Endangered and Threatened Wildlife and Plants; 12-Month Finding on Five Petitions To List Seven Species of Hawaiian Yellow-faced Bees as Endangered, 55170-55203 [2011-22433]
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FOR FURTHER INFORMATION CONTACT:
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R1–ES–2010–0012; MO
92210–0–008]
Endangered and Threatened Wildlife
and Plants; 12-Month Finding on Five
Petitions To List Seven Species of
Hawaiian Yellow-faced Bees as
Endangered
AGENCY:
Fish and Wildlife Service,
Interior.
Notice of 12-month petition
finding.
ACTION:
We, the U.S. Fish and
Wildlife Service (Service), announce a
12-month finding on five petitions to
list seven species of Hawaiian yellowfaced bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana) as
endangered and to designate critical
habitat under the Endangered Species
Act of 1973, as amended (Act). After
review of all available scientific and
commercial information, we find that
listing these seven species of Hawaiian
yellow-faced bees is warranted.
Currently, however, listing these seven
species of Hawaiian yellow-faced bees is
precluded by higher priority actions to
amend the Lists of Endangered and
Threatened Wildlife and Plants. Upon
publication of this 12-month petition
finding, we will add these seven species
of Hawaiian yellow-faced bees to our
candidate species list. We will develop
a proposed rule to list these seven
species of Hawaiian yellow-faced bees
as our priorities allow. We will make
any determinations on critical habitat
during development of the proposed
listing rule. In any interim period we
will address the status of the candidate
taxa through our annual Candidate
Notice of Review (CNOR).
DATES: The finding announced in this
document was made on September 6,
2011.
ADDRESSES: This finding is available on
the Internet at https://
www.regulations.gov at Docket Number
FWS–R1–ES–2010–0012. Supporting
documentation we used in preparing
this finding is available for public
inspection, by appointment, during
normal business hours at the U.S. Fish
and Wildlife Service, Pacific Islands
Fish and Wildlife Office, 300 Ala Moana
Boulevard, Room 3–122, Honolulu, HI
96850. Please submit any new
information, materials, comments, or
questions concerning this finding to the
above street address.
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SUMMARY:
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Loyal Mehrhoff, Field Supervisor,
Pacific Islands Fish and Wildlife Office
(see ADDRESSES); by telephone at 808–
792–9400; or by facsimile at 808–792–
9581. If you use a telecommunications
device for the deaf (TTD) please call the
Federal Information Relay Service
(FIRS) at 800–877–8339.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Act (16
U.S.C. 1531 et seq.) requires that, for
any petition to revise the Federal Lists
of Endangered and Threatened Wildlife
and Plants that contains substantial
scientific or commercial information
that listing a species may be warranted,
we make a finding within 12 months of
the date of receipt of the petition. In this
finding, we determine whether the
petitioned action is: (a) Not warranted,
(b) warranted, or (c) warranted, but
immediate proposal of a regulation
implementing the petitioned action is
precluded by other pending proposals to
determine whether species are
endangered or threatened, and
expeditious progress is being made to
add or remove qualified species from
the Federal Lists of Endangered and
Threatened Wildlife and Plants. Section
4(b)(3)(C) of the Act requires that we
treat a petition for which the requested
action is found to be warranted but
precluded as though resubmitted on the
date of such finding, that is, requiring a
subsequent finding to be made within
12 months. We must publish these 12month findings in the Federal Register.
Previous Federal Actions
On March 23, 2009, we received five
petitions dated March 23, 2009, from
Scott Hoffman Black, Executive Director
of the Xerces Society, requesting that
seven species of Hawaiian yellow-faced
bees be listed as endangered under the
Act and critical habitat be designated.
Each petition contained information
regarding the species’ taxonomy and
ecology, historical and current
distribution, present status, and current
and potential threats. We acknowledged
the receipt of the petitions in a letter to
Mr. Black, dated May 8, 2009. In that
letter we also stated that issuing an
emergency regulation temporarily
listing the species under section 4(b)(7)
of the Act was not warranted at that
time. We published the 90-day finding
in the Federal Register on June 16, 2010
(75 FR 34077). This notice constitutes
the 12-month finding on the March 23,
2009, petitions to list the seven species
of Hawaiian yellow-faced bees as
endangered.
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Species Information
Overview of the Genus Hylaeus
The seven species of bees described in
this finding 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 endemic to
single islands (Magnacca 2007a, p. 174).
All 60 Hawaiian species are in the
subgenus Nesoprosopis (Magnacca and
Danforth 2006, p. 393). The Hawaiian
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 family
is comprised of over 2,000 species, all
of which are solitary nesting (unlike
social wasps and bees), although a few
do nest in close vicinity to each other.
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).
A great deal of our knowledge on
Hawaiian Hylaeus bees is based upon
surveys by Robert Cyril Layton Perkins,
a distinguished British entomologist and
naturalist renowned for his pioneering
work on the insects of the Hawaiian
Islands, particularly the Hymenoptera
(sawflies, wasps, bees, and ants), in the
early 20th Century. His surveys were
conducted between 1892 and 1906, and
form the basis for most of the historical
records of Hylaeus in the Hawaiian
Islands. According to Perkins, Hylaeus
species were ‘‘almost the most
ubiquitous of any Hawaiian insects’’
(Perkins 1913, p. lxxxi). However, there
are about 90 years between Perkins’
surveys and the most recent surveys
conducted in the late 1990s for Hylaeus
bees in the Hawaiian Islands.
