmqvist 2000, p. 1,238; Sahli et al. 2008, p. 1) have
demonstrated that Hawaiian Hylaeus species almost exclusively visit
native plants to collect nectar and pollen and, in the process,
pollinate these plants. Hylaeus bees are very rarely found visiting
nonnative plants for nectar and pollen (Magnacca 2007, pp. 186, 188),
and are almost completely absent from habitats dominated by nonnative
plant species (Daly and Magnacca 2003, p. 11). Sahli et al. (2008, p.
1) quantified pollinator visitation rates to all of the flowering plant
species in communities on a Hawaiian lava flow dating from 1855 to
understand how pollination webs and the integration of native and alien
species changes with elevation. In that study, eight flowering plants
were observed at six sites, which ranged in elevation from 880 to 2400
meters (m) (2,887 to 7874 feet (ft)). The study also found that the
proportion of native pollinators changed along the elevation gradient;
at least 40 to 50 percent of visits were from alien pollinators at low
elevation, as opposed to 4 to 20 percent of visits by alien pollinators
at mid to high elevations. Hylaeus bees were less abundant at lower
elevations, and there were lower visitation rates of any pollinators to
native plants at lower elevations, which suggests that Hylaeus may not
be easily replaceable by nonnative pollinators (Sahli et al., 2008, p.
1). Because Hylaeus species are likely critical pollinators of one or
more native Hawaiian plant species, it is believed that their decline
or eventual extinction may negatively impact dependent native plant
species (Hopper et al. 1996, p. 8; Cox and Elmqvist 2000, p. 1238).
Taxonomy and Description of Each Petitioned Hylaeus Species
Unless clearly stated that the information is from our files, all
information, statements, and references cited regarding the taxonomy,
descriptions, life history, and range and distribution are based on
information submitted in the petitions.
Hylaeus anthracinus
Taxonomy
Hylaeus anthracinus was first described as Prosopis anthracina by
Smith in 1873 (Daly and Magnacca 2003, p. 55), and transferred to
Nesoprosopis 20 years later (Perkins 1899, pp. 75), and then
Nesoprosopis was reduced to a subgenus of Hylaeus in 1923 (Meade-Waldo
1923, p. 1). Although the distinctness of this species is unquestioned,
recent genetic evidence (Magnacca and Brown, submitted) suggests that
H. anthracinus may be composed of three cryptic (not recognized)
species which represent the populations on Hawaii; Maui and Kahoolawe;
and Molokai and Oahu.
Description
Hylaeus anthracinus is a medium-sized black bee with clear to smoky
wings and black legs. The male has a single large yellow spot on his
face, while below the antennal sockets the face is yellow. The female
is entirely black and can be distinguished by the black hairs on the
end of the abdomen and an unusual mandible that has three teeth, a
characteristic that is shared only with H. flavifrons, a closely
related species on Kauai (Daly and Magnacca 2003, p. 53).
Life History
The diet of the larval stage of Hylaeus anthracinus is unknown,
although the larvae are presumed to feed on stores of pollen and nectar
collected and deposited in the nest by the adult female. Likewise, the
nesting habits of H. anthracinus are not known, but the species is
thought to nest within the stems of coastal shrubs (Magnacca 2005a, p.
2).
Hylaeus anthracinus adults have been observed visiting the flowers
of Sesbania tomentosa (oahi), Scaevola sericea (naupaka kahakai), Sida
fallax (ilima), Argemone glauca (pua kala), Chamaesyce celastroides
(akoko), Chamaesyce degeneri (akoko), Heliotropium anomalum (hinahina),
and Myoporum sandwicense (naio). This species has also been collected
from inside the fruit capsule of Kadua coriacea (kioele) (Magnacca
2005a, p. 2). Hylaeus anthracinus has also been observed visiting
Tournefortia argentea (tree heliotrope), a tree native to tropical
Asia, Madagascar, tropical Australia, and Polynesia, for nectar and
pollen (Wagner et al. 1999, p. 398; Daly and Magnacca 2003, p. 55;
Magnacca 2007a, p. 181). The species was first collected on Oahu in
1864-1865, and is naturalized and documented from all of the main
islands except Kahoolawe (Wagner et al. p. 398). It is described as
introduced by Magnacca (2007, p. 181). Hylaeus anthracinus commonly
occurs alongside other Hylaeus species, including H. longiceps and H.
flavipes.
Range and Distribution
Hylaeus anthracinus was historically known from numerous coastal
strand and lowland dry forest locations up to 2,000 feet (ft) (610
meters (m)) in elevation on the islands of Hawaii, Lanai, Maui,
Molokai, and Oahu. According to the petition, between 1997 and 2008,
surveys for Hawaiian Hylaeus were conducted at 43 sites throughout the
Hawaiian Islands that were either historic collecting localities for H.
anthracinus, or potentially suitable habitat for this species. Hylaeus
anthracinus was observed at 14 of the 43 survey sites, but had
disappeared from each of the 9 historically occupied sites that were
surveyed (petition p. 7). Several of the historical collection sites,
such as Honolulu and Waikiki on Oahu, and Kealakekua Bay on Hawaii, no
longer contain Hylaeus habitat, which has been replaced by urban
development or is dominated by nonnative vegetation (Liebherr and
Polhemus 1997, pp. 346-347; Daly and
[[Page 34080]]
Magnacca 2003, p. 55; Magnacca 2007, pp. 186-188).
Hylaeus anthracinus is currently restricted to small populations in
a few small patches of coastal and lowland dry habitat (Magnacca 2005a,
p. 2); one location on Kahoolawe; five locations on the island of
Hawaii, two locations on Maui, three locations on Molokai, and two
locations on Oahu (Xerces 2009a, pp. 9-10). The petition does not
define the context applied to the term ``small,'' and we have no
additional information in our files. Accordingly, we are presenting the
population information as characterized by the petitioner. The 2004 H.
anthracinus collection on the island of Hawaii occurred in montane dry
forest (Magnacca 2005a, p. 2). Although it was previously unknown from
the island of Kahoolawe, H. anthracinus was observed at one location on
the island in 2002 (Daly and Magnacca 2003, p. 55). According to the
petition, it is believed to be extirpated from Lanai (Daly and Magnacca
2003, p. 55).