Surveys in more recent years (1998–
2010) for Hylaeus spp. in the Hawaiian
Islands have largely involved targeted
collecting on specific flowering plants
(Daly and Magnacca 2003, pp. 217–233;
Magnacca in litt. 2011, p. 5), rather than
survey methods such as pan trapping or
Pollard walks (see below). While this
means the numbers of individuals and
species observed are not strictly
quantifiable by effort, the probability of
collecting species actually present is
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higher (Magnacca in litt. 2011, p. 5).
Because the number and diversity of
Hylaeus spp. tend to be locally
concentrated rather than widely
distributed, randomized and more
quantifiable surveys such as pan
trapping and Pollard walks are actually
less effective means of locating Hylaeus
spp. (Magnacca in litt. 2011, p. 5). Pan
trapping involves the use of shallow
pans of fluid, and relies on the organism
falling or flying into the fluid
preservative. Pollard walks involve
observers walking along a fixed transect
route and recording the insects
observed.
The recent Hylaeus spp. survey efforts
are not easily comparable to Perkins’
collections, which are considered now
to have been conducted
opportunistically. For example, Perkins
collected higher numbers of individuals
and species in certain areas, including
coastal areas that were much less
disturbed at the time, and some species,
such as H. facilis, were formerly very
common but have almost entirely
disappeared (Magnacca in litt. 2011, p.
5).
Life History of Genus Hylaeus
The following discussion includes all
Hawaiian Hylaeus species, and specific
information about the seven petitioned
Hylaeus species.
Hawaiian Hylaeus species are
grouped within two categories: Groundnesting species that require relatively
dry conditions, and wood-nesting
species that are often found within
wetter areas (Zimmerman 1972, p. 533;
Daly and Magnacca 2003, p. 11). Nests
of Hylaeus species are usually
constructed opportunistically within
dead twigs or plant stems, or other
similarly small natural cavities under
bark or rocks (i.e., they seek out existing
cavities that they suit to their own
needs). This is unlike the nests of many
other bee species, which are
purposefully excavated or constructed
underground. Like other Hylaeus,
Hawaiian Hylaeus 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 female
Hylaeus bee lays eggs in brood cells 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 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
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).
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. 2,524–2,528; Cox
and Elmqvist 2000, p. 1,238; Sahli et al.
2008, p. 1) have demonstrated Hawaiian
Hylaeus species almost exclusively visit
native plants to collect nectar and
pollen, pollinating those plants in the
process. Hylaeus bees are very rarely
found visiting nonnative plants for
nectar and pollen (Magnacca 2007a, 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 nonnative
species changes with elevation. In that
study, eight flowering plants were
observed at six sites, which ranged in
elevation from approximately 2,900 to
7,900 feet (ft) (approximately 880 to
2,400 meters (m)). The study also found
the proportion of native pollinators
changed along the elevation gradient; at
least 40 to 50 percent of visits were from
nonnative pollinators at low elevation,
as opposed to 4 to 20 percent of visits
by nonnative 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 suggest Hylaeus may
not be easily replaceable by nonnative
pollinators (Sahli et al. 2008, p. 1).
For some of the seven Hawaiian
yellow-faced bees addressed in this
finding, we have information about the
specific host plants they visit for nectar
and pollen. For some species, we have
also identified primary host plants
visited (see description of the species
where noted). However, for others, we
lack detailed information on the specific
host plants visited for foraging.
Nonetheless, researchers believe native
plants both endemic and indigenous to
the Hawaiian Islands are essential to the
survival of the Hylaeus species (Hopper
et al. 1996, pp. 8–9; Daly and Magnacca
2003, pp. 217–229; Magnacca 2007a, pp.
185–186).
Hawaiian Island Ecosystems
The five Hawaiian Island ecosystems
that support the seven Hawaiian yellowfaced bees addressed in this 12-month
finding are described in the following
section. See Table 1 below for a list of
the ecosystems from which each species
is reported. Because Hawaiian Hylaeus
spp., including these seven, are believed
to be essential pollinators of the native
Hawaiian plant fauna, we are providing
this background information on the
different ecosystems in which they
occur to better elaborate upon the
specific threats found in the five
ecosystem types.
TABLE 1—CURRENT (AND HISTORICAL) DISTRIBUTION OF THE SEVEN YELLOW–FACED BEES BY ECOSYSTEM TYPE AND
ISLAND
Species and number of current
populations
Ecosystems
Coastal
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H. anthracinus, 13 populations
HI, MA, MO,
OA.
H. assimulans, 5 populations ...
KAH, (*MA),
(*OA).
(*MA), MO,
(*OA).
(*LA), (*MA),
MO.
N/A ..................
LA, MA, MO,
OA.
H. facilis, 2 populations ............
H. hilaris, 1 population ..............
H. kuakea, 2 populations ..........
H. longiceps, 6 populations ......
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Lowland dry
Lowland mesic
Lowland wet
Montane mesic
HI, KAH, (*LA),
MA, (*MO),
(*OA).
LA, MA, (*OA)
N/A ..................
N/A ..................
............................
HI.
N/A ..................
N/A ..................
............................
N/A.
(*LA), (*OA) .....
(*LA), (*MA),
(*OA).