Hylaeus assimulans
Taxonomy
Hylaeus assimulans was first described as Nesoprosopis assimulans
(Perkins 1899, pp. 75, 101-102), and then Nesoprosopis was reduced to a
subgenus of Hylaeus in 1923 (Meade-Waldo 1923, p. 1). The species was
most recently described as Hylaeus assimulans by Daly and Magnacca in
2003 (pp. 55-56).
Description
Hylaeus assimulans is distinguished by its large size relative to
other coastal Hylaeus species and slightly smoky to smoky-colored
wings. The male is black with yellow face marks, with an almost
entirely yellow clypeus (lower face region) with additional marks on
the sides that narrow dorsally (towards the top). The male also has
brown appressed (flattened) hairs on the tip of the abdomen. The female
is entirely black, large-bodied, and has no distinct punctuation on the
abdomen (Daly and Magnacca 2003, p. 56).
Life History
The diet of the larval stage of Hylaeus assimulans is unknown,
although the larvae are presumed to feed on stores of pollen and nectar
collected and deposited in the nest by the female adult (Xerces 2009b,
p. 11). Likewise, the nesting habits of H. assimulans are not known,
but the species is thought to nest underground, as do other closely
related species (Magnacca 2005b, p. 2).
Hylaeus assimulans adults have been observed visiting the flowers
of Lipochaeta lobata (nehe) and Sida fallax (ilima), this species'
likely primary host plant (Xerces 2009b, p. 10). Hylaeus assimulans
appears to be closely associated with plants in the genus Sida, and the
petitioner suggests this yellow-faced bee species may be more common
where this plant is abundant (Daly and Magnacca 2003, pp. 58, 217;
Magnacca 2007, p. 183). The petition contains information indicating
that in recent collections, H. assimulans seems to be more common in
dry forest at relatively higher elevations, and is less often found in
coastal strand habitat. The petition states that the frequency of H.
assimulans observations in higher, dry forest may be related to the
abundance of Sida in the understory (Magnacca 2005b, p. 2). The
petitioner also states that it is likely that H. assimulans visits
several other native plants, including Acacia koa (koa), Metrosideros
polymorpha (ohia), Styphelia tameiameiae (pukiawe), and species of
Scaevola (naupaka) and Chamaesyce (akoko), which are frequented by
other Hylaeus species as well.
Range and Distribution
Historically, Hylaeus assimulans was known from numerous coastal
strand and lowland dry locations up to 2,000 ft (610 m) in elevation on
the islands of Lanai, Maui, and Oahu. Although there are no collections
from Molokai, the petition states that H. assimulans also occurred
there because all other species of Hylaeus known from Maui, Lanai, and
Oahu also occurred on Molokai (Xerces 2009b, p. 6). Between 1997 and
2008, surveys for Hawaiian Hylaeus were conducted in 25 sites on Oahu,
Maui, Kahoolawe, Lanai, and Molokai. Hylaeus assimulans was absent from
six of its historic localities on Oahu, Maui, and Lanai (Xerces 2009b,
pp. 6-7). Hylaeus assimulans was not observed at 19 other sites with
potentially suitable habitat on Oahu, Maui, Lanai, and Molokai,
including several sites from which other native Hylaeus species have
been recently collected (Daly and Magnacca 2003, p. 56; Xerces 2009b,
p. 7).
Currently, Hylaeus assimulans is found in a few small patches of
coastal and lowland dry habitat at two locations on Lanai, two
locations on Maui, and one location on Kahoolawe (Daly and Magnacca
2003, p. 58; Magnacca 2005, p. 2). According to the petition, this
species has likely been extirpated from Oahu since it was absent from
the island's best extant coastal strand habitat at Kaena Point (Kaena
Point Natural Area Reserve (NAR)) (Magnacca 2005, p. 2).
Hylaeus facilis
Taxonomy
According to the petitioner, Hylaeus facilis is a member of the H.
difficilis species group, and is closely related to H. chlorostictus
and H. simplex. Hylaeus facilis was first described as Prosopis facilis
by Smith in 1879 (Daly and Magnacca, p. 80), based on a specimen
erroneously reported from Maui. According to Blackburn and Cameron
(1886 and 1887), the species' type locality was Pauoa Valley on Oahu
(Daly and Magnacca 2003, p. 80). The species was later transferred to
the genus Nesoprosopis (Perkins 1899, pp. 75, 77). Nesoprosopis was
subsequently reduced to a subgenus of Hylaeus (Meade-Waldo 1923, p. 1).
The species was most recently recognized by Daly and Magnacca (2003, p.
80) as Hylaeus facilis.
Description
Hylaeus facilis is a medium-sized bee with smoky colored wings. The
male has an oval yellow mark on its face that covers the entire clypeus
(lower face region), and a narrow stripe beside the eyes, but is
otherwise unmarked. The large, externally visible gonoforceps (paired
lateral outer parts of the male genitalia) distinguish H. facilis from
the closely related H. simplex (Daly and Magnacca 2003, p. 83). The
female is entirely black, and indistinguishable from females of H.
difficilis and H. simplex (Daly and Magnacca 2003, pp. 81-82).
Life History
The diet of the larval stage of Hylaeus facilis is unknown,
although the larvae are presumed to feed on stores of pollen and nectar
collected and deposited in the nest by the adult female. The nesting
habits of H. facilis have not been observed, but the species is thought
to nest underground as do the closely related species H. chlorostictus
and H. simplex (Daly and Magnacca 2003, p. 83; Magnacca 2005c, p. 2).
According to the petition, the native host plants of adult Hylaeus
facilis are unknown, but it is likely that this species visits several
plants that other Hylaeus species are known to frequent, including
Acacia koa (koa), Metrosideros polymorpha (ohia), Styphelia tameiameiae
(pukiawe), Scaevola spp. (naupaka), and Chamaesyce spp. (akoko).
Hylaeus facilis has also been observed visiting the nonnative
Tourneforia argentea (tree heliotrope) for nectar and pollen (Magnacca
2007, p. 181).
[[Page 34081]]
Range and Distribution
Hylaeus facilis was historically known from Lanai, Maui, Molokai,
and Oahu, in dry shrubland to wet forest, from coastal to montane
habitat up to 3,281 ft (1,000 m) in elevation (Gagne and Cuddihy 1999,
p. 93; Daly and Magnacca 2003, pp. 81, 83). Perkins (1899, p. 77)
remarked that H. facilis was among the most common and widespread
Hylaeus species on Oahu and all of Maui Nui (Lanai, Molokai, and Maui)
(Magnacca 2007, p. 183). The petitioner contends that although the
species was widely collected within a diverse range of habitats, it
probably prefers dry to mesic forest and shrubland (Magnacca 2005c, p.
2), which are increasingly rare and patchily distributed habitats
(Smith 1985, pp. 227-233; Juvik and Juvik 1998, p. 124; Wagner et al.
1999, pp. 66-67, 75; Magnacca 2005c, p. 2).
The petition states that Hylaeus facilis has almost entirely
disappeared from most of its historical range (Daly and Magnacca 2003,
p. 7; Magnacca 2007, p. 183), and the abundance of specimens in the
collections at the Bishop Museum in Honolulu demonstrates the historic
prevalence of this species in a diverse array of habitats and
elevations (Magnacca 2007, p. 183). Between 1998 and 2006, 39 sites on
Oahu, Maui, Lanai, and Molokai were surveyed; H. facilis was absent
from each of the 13 historical localities that were revisited (Magnacca
2007, p. 183). Hylaeus facilis was not observed at 26 other sites with
potentially suitable habitat, including many sites from which other
native Hylaeus species have been recently collected (Daly and Magnacca
2003, pp. 7, 81-82; Magnacca 2007, p. 183).
Currently, Hylaeus facilis is only known from three sites, one each
on the islands of Maui, Molokai, and Oahu (Daly and Magnacca 2003, pp.
81-82; Magnacca 2005c, p. 2). According to the petitioner, this species
is likely extirpated from Lanai (Xerces 2009c, p. 7).
Hylaeus hilaris
Taxonomy
Hylaeus hilaris was first described as Prosopis hilaris by Smith in
1879 (Daly and Magnacca 2003, pp. 103-104), transferred to the genus
Nesoprosopis 20 years later (Perkins 1899, pp. 75), and then
Nesoprosopis was reduced to a subgenus of Hylaeus in 1923 (Meade-Waldo
1923, p. 1). In 2003, Daly and Magnacca described the species as
Hylaeus hilaris (Daly and Magnacca 2003, pp. 103-104).
Description
Hylaeus hilaris is distinguished by its large size (male wing
length is 0.185 inches (in) (4.7 millimeters (mm)) relative to other
coastal Hylaeus species. The wings of this species are slightly smoky
to smoky colored, and it is the most colorful of the Hawaiian Hyaleus
species. The face of the male is almost entirely yellow, with yellow
markings on the legs and thorax, and the metasoma (middle portion of
the abdomen) are usually predominantly red. Females are drably colored,
with various brownish markings. As with other cleptoparasitic (see
``Life History'' below) species, H. hilaris lacks the specialized
pollen-sweeping hairs of the front legs (Daly and Magnacca 2003, pp. 9,
106). It is also one of only two Hawaiian Hyaleus species to possess
apical (at the end or tip of a structure) bands of fine white hairs on
the segments of the metasoma.
Life History
Most adult Hawaiian Hylaeus species consume nectar for energy;
however, Hylaeus hilaris has yet to be observed actually feeding from
flowers. Hylaeus hilaris and the four species related to it (H.
hostilis, H. inquilina, H. sphecodoides, and H. volatilis) are known as
cleptoparasites or cuckoo bees. The mated female does not construct a
nest or collect pollen, but instead enters the nest of another species
and lays an egg in a partially provisioned cell. Upon emerging, the
cleptoparasitic larva kills the host egg and consumes the provisions,
pupates, and eventually emerges as an adult. As a result of this
lifestyle shift, H. hilaris bees have lost the pollen-collecting hairs
that other species possess on the front legs. Cleptoparasitism is
actually quite common among bees: approximately 25 percent of known bee
species have evolved to become cleptoparasites. Among the world's bees,
other than the Hawaiian Hylaeus group, no cleptoparasites are known
from the family Colletidae (Daly and Magnacca 2003, p. 9). The larvae
of H. hilaris and their diet are unknown (Magnacca 2005d, p. 2);
however, the species is known to lay its eggs within the nests of H.
anthracinus, H. assimulans, and H. longiceps (Perkins 1913, p. lxxxi).
Although the species has never been observed at flowers, H. hilaris
adults presumably consume nectar as a food source (Xerces 2009 d, p.
9). Hylaeus hilaris depends on a number of related Hylaeus host species
for its parasitic larvae, and its population size is inherently much
smaller than its host species (Xerces 2009d, p. 9).
Range and Distribution
Hylaeus hilaris was historically known from coastal strand habitat
on the islands of Lanai, Maui, and Molokai. The petitioner states that
it is believed to have occurred along much of the coast of these
islands since its primary hosts, H. anthracinus, H. assimulans, and H.
longiceps, likely extended throughout this habitat. According to the
petition, nearly all of the coastal strand habitat has been either
developed or degraded, and is no longer suitable for H. hilaris
(Liebherr and Polhemus 1997, pp. 346-347; Magnacca 2007, pp. 186-188).
Hylaeus hilaris was absent from three of its historical population
sites revisited by researchers between 1998 and 2006. It was also not
observed at 10 additional sites with potentially suitable habitat where
other native Hylaeus species have been recently collected (Daly and
Magnacca 2003, pp. 103, 106).