N/A ..................
(*MA), OA .......
(*MO) .................
N/A.
N/A ..................
............................
N/A.
OA ...................
N/A ..................
N/A ..................
N/A ..................
............................
............................
N/A.
N/A.
(*MA) ...............
N/A ..................
LA, (*MA),
(*MO).
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TABLE 1—CURRENT (AND HISTORICAL) DISTRIBUTION OF THE SEVEN YELLOW–FACED BEES BY ECOSYSTEM TYPE AND
ISLAND—Continued
Ecosystems
Species and number of current
populations
Coastal
Lowland dry
Lowland mesic
Lowland wet
Montane mesic
H. mana, 1 population ..............
N/A ..................
N/A ..................
OA ...................
N/A ..................
............................
Montane dry
N/A.
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HI = Hawaii (Island); KAH = Kahoolawe; LA = Lanai; MA = Maui; MO = Molokai; OA = Oahu;
(*XX) denotes a historical population; N/A means no population records
Coastal Ecosystem
The coastal ecosystem is found on all
of the main Hawaiian Islands, with the
highest species diversity found in the
least populated coastal areas of Hawaii,
Maui, Molokai, Kahoolawe, Oahu, and
Kauai, and their associated islets, and
extends from sea level to approximately
1,000 ft (approximately 300 m) in
elevation. The coastal vegetation zone is
typically dry, with annual rainfall of
less than approximately 20 inches (in)
(50.8 centimeters (cm)); however
windward rainfall may be high enough
(up to approximately 40 in (1,000 mm))
to support mesic-associated and
sometimes wet-associated vegetation
(Gagne and Cuddihy 1999, pp. 54–66).
Compared to dry and mesic ecosystems,
biological diversity (number of species)
is low to moderate in the coastal
ecosystem, but may include some
specialized plants and animals such as
nesting seabirds and the rare native
plant Sesbania tomentosa (ohai) (The
Nature Conservancy (TNC) 2006a).
Sesbania tomentosa formerly occurred
widely in lower elevation dry habitat on
all of the main islands and at least on
Necker and Nihoa of the Northwestern
Hawaiian Islands. The species is now
scattered throughout its former range,
and is restricted to relic populations on
sandy beaches, on dunes, on soil
pockets on lava, and along pond
margins (Wagner et al. 1990, p. 705).
The dominant native vegetation in
coastal ecosystems is the shrub Scaevola
sericea (naupaka kahakai) (Alpha et al.
1996, p. 86). Other common native plant
species include Ipomoea pes-caprae
(beach morning-glory), Sporobolus
virginicus (beach dropseed),
Jacquemontia ovata (pau o Hiiaka), and
Sesuvium portulacastrum (akulikuli or
sea purslane) (Wagner et al. 1999, pp.
57–59). Among the Hylaeus species
addressed in this finding, five are
known from coastal ecosystems,
including H. anthracinus, H.
assimulans, H. facilis, H. hilaris, and H.
longiceps.
Lowland Dry Ecosystem
The lowland dry ecosystem includes
shrublands and forests below
approximately 3,300 ft (1,000 m) in
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elevation that receive less than 50 in
(127 cm) annual rainfall, or are in
otherwise prevailingly dry substrate
conditions. Areas consisting of
predominantly native species in the
lowland dry ecosystem are now rare.
This ecosystem is found on the islands
of Hawaii, Maui, Molokai, Lanai,
Kahoolawe, Oahu, and Kauai, and is
best represented on the leeward sides of
the islands (Gagne and Cuddihy 1999, p.
67). Biological diversity is low to
moderate in this ecosystem, and
includes specialized animals and plants
such as the Hawaiian owl or pueo (Asio
flammeus sandwichensis) and Santalum
ellipticum (iliahialoe) (Wagner et al.
1999, pp. 1,220–1,221; TNC 2006b).
Hylaeus anthracinus, H. assimulans,
H. facilis, and H. longiceps are known
from lowland dry forests. These forests
are typically dominated by Diospyros
sandwicensis (lama), Erythrina
sandwicensis (wiliwili), Nestegis
sandwicensis (olopua), or Metrosideros
polymorpha (ohia) and a diversity of
native shrubs growing within the
understory (Gagne and Cuddihy 1999,
pp. 72–74).
Lowland Mesic Ecosystem
The lowland mesic ecosystem
includes a variety of grasslands,
shrublands, and forests, below
approximately 3,300 ft (1,000 m) in
elevation, that receive between 50 and
75 in (127 and 191 cm) annual rainfall,
or are in otherwise mesic substrate
conditions (TNC 2006c). In the
Hawaiian Islands, this ecosystem is
found on Hawaii, Maui, Molokai, Lanai,
Oahu, and Kauai, on both windward
and leeward sides of the islands.
Biological diversity is high in this
system (TNC 2006c).
Lowland mesic forests are typically
dominated by Acacia koa (koa),
Diospyros sandwicensis, Metrosideros
polymorpha, or Nestegis sandwicensis,
and a diversity of understory trees and
native shrubs growing below the canopy
species (Gagne and Cuddihy 1999, p.
80–82). Historically, Hylaeus facilis was
known from lowland mesic forest, but
currently only H. kuakea and H. mana
are found in this habitat.