The petitioner states that this species has been collected only
twice in the last 70 years, but acknowledges a gap of about 70 years
between major collecting efforts (Xerces 2009d, p. 6). Hylaeus hilaris
has recently been collected on two occasions; once in 1989 and again in
1999. The species was absent from each of its historical localities
that were revisited between 1998 and 2006 (Xerces 2009d, p. 6).
Currently, the only known population of H. hilaris is located on The
Nature Conservancy's Moomomi Preserve on Molokai (Daly and Magnacca
2003, pp. 103, 106; Magnacca 2005d, p. 2). According to the petition,
it is no longer extant on Lanai (Xerces 2009d, p. 6).
Hylaeus kuakea
Taxonomy and Description
Hylaeus kuakea was first described by Daly and Magnacca (2003, pp.
1, 125-127) from specimens collected in 1997 in the Waianae Mountains
on Oahu. Hylaeus kuakea is a small, black bee with slightly smoky
colored wings. This species does not fit into any of the well-defined
Hylaeus species groups. Its facial marks are similar to those of the H.
difficilis group and to H. anthracinus, but it can be distinguished by
its unusual ivory facial marking covering the clypeus (the lower face
region). Hylaeus kuakea also resembles H. anthracinus, but has a
denser, more distinct arrangement of setae (sensory hairs) on the head
and generally narrower marks next to the compound eyes (Daly and
Magnacca 2003, p. 125; Magnacca 2005e, p. 2). Only two adult
[[Page 34082]]
male specimens have been collected; females have yet to be collected or
observed.
Life History
The diet of the larval stage of Hylaeus kuakea is unknown, although
the larvae are presumed to feed on stores of pollen and nectar
collected and deposited in the nest by the adult female (Xerces 2009e,
p. 7). The nesting habits of H. kuakea have not been observed, but the
species is believed to be related to other wood-nesting Hawaiian
Hylaeus species (Magnacca and Danforth 2006, p. 403).
According to information in the petition, the native host plants of
the adult Hylaeus kuakea are unknown, but it is likely that this
species visits several plants that other Hylaeus species are known to
frequent, including Acacia koa, Metrosideros polymorpha, Styphelia
tameiameiae, Scaevola spp., and Chamaesyce spp. (Magnacca 2005e, p. 2).
Range and Distribution
Hylaeus kuakea is only known from two collections made in Moho
Gulch Ridge, at the northern end of Honouliuli Preserve, at an
elevation of about 1,900 ft (579 m) in the Waianae Mountains on Oahu.
Hylaeus kuakea is found in lowland mesic forest, which is increasingly
rare and patchily distributed on Oahu (Smith 1985, pp. 227-233; Juvik
and Juvik 1998, p. 124; Wagner et al. 1999, pp. 66-67, 75).
According to the petitioner, although there is potentially suitable
lowland mesic habitat in Honouliuli Preserve, no other individuals of
Hylaeus kuakea were found in surveys subsequent to the type collection
in 1997 (Magnacca 2007, p. 184). In addition, Perkins did not collect
this species in surveys in the Honouliuli Preserve vicinity or in
nearby areas in 1899, 1910, and 1911 (Xerces 2009e, p. 6). The
petitioner therefore concludes that the extreme rarity of this species,
its absence from nearby sites, and the fact that it was not discovered
until very recently suggest that very few populations remain (Magnacca
2005e, p. 2).
Hylaeus longiceps
Taxonomy
Hylaeus longiceps was first described in 1899 as Nesoprosopis
longiceps (Perkins 1899, pp. 75, 98), and then Nesoprosopis was reduced
to a subgenus of Hylaeus in 1923 (Meade-Waldo 1923, p. 1). Daly and
Magnacca (2003, pp. 133-134) most recently described the species as
Hylaeus longiceps.
Description
Hylaeus longiceps is a small to medium-sized, black bee with clear
to slightly smoky colored wings. Its distinguishing characteristics are
its long head and the facial marks of the male. The lower face of the
male is marked with a yellow band that extends at the sides of the face
in a broad stripe above the antennal sockets. The area above the
clypeus (lower face region) is very long and narrow, and the scape (the
first antennal segment) is noticeably twice as long as it is wide. The
female is entirely black and unmarked (Daly and Magnacca 2003, p. 133).
Life History
The diet of the larval stage of Hylaeus longiceps is unknown,
although the larvae are presumed to feed on stores of pollen and nectar
collected and deposited in the nest by the female adult (Xerces 2009a,
p. 15). The nesting habits of H. longiceps are unknown, but the species
is thought to nest underground, as in other closely related species
(Magnacca 2005f, p. 2).
Hylaeus longiceps adults have been observed visiting the flowers of
a wide variety of plants, including Scaevola coriacea (dwarf naupaka),
Sida fallax, Scaevola spp. (naupaka kahakai), Sesbania tomentosa
(ohai), Myoposum sandwicense (naio), Santalum ellipticum (iliahialoe,
coast sandalwood), Chamaesyce degeneri (akoko), and Vitex rotundifolia
(pohinahina) (Xerces 2009a, p. 14). The petitioner reports that it is
likely that H. longiceps visits several plant species that other
Hylaeus species are known to frequently visit, including Scaevola spp.,
Chamaesyce spp., Tournefortia argentea, Jacquemontia ovata (pau o
hiiaka), and Sida fallax (Magnacca 2005f, p. 2).
Range and Distribution
Hylaeus longiceps is historically known from numerous coastal
strand and lowland dry shrubland locations up to 2,000 ft (610 m) in
elevation on the islands of Lanai, Maui, Molokai, and Oahu. The species
is primarily known from coastal habitat, but is infrequently collected
in dry shrubland. Hylaeus longiceps is rarely observed in higher
elevation dry forests. Perkins (1899, p. 98) noted that H. longiceps
was locally abundant, and probably occurred historically throughout
much of the leeward and lowland areas on Maui Nui (Maui, Molokai,
Lanai, and Kahoolawe) and Oahu, since its host plants, Sida fallax,
Chamaesyce spp., Scaevola spp., and Jaquemontia ovata, occurred
throughout these areas (Magnacca 2005f, p. 2). The petitioner states
that most of the habitat in these areas has been either developed or
degraded and is no longer suitable for H. longiceps (Liebherr and
Polhemus 1997, pp. 346-347; Magnacca 2007, pp. 186-188).