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Lowland Wet Ecosystem
The lowland wet ecosystem is
generally found below approximately
3,300 ft (1,000 m) in elevation on the
windward sides of the main Hawaiian
Islands, except Kahoolawe (Gagne and
Cuddihy 1999, p. 85; TNC 2006d).
These areas include a variety of wet
grasslands, shrublands, and forests that
receive greater than 75 in (191 cm)
annual precipitation, or are in otherwise
wet substrate conditions (TNC 2006d).
Biological diversity is high in this
system (TNC 2006d). The majority of
lowland wet forests are dominated by
Metrosideros polymorpha, with
understory trees such as Psychotria spp.
(kopiko) and Antidesma platyphyllum
(hame) (Gagne and Cuddihy 1999, p.
87). Currently, Hylaeus facilis is known
from lowland wet forest (Daly and
Magnacca 2003, p. 81).
Montane Dry Ecosystem
The montane dry ecosystem is
composed of natural communities
(shrublands, grasslands, forest) found at
elevations between approximately 3,300
and 6,600 ft (1,000 and 2,000 m), in
areas where annual precipitation is less
than 50 in (127 cm), or otherwise in dry
substrate conditions (TNC 2006g).
Montane dry forests occur on the
leeward sides of the islands of Maui and
Hawaii, and biological diversity is
moderate (Gagne and Cuddihy 1999, p.
93; TNC 2006g). Montane dry forests are
dominated by some combination of
Acacia koa, Sophora chrysophylla)
(mamame), Metrosideros polymorpha,
and rarely, Chamaesyce olowaluana
(akoko) (Gagne and Cuddihy, p. 95). In
2004, a single individual of H.
anthracinus was collected in montane
dry forest on Hawaii Island.
Specific Information on Hylaeus
anthracinus
Taxonomy and Description
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). Nesoprosopis was
reduced to a subgenus of Hylaeus in
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1923 (Meade-Waldo 1923, p. 1).
Although the distinctness of this species
remains unquestioned, recent genetic
evidence (Magnacca and Brown 2010,
pp. 5–7) suggests H. anthracinus may be
composed of three cryptic (not
recognized) species or subspecies that
represent the populations on Hawaii,
Maui and Kahoolawe, and Molokai and
Oahu. However, this has not been
established scientifically; therefore, we
treat H. anthracinus as a single species
in this finding.
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 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, Scaevola sericea,
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). Tournefortia argentea was first
collected on Oahu in 1864–1865, and is
naturalized and documented from all of
the main islands except Kahoolawe
(Wagner et al. 1999, p. 398). 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 and
lowland dry forest habitats up to 2,000
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ft (610 m) in elevation on the islands of
Hawaii, Lanai, Maui, Molokai, and
Oahu. Between 1997 and 2008, surveys
for Hawaiian Hylaeus were conducted at
43 sites throughout the Hawaiian
Islands that were either historical
collecting localities for H. anthracinus,
or potentially suitable habitat for this
species. Hylaeus anthracinus was
observed at 13 of the 43 survey sites, but
had disappeared from each of the 9
historically occupied sites surveyed
(Daly and Magnacca 2003, p. 217;
Magnacca 2007b, p. 44). 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 Magnacca 2003, p. 55;
Magnacca 2007a, pp. 186–188).
Hylaeus anthracinus is currently
known from 13 small patches of coastal
and lowland dry forest habitat
(Magnacca 2005a, p. 2): five locations on
the island of Hawaii; one location on
Kahoolawe; two locations on Maui;
three locations on Molokai; and two
locations on Oahu (Daly and Magnacca
2003, p. 217; Magnacca 2005a, p. 2;
Magnacca 2007b, p. 44). These 13
locations supported small populations
of H. anthracinus, but the number of
individual bees is unknown. In 2004, a
single individual was collected in
montane dry forest on the island of
Hawaii; however, the presence of
additional individuals has not been
confirmed at this site (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). The species is
believed to be extirpated from Lanai
(Daly and Magnacca 2003, p. 55).
Additionally, during surveys between
1997 and 2008, H. anthracinus was
absent from 17 other sites on Hawaii,
Maui, Molokai, and Oahu with
potentially suitable habitat from which
other species of Hylaeus were collected
(Daly and Magnacca 2003; Magnacca,
University of Hawaii at Hilo, pers.
comm. 2008a).
Hawaii Island
Hylaeus anthracinus was first
described by Perkins (1899, p. 100) from
specimens collected by F. Smith on the
Kona (west) coast at Kealakekua Bay. In
the intervening 99 years, H. anthracinus
appears to have declined significantly
throughout its historical range on this
coastline. Between 1997 and 2008,
researchers thoroughly surveyed the
area around Kealakekua Bay and Keei to
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the south, but found no species of
Hylaeus and observed that most of these
areas are either dominated by invasive,
nonnative plants, such as Leucaena
leucephala (koa haole), or lack
vegetation entirely (Magnacca, pers.
comm. 2008a). Hylaeus anthracinus is
currently found in five locations in
coastal and lowland dry forest on the
leeward (west) side of the island,
including Kohanaiki; KalokoHonokohau National Historic Park
(NHP); Makalawena Beach; the
Mahaiula section of Kekaha Kai (Kona
Coast) State Park; and Kaulana Bay near
Ka Lae (South Point). In addition, there
is one recent collection from montane
dry forest in the U.S. Army’s Pohakuloa
Training Area, in the northern part of
the island. Collection reports from these
six areas follow:
(A) Kohanaiki: Hylaeus anthracinus
was collected in coastal habitat on
Tournefortia argentea at this location
near Puhili Point by Magnacca (2007b,
p. 44). Kohanaiki is an area of land
granted to indigenous Hawaiians in
1995 for cultural and recreational
preservation and pursuits (Kohanaiki
Ohana 1995 (https://www.kohanaiki.org/
)). There is some possibility for
increased recreational impact to the
area, if and when adjacent privately
owned parcels are developed, as is
currently planned (Kohanaiki Ohana
1995 (https://www.kohanaiki.org/)).