Hylaeus longiceps is now restricted to small populations in small
patches of coastal and lowland habitat on Lanai, Maui, Molokai, and
Oahu (Magnacca 2005f, p. 2). Twenty-five sites that were either
historic collecting localities for H. longiceps or contained
potentially suitable habitat for this species were surveyed between
1997 and 2008. Hylaeus longiceps was observed at only six of the
surveyed sites: three sites on Lanai, one site on Maui, one site on
Molokai, and one site on Oahu. Only one historic location, Waieu Dune
on Maui, still supports a population of H. longiceps (Daly and Magnacca
2003, p. 135).
Hylaeus mana
Taxonomy and Description
Hylaeus mana was first described by Daly and Magnacca (2003, pp.
135-136) from four specimens collected in 2002 on the leeward side of
the Koolau Mountains on Oahu. This species is an extremely small,
gracile (gracefully slender) black bee with yellow markings on the
face. The smallest of all Hawaiian Hylaeus species, H. mana is a member
of the dumetorum species group. The face of the male is largely yellow
below the antennae, extending dorsally in a narrowing stripe. The
female's face has three yellow lines, one against each eye, and a
transverse stripe at the apex of the clypeus (lower face region). The
female's other markings are the same as the male's (Daly and Magnacca
2003, p. 135). Hylaeus mana can be distinguished from H. mimicus and H.
specularis, with whom its range overlaps, by its extremely small size,
the shape of the male's genitalia, the female's extensive facial marks,
and a transverse rather than longitudinal clypeal marking (Daly and
Magnacca 2003, p. 138).
Life History
The diet of the larval stage of Hylaeus mana is unknown, although
the larvae are presumed to feed on stores of pollen and nectar
collected and deposited in the nest by the adult female (Xerces 2009e,
p. 7). The nesting habits of H. mana are not well known, but it is
assumed the species is closely related to other wood-nesting Hawaiian
Hylaeus species (Magnacca and Danforth 2006, p. 403).
Adult specimens of Hylaeus mana were collected while they visited
[[Page 34083]]
flowers of Santalum freycinetianum var. freycinetianum (iliahi,
sandalwood), a native Hawaiian plant found only on Oahu and Molokai
(Wagner et al. 1999, p. 1221). The petitioner asserts that it is likely
that H. mana visits several other native plant species, including
Acacia koa, Metrosideros polymorpha, Styphelia tameiameiae, Scaevola
spp., and Chamaesyce spp. (Magnacca 2005g, p. 2).
Range and Distribution
Hylaeus mana is only known from lowland mesic forest located along
the Manana Trail in the Koolau Mountains on Oahu, at an elevation of
about 1,400 ft (427 m). Few Hylaeus bees have been found in this type
of koa-dominated, lowland mesic forest on Oahu (Daly and Magnacca 2003,
p. 138). This type of forest is increasingly rare and patchily
distributed on Oahu (Smith 1985, pp. 227-233; Juvik and Juvik 1998, p.
124; Wagner et al. 1999, pp. 66-67, 75).
According to the petition, because the first collection of Hylaeus
mana was made in 2002, the historic range and current distribution,
other than the collection on Manana Trail, are unknown at this time
(Magnacca 2005g, p. 2). This species was not found in surveys of
potentially suitable habitat in the same general area by Perkins in
1899, 1910, and 1911 (Xerces 2009e, p. 6). The petitioner therefore
concludes that the extreme rarity of this species, its absence from
nearby sites, and the fact that it was not discovered until very
recently suggest that very few populations remain (Magnacca 2005g, p.
2).
We accept the characterization of the seven species of Hawaiian
yellow-faced bees (Hylaeus anthracinus, H. assimulans, H. facilis, H.
hilaris, H. kuakea, H. longiceps, and H. mana) as described in the
information provided by the petitioner.
Factors Affecting the Species
Section 4 of the Act (16 U.S.C. 1533), and its implementing
regulations at 50 CFR 424, set forth 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; or (E) other natural or manmade factors affecting its
continued existence. In making this 90-day finding, we evaluated
whether information on threats to the seven Hawaiian Hylaeus bee
species presented in the petitions and available in our files at the
time of the review of the petitions constitute substantial scientific
or commercial information such that listing the species may be
warranted. Our evaluation of this information is discussed below.
Unless clearly stated that the information is from our files, all
threats described below and their effects on the seven Hawaiian Hylaeus
bee species are based on information submitted in the petitions. Any
references provided in support of particular statements related to
potential threats are the petitioner's references, unless specifically
identified otherwise.
A. Present or Threatened Destruction, Modification, or Curtailment of
the Species' Habitat or Range
According to the petitions, degradation and loss of coastal and
lowland habitat used by Hylaeus bees on all of the main Hawaiian
Islands is the primary threat to these seven species (Cuddihy and Stone
1990, pp. 60-61; Daly and Magnacca 2003, pp. 55, 173). Coastal and
lowland habitats have been severely altered and degraded, partly
because of past and present land management practices, including
agriculture, grazing, and urban development; the deliberate and
accidental introductions of nonnative animals and plants; and
recreational activities. In addition, the petitions present information
indicating that fire is a potential threat to the habitat of these
seven species in some locations.
Habitat Destruction and Modification by Urbanization and Land Use
Conversion
Increased access to coastal areas, and resulting habitat
disturbance, has been facilitated by coastal development and
roadbuilding (Cuddihy and Stone 1990, pp. 94-95). As described in the
petitions, five species (Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, and H. longiceps) were once widespread and common
in coastal strand habitat (Perkins 1912, p. 688). These five Hylaeus
species are now absent from all of Perkins' coastal collection
localities. Hylaeus facilis has recently been collected in coastal
habitat at Kuololimu Point, and H. hilaris has recently been collected
in coastal habitat at Moomomi Preserve, Molokai (Xerces 2009c, p. 9).