(B) Kaloko-Honokohau NHP: In 2007,
researchers collected Hylaeus
anthracinus in coastal habitat in
Kaloko-Honokohau NHP, which is just
south of Kohanaiki, and managed by the
National Park Service (NPS) (P. Aldrich,
University of Hawaii at Manoa, pers.
comm. 2008a; Magnacca, pers. comm.
2008c).
(C) Makalawena Beach: Researchers
collected Hylaeus anthracinus in coastal
habitat in south Kona at Makalawena
Beach in 2007 (P. Aldrich, pers. comm.,
July 2008a). Inaccessible by motor
vehicle, visitors must hike to the beach
on a trail that begins in nearby Kekaha
Kai State Park. Makalawena Beach is
located on private land owned by
Kamehameha Schools.
(D) Mahaiula Section of Kekaha Kai
State Park: Researchers collected
Hylaeus anthracinus in coastal habitat
in the Mahaiula section of Kekaha Kai
State Park in 2007 (P. Aldrich,
unpublished data). The park is managed
by the Hawaii Department of Land and
Natural Resources’ (DLNR) Division of
State Parks, and is open to the public
daily. This section of the park is
accessed by a 1.5-mile (mi) (1.6kilometer (km)) unpaved road from the
main highway (Queen Kaahumanu
Highway (Hwy 19)), and offers public
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recreational opportunities for swimming
and beach-related activities, such as
hiking, picnicking, and boating (https://
www.hawaiistateparks.org/
hawaiistateparks/parks/hawaii/
index.cfm?park_id=47).
(E) Kaulana Bay: Hylaeus anthracinus
appears to be restricted to an area of
5,000–10,000 year-old lava flows east of
Ka Lae at Kaulana Bay, where it and
other species of Hylaeus were collected
in 1999 and 2002 (Magnacca 2007a, p.
181). The substrate of these lava flows
is distinct from the surrounding areas
covered by Pahala ash (Magnacca, pers.
comm. 2010b). The area near Ka Lae, at
the southernmost tip of the island of
Hawaii, is believed to be the best coastal
habitat for Hylaeus on the island.
However, H. anthracinus was absent
from several sites with potentially
suitable vegetation near Ka Lae and
other sites to the east along the coast,
including Kalu, Kaalualu, and Mahana,
where other Hylaeus species were
collected. The population of H.
anthracinus at Kaulana Bay appears
highly localized, and may have more
stringent habitat requirements related to
localized substrate type than other
species of Hawaiian Hylaeus found in
nearby areas (e.g., H. difficilis and H.
flavipes). The Ka Lae area, including
Kaulana Bay, is registered as a National
Historic Landmark District and a large
portion of the area is primarily owned
by the State’s Department of Hawaiian
Home Lands (DHHL), although a smaller
portion is privately owned. Public
access to Kaulana Bay is not restricted,
and the area is used for recreational
activities such as off-road vehicle use
(Magnacca, pers. comm. 2008a).
(F) U.S. Army’s Pohakuloa Training
Area (PTA): In 2004, one male Hylaeus
anthracinus was collected on the
southern slopes of Mauna Kea in
montane dry forest habitat in the U.S.
Army’s PTA at approximately 5,200–
5,400 ft (1,590–1,650 m) in elevation
(Magnacca 2007b, p. 44). The specimen
was found inside the fruit capsule of the
federally endangered plant, Hedyotis
coriacea (kioele). Hylaeus anthracinus
has not been observed at the PTA since
the collection made in 2004 (Magnacca
2007b, p. 44). It is unknown if this
collection was a single vagrant
individual or from an established
population at the PTA (Magnacca
2007b, p. 44).
Kahoolawe Island
Previously unknown on Kahoolawe, a
population of Hylaeus anthracinus was
discovered in 2002 in coastal habitat at
Pali o Kalapakea, where four specimens
were collected at an elevation of 1,000
ft (300 m) (Daly and Magnacca 2003;
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Magnacca, pers. comm. 2008a).
However, this species was absent from
potentially suitable habitat located at
Kamohio on the southeastern coast of
the island where other Hylaeus species
were collected. Overgrazing by
introduced cattle and goats, and
bombing and target practice by the U.S.
military, have led to soil erosion
resulting in the loss of almost all of the
coastal and lowland dry forest habitat
on this island (Warren 2004, p. 461). In
1993, Congress ended military use on
Kahoolawe, and the Kahoolawe Island
Reserve Commission (KIRC) was created
to manage land use and restore
Kahoolawe’s natural resources (Dept. of
Defense, p. 1). Access to the island is
limited and controlled by KIRC, and
activities conducted on the island
include fishing, habitat restoration,
historical preservation, and education.
Commercial enterprises are currently
prohibited on the island (Warren 2004,
p. 1).