The petitioner states that lowland dry forests and shrublands have
been heavily impacted by urbanization and conversion to agriculture or
pasture throughout the Hawaiian Islands, with the estimated loss of
more than 90 percent of dry forests and shrublands (Bruegmann 1996, p.
26; Juvik and Juvik 1998, p. 124). Four species (Hylaeus anthracinus,
H. assimulans, H. facilis, and H. longiceps) were once widespread and
found within lowland dry habitat on several islands, including Hawaii,
Lanai, Maui, Molokai, and Oahu, but are largely absent from their
historical population sites on these islands.
Mesic forest, once abundant and considered the most diverse of all
Hawaiian forest types (Rock 1913, p. 9), is now very rare, with much of
it converted to pasture, or military or agricultural use, or lost to
urbanization (Cuddihy and Stone 1990, p. 61; Magnacca 2007, p. 187).
Fire has also negatively impacted this habitat type, as is discussed
below. Hylaeus facilis was historically the most wide ranging of the
seven bee species in terms of the variety of habitats from which it was
recorded, which included mesic forest on Lanai, Maui, Molokai, and
Oahu. This species is now restricted to single locations on the islands
of Molokai and Oahu.
The petitioner identified the loss of coastal, dry lowland, and
montane wet forest habitat on Oahu, Lanai, Maui, and Molokai as a
contributing factor to the decline of H. facilis, but acknowledges that
``although recorded from several sites currently considered to be wet
forest, it is possible that H. facilis would not normally inhabit this
[habitat] in a natural state.'' The petitioner attributes the current
observation of this species at sites now known to be wetter than they
were during the early Perkins' collecting period to the more open
understory vegetation (Perkins 1899, p. 76). It is conceivable that the
loss of mesic forest habitat used by H. kuakea is due to urbanization
and land use conversion, although the petitioner presents no
information in this regard, nor do we have information in our files
regarding the historical locations of these two species, both of which
were only recently collected (H. kuakea in 1997; H. mana in 2002).
Habitat Destruction and Modification by Nonnative Plants
The petitioner states that the spread of nonnative plant species is
one of the primary causes of decline, and a current threat to the
existing populations of each of the seven Hylaeus bee species, because
they depend closely on native vegetation for nectar and pollen, and the
bees are almost entirely absent from habitat dominated by invasive,
nonnative vegetation (Sakai et al. 2002, pp. 276, 291; Daly and
Magnacca 2003, p. 11; Liebherr 2005, p. 186). According to information
available in our files and
[[Page 34084]]
presented by the petitioner, the native flora within a majority of
lowland habitat on the Hawaiian Islands below 1,969 ft (600 m) is being
replaced by aggressive, nonnative plant species (Cuddihy and Stone
1990, pp. 73-74; Wagner et al. 1999, p. 52). The petitioner states that
many native plant species that are replaced by nonnative plants were
once foraging resources for numerous Hylaeus species (Cox and Elmqvist
2000, p. 1238; Daly and Magnacca 2003, p. 11; USFWS 1999, pp. 145, 163,
171, 180; USFWS 2008, pp. 7, 9). Six of the seven Hylaeus bee species
(Hylaeus anthracinus, H. assimulans, H. facilis, H. kuakea, H.
longiceps, and H. mana) are most often found in dry and mesic forest
and shrubland habitat (Daly and Magnacca 2003, p. 11), and the greatest
proportion of endangered or at-risk Hawaiian plant species are also
limited to these same habitats; 25 percent of Hawaiian listed plant
species are from dry forest and shrubland alone (Sakai et al. 2002, pp.
276, 291, 292). The petitioner asserts that lowland dry and mesic
forests now support less-diverse Hylaeus communities because many
native plants used for foraging are extirpated from these habitats
(Magnacca 2007, pp. 186-187).
The petitioner states that besides Scaevola sericea (naupaka
kahakai), native vegetation is lacking along most of the coastline of
the major Hawaiian Islands, and that Hylaeus bees cannot survive on
this plant alone (Magnacca 2007, p. 187). The petitioner also states
that native coastal vegetation in many areas, such as Moomomi Preserve
on Molokai, which currently is the only known location for Hylaeus
hilaris, is threatened by Prosopis pallida (kiawe), an invasive,
nonnative, deciduous thorny tree (Xerces 2009a, p. 25; 2009b, p. 17;
2009c, p. 21; 2009d, p. 11).
According to the petitions, many of the native plants that serve as
foraging resources for the adults of the seven Hylaeus bee species are
declining due to a lack of pollinators (Daly and Magnacca 2003, p. 11;
USFWS 2008, pp. 7, 9) and are found only in very small populations
(USFWS 1999, pp. 145, 163, 171, 180; Cox and Elmqvist 2000, p. 1238).
The petitioner points out, for example, that H. longiceps and H.
anthracinus are known to forage on the federally endangered plant
Sesbania tomentosa (ohai). Both H. longiceps and H. anthracinus also
visit Chamaesyce celastroides var. kaenana (akoko), a federally
endangered plant endemic to coastal dry shrubland on Oahu (Daly and
Magnacca 2003, pp. 55, 74). Hylaeus longiceps is also known to forage
on the endangered Scaevola coriacea (dwarf naupaka) (USFWS 1999, p.
145; Daly and Magnacca 2003, pp. 55, 135). In addition, H. anthracinus
has been collected from inside the fruit capsule of Kadua coriacea
(kioele), a federally endangered dry forest plant, known from fewer
than 300 individuals on the island of Hawaii (USFWS 2008, p. 5;
Christian Torres, USFWS, pers. comm. 2009).