(B) Kanaio Natural Area Reserve:
Hylaeus anthracinus was collected in
1999 in remnant native lowland dry
forest in the State’s Kanaio Natural Area
Reserve (NAR) on the southern slopes of
Haleakala at 2,000 ft (600 m) in
elevation (Daly and Magnacca 2003, p.
217). Kanaio NAR is a State-protected
area of approximately 876 ac (355 ha),
and contains patches of lowland dry
forest and shrub lands. The State plans
to rehabilitate habitat in the Kanaio
NAR by excluding feral ungulates with
fencing, managing weeds, and planting
native species (https://hawaii.gov/dlnr/
dofaw/rpc/projects-on-maui).
(C) Manawainui Gulch: In 1999,
Hylaeus anthracinus was collected at
this coastal site on land owned by the
State’s DHHL (Magnacca, pers. comm.
2008a). The site is east of Kahikinui,
and should not be confused with the
Manawainui Valley, which is east of
Kaupo, or Manawainui Gulch at
Ukumehame on west Maui.
Maui
Perkins (1899, p. 100) originally
described Hylaeus anthracinus as
abundant in coastal and lowland habitat
on the island of Maui, where it was
known from four sites. Perkins’ primary
collection site for coastal bees on Maui
was the Wailuku sandhills, which once
supported a diverse bee fauna. Lacking
adequate descriptions, researchers were
unable to relocate two of the Perkins
collection sites during recent surveys,
but two sites were relocated and
surveyed in 1999 and 2001 (Magnacca
2007a, p. 173). Hylaeus anthracinus has
also been collected at Kanaio on the
lower southern slopes of Haleakala, an
unusual location for this otherwise
exclusively coastal species. The species
was also collected at the coast nearby,
at Manawainui. Descriptions of these
three sites follows:
(A) Wailuku Sand Hills: Formerly a
large expanse of coastal dune habitat,
the Wailuku sand hills remain as small
remnant dunes and only one, at Waiehu,
contains intact native vegetation
potentially suitable for Hylaeus bees.
This remnant coastal sand dune covers
less than 2.5 acres (ac) (1 hectare (ha))
on State lands near a golf course.
Hylaeus anthracinus was not observed
during the 1999 and 2001 surveys in
this location (Daly and Magnacca 2003,
p. 217). The rest of the dunes have been
destroyed by development or are
overgrown with the nonnative plant
Prosopis pallida (kiawe). Researchers
observed that the Kahului section of the
dunes, located south of the native
remnant dune, no longer contains
potentially suitable habitat for species of
Hylaeus (Magnacca 2007a, p. 182).
Molokai
Perkins collected Hylaeus
anthracinus at Kaulawai [Kauluwai] and
two unknown sites: the lower slopes of
the north Molokai mountains and the
‘‘Molokai plains’’ (Perkins 1899; Daly
and Magnacca 2003, p. 55). Hylaeus
anthracinus occurred in three of five
sites surveyed between 1999 and 2005.
These locations include TNC’s
Moomomi Preserve on Molokai’s
northwest coast, and Hoolehua Beach
and Kaupikiawa, both located on the
Kalaupapa peninsula (Magnacca, pers.
comm. 2008a). This species was not
observed at several other sites with
potentially suitable habitat, including
sand dune habitat near the Kaluakoi
resort on Molokai’s west coast
(Magnacca, pers. comm. 2008a).
Collection reports of these sites follow:
(A) Moomomi Preserve: Between 1999
and 2001, researchers collected H.
anthracinus and H. longiceps from an
area of native vegetation in coastal dune
habitat within Moomomi Preserve
(Magnacca 2007a, p. 181). Moomomi
Preserve contains intact coastal dunes
dominated by native vegetation, as well
as dune and inland areas dominated by
nonnative vegetation.
(B) Hoolehua Beach and Kaupikiawa:
In 2005, Hylaeus anthracinus was
collected at a coastal site above
Hoolehua Beach near the tip of the
Kalaupapa peninsula, and at
Kaupikiawa, just to the east (Magnacca
2007b, p. 181). Both sites are located
within Kalaupapa NHP, which is
cooperatively managed by the NPS,
DHHL, and the State’s DLNR and
Departments of Health (DOH) and
Transportation (DOT). The areas on the
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east side of the Kalaupapa peninsula are
largely rocky and devoid of vegetation,
but contain scattered patches of native
coastal vegetation, similar to Ka Lae on
the island of Hawaii (Magnacca 2007a,
p. 181).
Oahu
Hylaeus anthracinus was historically
known from seven sites on the island of
Oahu, although two of the coastal sites
were not conclusively identified by
Perkins and the exact locations cannot
now be determined (Perkins 1899, p.
100). This species appears to have
declined precipitously since Perkins’
collecting period on Oahu (1892–1906)
and is currently only known from two
sites, Kaena Point NAR and Mokuauia
(Goat Island). Between 1997 and 2008,
H. anthracinus was not found during
surveys of five of its historical Perkinsera collection sites. Several of these sites
no longer provide suitable habitat for
Hylaeus species because native
vegetation has been removed during
urbanization, or the sites are dominated
by invasive, nonnative vegetation. These
sites include Honolulu, Waikiki, ‘‘the
Honolulu mountains,’’ Waialua, and the
Waianae coast (Liebherr and Polhemus
1997, pp. 345–347; Daly and Magnacca
2003, p. 55). Between 1999 and 2002,
researchers searched coastal habitat at
Makapuu and Kalaeloa (Barber’s Point),
but did not find any species of Hylaeus
(Magnacca, pers. comm. 2008a). The
coastal habitat at both sites is degraded
and dominated by nonnative vegetation.