Habitat Destruction and Modification by Nonnative Ungulates
The petitioner claims that the decline of native plant communities
has likely had a negative impact on Hawaii's Hylaeus bee species
(Cuddihy and Stone 1990, pp. 59-66, 88-94, 73-76; USFWS 2006, p. 2684).
The presence of nonnative mammals, such as feral pigs (Sus scrofa),
cattle (Bos taurus), goats (Capra hircus), and axis deer (Axis axis),
is considered one of the primary factors underlying the alteration and
degradation of native vegetation and habitat in the Hawaiian Islands
(Stone 1985, pp. 262-263; Cuddihy and Stone 1990, pp. 60-66; 73 FR
73801). Beyond the direct effects of trampling and consuming native
plants, nonnative ungulates contribute significantly to increased
erosion, and their behavior (i.e., rooting, moving across large
expanses) facilitates the spread and establishment of competing,
invasive, nonnative plant species (Xerces 2009a, p. 26; 2009b, p. 18;
2009c, pp. 21-22, 2009d, pp. 12-13, 2009e, p. 10). Several endangered
coastal and lowland plant species that are threatened by the browsing,
trampling, and digging activities of nonnative ungulates are confirmed
foraging sources for Hylaeus species and, therefore, are likely
foraging sources for these seven Hylaeus species (USFWS 1999, pp. 145,
163, 171, 180; Daly and Magnacca 2003, pp. 11, 13).
Habitat Destruction and Modification by Fire
The petitions state that fire can dramatically alter the species
composition of plant communities in coastal and lowland habitats
(Hughes et al. 1991, p. 743; Blackmore and Vitousek 2000, p. 625), and
thus potentially impact Hylaeus populations. The petitioner also
suggests that ordnance-induced fires on the Army's Pohakuloa Training
Area on the island of Hawaii may threaten the dry forest habitat of
Hylaeus anthracinus. Fires were uncommon in the Hawaiian Islands until
the arrival of humans about 2,000 years ago (Smith and Tunison 1992,
pp. 394-395). Native habitat in the Hawaiian Islands has been
increasingly colonized by fire-adapted invasive plant species that take
the place of, and permanently replace, native plant species (Cuddihy
and Stone, pp. 88-94; Smith and Tunison 1992, pp. 394-395; D'Antonio et
al. 2000, pp. 73-74). This process has been facilitated by nonnative
ungulates, which alter the floral composition of native habitats,
making conditions more conducive to fire. This impact occurs because of
the browsing and trampling of native vegetation, and the spreading of
seeds of nonnative, fire-adapted plant species such as Melinis
minutiflora (molasses grass) and Schizachyrium condensatum (tufted
beardgrass) (D'Antonio et al. 2000, pp. 73-74).
Habitat Destruction and Modification by Recreational Activities
The petitions state that some of the best habitat areas for Hylaeus
species are also popular recreational sites, particularly those areas
located within coastal strand habitat (Xerces 2009a, p. 27; 2009c, p.
22). Human impacts at recreational sites may include removal or
trampling of vegetation on or near trails and the compaction of
vegetation by off-road vehicles (Xerces 2009a, p. 27; 2009c, p. 22). In
particular, the petitioner claims that Hylaeus facilis habitat may be
threatened by recreational activities, such as hunting and hiking on
the Poamoho Trail on Oahu (Hawaii Department of Land and Natural
Resources 2000, p. 15; Xerces 2009c, p. 22). According to the
petitions, some of the best remaining habitat for H. anthracinus and H.
longiceps includes Kaena Point (on Oahu), Kona Coast State Park,
Makalawena, Mokuauia, and South Point (on the island of Hawaii), areas
that are popular recreational sites with largely unregulated access
(Xerces 2009a, p. 27).
Habitat Destruction and Modification by Climate Change
The petitioner asserts that a changing climate may cause shifts in
the range of Hylaeus host plant species, which can be especially
detrimental to dependent pollinators like these seven species when
combined with habitat loss (National Research Council 2007, p. 102).
Most bees have difficulty crossing large geographical barriers
(Michener 2000, p. 103), and successive generations of solitary species
like Hylaeus tend to nest in the same area year after year. The
petitioner points out that the seven Hylaeus bee species are restricted
to habitat patches where native host plant species are present, and
argues that they are not likely to disperse far to find new habitat
(Xerces 2009a, p. 30; 2009b, p. 21; 2009c, p. 25; 2009d, p. 14; 2009e,
p. 13). Thus, the
[[Page 34085]]
ecology of these seven species, combined with the patchy distribution
of their remaining habitat, may hinder their dispersal if relocation
becomes necessary due to climate-influenced changes in distribution of
host plant species (Magnacca 2007, pp. 173, 181-183, 188) and cause the
extirpation of remaining populations of Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H. kuakea, H. longiceps, and H.
mana.
The petitioner states that climate change may also have a
deleterious effect upon the seven Hylaeus bee species due to climate-
induced changes in rainfall patterns, since these species prefer
relatively dry habitats, some of which lack groundwater sources. The
petitioner presents a concern that a predicted rise in sea level in the
Hawaiian Islands (Baker et al. 2006, p. 1) might threaten coastal
strand populations of the seven Hylaeus bee species. The petitions cite
one study that predicted that sea level rise in the Northwestern
Hawaiian Islands will cause a median projected loss of land of 3 to 65
percent with a 19-in (48-cm) sea level rise, and a maximum projected
loss of land of 5 to 75 percent with a 35-in (88-cm) sea level rise
(Baker et al. 2006, p. 1). Although none of the seven Hylaeus bees
occurs on the Northwestern Hawaiian Islands, the petitioner concludes
that sea level rise will also impact the populations of the five
species of Hawaiian yellow-faced bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H. longiceps) inhabiting
coastal sites on the main Hawaiian Islands.