Descriptions of the two known sites
follow:
(A) Kaena Point NAR: Between 1998
and 2008, Hylaeus anthracinus was
collected at Kaena Point, which is
located on Oahu’s northwest-most point
(Daly and Magnacca 2003, p. 55; Sahli,
University of Hawaii at Manoa, pers.
comm. 2008). Kaena Point contains the
best intact native coastal habitat on
Oahu, and is an excellent example of
that type of ecosystem in the
Northwestern Hawaiian Islands. It
provides habitat for nesting seabirds,
monk seals, native plants, and other
native species (Magnacca 2007a, p. 181).
The primary activities within this NAR
include recreation, hiking, nature study,
education, and the observation of
wildlife (DLNR 2007, p. 20). While
illegal off-road driving was once a
concern, a physical barrier is now in
place that prevents vehicular access,
and native vegetation is regenerating
and being restored by the Kaena Point
Ecosystem Restoration Project (DLNR
2007, p. 20; Magnacca 2007a, p. 181). In
partnership with several agencies
including the Service, the DLNR is
building a predator-proof fence to
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prevent nonnative species, such as cats
and dogs that threaten nesting seabirds,
from entering 59 ac (24 ha) of coastal
habitat within Kaena Point NAR
(https://www.state.hi.us/dlnr/dofaw/
kaena/index.htm).
(B) Mokuauia (Goat Island): From the
lack of records, it appears Perkins and
other early naturalists did not search
Mokuauia or Oahu’s other offshore islets
for yellow-faced bees. Recently, Hylaeus
anthracinus was found on this islet by
Service biologists during general
surveys of the islet (S. Plentovich,
Service, pers. comm. 2008). Mokuauia,
an offshore islet in Laie Bay located on
Oahu’s northeast coast, encompasses 13
ac (5.3 ha) and reaches a maximum
elevation of 15 ft (4.5 m). The entire
islet is a State Seabird Sanctuary and is
managed by the State’s Department of
Forestry and Wildlife (DOFAW). The
entire islet was designated as critical
habitat for the endangered plant
Sesbania tomentosa in 2003, and the
DOFAW is actively restoring native
vegetation and controlling nonnative
species. Mokuauia is easily accessed by
the public and is a popular destination
for small boats, kayaks, and swimmers
on weekends.
Lanai
Hylaeus anthracinus has not been
observed on Lanai for over 100 years
and is likely extirpated from this
privately owned island. This species
was not observed at any of the recently
surveyed sites, including Manele Bay,
where it was collected by Perkins in
1899 (Magnacca 2007a, p. 182;
Magnacca, pers. comm. 2008a).
However, other Hylaeus species were
collected at seven of the eight locations
surveyed (Daly and Magnacca 2003, pp.
217–229).
Summary of Hylaeus anthracinus Range
and Distribution
Hylaeus anthracinus was historically
known from numerous coastal and
lowland dry forest habitats up to 2,000
ft (600 m) in elevation, on the islands of
Hawaii, Lanai, Maui, Molokai, and
Oahu. Currently, this species is known
from a total of 13 sites in a few small
patches of coastal and lowland dry
forest habitat: one location on
Kahoolawe, five locations on the island
of Hawaii, two locations on Maui, three
locations on Molokai, and two locations
on Oahu. In addition, in 2004 a single
individual of H. anthracinus was
collected in montane dry forest habitat
on the island of Hawaii. It is unknown
if this collection was a single vagrant
individual or from an established
population. The lands on which H.
anthracinus occurs are under a variety
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of jurisdictions, including private (e.g.,
TNC), State (e.g., DHHL, DOFAW,
NARs, State Park, Seabird Sanctuary),
and Federal (U.S. Army, NPS).
Specific Information on Hylaeus
assimulans
Taxonomy and Description
Hylaeus assimulans was first
described as Nesoprosopis assimulans
(Perkins 1899, pp. 75, 101–102);
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). 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 (Magnacca 2005b, p. 2). Likewise,
the nesting habits of H. assimulans are
not known, but because the species is
genetically related to other ground
nesting Hylaeus spp., it is thought to be
a ground nester (Magnacca 2005b, p. 2).
Hylaeus assimulans adults have been
observed visiting the flowers of
Lipochaeta lobata (nehe) and its likely
primary host plant, Sida fallax (Daly
and Magnacca 2003, p. 58). Hylaeus
assimulans appears to be closely
associated with plants in the genus
Sida, and studies thus far suggest this
yellow-faced bee species may be more
common where this plant is abundant
(Daly and Magnacca 2003, pp. 58, 217;
Magnacca 2007a, p. 183). In recent
survey efforts, H. assimulans seems to
be more common in dry forest at
relatively higher elevations, which may
be related to the abundance of Sida in
the understory (Magnacca 2005b, p. 2).
Sida spp. were less often found in
coastal habitat. It is likely H. assimulans
visits several other native plants,
including Acacia koa, Metrosideros
polymorpha, Styphelia tameiameiae
(pukiawe), and species of Scaevola
(naupaka) and Chamaesyce (akoko),
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which are frequented by other Hylaeus
species as well (Magnacca, pers. comm.