Summary of Factor A
In summary, we find that the information provided in the petitions
presents substantial scientific or commercial information indicating
that the petitioned actions may be warranted due to the present or
threatened destruction, modification, or curtailment of the species'
habitat or range. The petitioner has provided no information, and we
have no information in our files to substantiate the claim that there
will be climate-induced changes in rainfall patterns in the areas where
the seven species occur, or that relatively dry habitats will be
negatively impacted. The petitions did identify numerous potential
factors that may be affecting Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H. longiceps, and H. mana, including
habitat loss and degradation due to urbanization and land conversion;
replacement of native host plants by nonnative plants caused by the
browsing, trampling, and rooting activities of nonnative ungulates,
which facilitates the establishment of nonnative plants in disturbed
areas; conversion by fire of native plant communities to plant
communities dominated by nonnative, fire-adapted plants; and the
removal or trampling of native vegetation by people and compaction of
native vegetation by off-road vehicles in popular recreational areas,
particularly in coastal strand habitat. Information in our files also
indicates these factors may present a threat to the seven species of
Hylaeus. We, therefore, conclude the petitions present substantial
information to indicate that the present or threatened destruction or
modification of habitat may present a threat to H. anthracinus, H.
assimulans, H. facilis, H. hilaris, H. kuakea, H. longiceps, and H.
mana.
B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
According to the petitioner, Hylaeus facilis, H. hilaris, H.
kuakea, and H. mana, each with two or fewer known populations, are
especially vulnerable to overcollection because the collection of even
a few individuals could significantly reduce the production of
offspring (Xerces 2009c, p. 23; 2009d, p. 13; 2009e, p. 11). However,
the petitions also acknowledge that because of the high fecundity of
individual insects, the collection of insects does not pose a threat to
their populations (Xerces 2009c, p. 23; 2009d, p. 11; 2009e, p. 11),
which introduces an element of uncertainty concerning this claim.
Insect collecting is a valuable component of research, including
taxonomic work, and is often necessary for documenting the existence of
populations and population trends. The petitioner has not presented
information with which we can evaluate whether the overcollection of
Hylaeus facilis, H. hilaris, H. kuakea, or H. mana may present a threat
to any of these species, or determine whether this activity has
resulted in population declines. In this regard, neither the petitions,
nor information available in our files, presents information that would
indicate overcollection may present a significant threat to H.
anthracinus, H. assimulans, H. facilis, H. hilaris, H. kuakea, H.
longiceps, or H. mana.
C. Disease or predation
Neither the petitions nor information in our files presents
information that would indicate disease is a current threat to Hylaeus
anthracinus, H. assimulans, H. facilis, H. hilaris, H. kuakea, H.
longiceps, or H. mana.
Predation by Nonnative Ants
Ants are known to prey upon Hylaeus species (Medeiros et al. 1986,
pp. 45-46; Reimer 1994, p. 17), thereby directly eliminating them from
specific areas. The petitions state that ants are not a natural
component of Hawaii's arthropod fauna, and the native species of the
islands evolved in the absence of predation pressure from ants. They
also state that ants can be particularly destructive predators because
of their high densities, recruitment behavior, aggressiveness, and
broad range of diet (Reimer 1993, pp. 17-18). The petitions also state
that the threat of ant predation on the seven Hylaeus bee species is
amplified by the fact that most ant species have winged reproductive
adults (Borror et al. 1989, p. 738) and can quickly establish new
colonies in suitable habitats (Staples and Cowie 2001, p. 55). In
addition, the petitions state that these attributes allow some ants to
destroy otherwise geographically isolated populations of native
arthropods (Nafus 1993, pp. 19, 22-23).
According to the petitions, at least 47 species of ants are known
to be established in the Hawaiian Islands (Hawaii Ants 2008, pp. 1-11).
Native insect fauna, likely including Hylaeus bees (Zimmerman 1948, p.
173; Reimer et al. 1990, pp. 40-43; HEAR database 2005, pp. 1-2), have
been severely impacted by at least four particularly aggressive ant
species: The big-headed ant (Pheidole megacephala), the long-legged ant
(also known as the yellow crazy ant) (Anoplolepis gracilipes),
Solenopsis papuana (no common name), and Solenopsis geminata (no common
name). The petitions state that numerous other species of ants are
recognized as threats to Hawaii's native invertebrates, and an unknown
number of new species of ants are established every few years (Staples
and Cowie 2001, p. 53). The petitions state that due to their
preference for drier habitat sites, ants are more likely to occur in
high densities in the dry and mesic habitat currently occupied by the
seven bees (Xerces 2009a, p. 28; 2009b, p. 19; 2009c, p. 24; 2009d, pp.
13-14; 2009e, pp. 11-12).
The petitions state that the long-legged ant appeared in Hawaii in
1952; now occurs on Kauai, Oahu, Maui, and Hawaii (Reimer et al. 1990,
p. 42); and inhabits low-to-mid-elevation (less than 2,000-ft (600-m))
rocky areas of moderate rainfall (less than 100 in (250 cm) annually)
(Reimer et al. 1990, p. 42). The petitioner also states that direct
observations indicate that Hawaiian arthropods are susceptible to
predation by this species; Gillespie and Reimer
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(1993, p. 21) and Hardy (1979, p. 34) documented the impacts to native
insects within the Kipahulu area on Maui after this area was invaded by
the long-legged ant. The petitioner concludes that although only
cursory observations exist, long-legged ants are thought to be a threat
to populations of the seven Hylaeus bees within dry to mesic areas
within their elevation range (Reimer et al. 1990, p. 42).
Solenopsis papuana is the only abundant, aggressive ant that has
invaded intact mesic to wet forest from sea level to over 2,000 ft (600
m) on all of the main Hawaiian Islands, and is still expanding its
range (Reimer 1993, p. 14). The petitions state that because of this
species' expanding range, it may threaten populations of Hylaeus
facilis in mesic areas up to 2,000 ft (600 m) in elevation (Reimer
1993, p. 14).
The petitions state that the presence of ants in nearly all of the
low-elevation habitat sites historically and currently occupied by the
seven Hylaeus bee species may increase the uncertainty of Hylaeus
recovery within these areas. Hylaeus populations are known to be
drastically reduced in ant-infested areas (Medeiros et al. 1986, pp.
45-46; Stone and