2008b).
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Range and Distribution
Historically, Hylaeus assimulans was
known from numerous coastal and
lowland dry forest habitats up to 2,000
ft (610 m) in elevation on the islands of
Lanai, Maui, and Oahu. There are no
collections from Molokai, although it is
likely H. assimulans also occurred there
because all other species of Hylaeus
known from Maui, Lanai, and Oahu also
occurred on Molokai (Daly and
Magnacca 2003, pp. 217–229). Between
1997 and 2008, surveys for Hawaiian
Hylaeus were conducted in 25 sites on
Kahoolawe, Lanai, Maui, Molokai, and
Oahu. Hylaeus assimulans was absent
from six of its historical localities on
Lanai, Maui, and Oahu (Xerces Society
2009b, p. 4). Hylaeus assimulans was
not observed at 19 other sites with
potentially suitable habitat on Lanai,
Maui, Molokai, and Oahu, including
several sites from which other native
Hylaeus species have been recently
collected (Daly and Magnacca 2003, pp.
56, 217; Magnacca 2005b, p. 2;
Magnacca 2007a, pp. 177, 181, 183).
Currently, Hylaeus assimulans is
known from a few small patches of
coastal and lowland dry forest habitat at
one location on Kahoolawe, two
locations on Lanai, and two locations on
Maui (Daly and Magnacca 2003, p. 58;
Magnacca 2005, p. 2). This species has
likely been extirpated from Oahu
because it has not been observed since
Perkins’ 1899 surveys and was not
found during recent surveys of
potentially suitable coastal habitat at
Kaena Point, Makapuu, and Kalaeloa
(Daly and Magnacca 2003, p. 217;
Magnacca 2005, p. 2; H. Sahli,
unpublished data).
Kahoolawe
Although not historically known from
Kahaoolawe (Daly and Magnacca 2003,
Magnacca, pers. comm. 2008a), Hylaeus
assimulans was discovered in 1997 near
the high cliffs of Kamohio Bay in the
center of the southern coast of the island
(Daly and Magnacca 2003, p 217). The
species was absent from one other site
on the island in lowland habitat on the
east coast at Pali o Kalapakea where
other Hylaeus species were collected
(Daly and Magnacca 2003, pp. 217–229).
Lanai
On Lanai, Perkins found Hylaeus
assimulans in low numbers within
uninhabited coastal habitat at Awalua in
northwest Lanai, and in the Koele
mountains at an elevation of 2,000 ft
(610 m) (Perkins 1899, p. 102). Between
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1998 and 2006, seven sites with
potentially suitable habitat on private
lands, including Mt. Koele and Awalua,
were surveyed, and H. assimulans was
found only near Manele Road and
Polihua Road in small pockets of native
vegetation (Magnacca, pers. comm.
2008b). Descriptions of these sites
follow:
(A) Manele Road: In 1999, Hylaeus
assimulans was collected in lowland
dry forest along Manele Road at 600 ft
(180 m) in elevation, north of Manele
Beach in southern Lanai (Daly and
Magnacca 2003, p. 217). Researchers
observed the canopy was dominated by
invasive Prosopis pallida trees and the
understory had a dense stand of Sida
fallax, the likely primary host plant of
H. assimulans (Magnacca, pers. comm.
2008b). However, with the exception of
a few stunted plants at the roadside
where moisture had accumulated, the
rest of the stand of Sida fallax had
senesced (reached maturity) or possibly
died. Native plants at this site appeared
to be drought-intolerant and probably
did not provide consistent habitat for
Hylaeus throughout the year (Magnacca
2007a, p. 183; Magnacca, pers. comm.
2008a).
(B) Polihua Road: In 1999, two
specimens of H. assimulans were
collected in lowland dry forest along
Polihua Road at 1,000 ft in elevation
(300 m) in central Lanai (Daly and
Magnacca 2003, p. 58). Both sites are on
private land, and we are unaware of any
recent or current land management in
these areas.
Maui
Perkins collected Hylaeus assimulans
from coastal habitat at the Wailuku sand
hills, and from an unknown site labeled
‘‘Maui’’ (Daly and Magnacca 2003, p.
58). Although other rare Hylaeus spp.
were collected from the Waiehu dunes
area during surveys conducted in 1999
and 2001, H. assimulans, as well as
several other species once collected
there by Perkins, were not found (Daly
and Magnacca 2003, pp. 217–229;
Magnacca, pers. comm. 2008a). Between
1998 and 2006, researchers surveyed six
potentially suitable habitat locations
island-wide, and H. assimulans was
found within small pockets of native
plants in only two of these sites (Daly
and Magnacca 2003, p. 217; Magnacca,
pers. comm. 2008a). However,
researchers believe H. assimulans may
exist in potentially suitable habitat in
rugged and inaccessible portions of west
Maui (Magnacca, in litt., 2010, p. 1).
Descriptions of these two sites follow:
(A) Lahainaluna: In 1999, Hylaeus
assimulans was collected in dry
lowland forest at 1,800 ft (550 m) in
elevation on the west side of Maui. The
site is with the State’s West Maui NAR.
Established in 1986, the NAR’s
management plan calls for the control
and removal of feral ungulates, and the
control of selected priority invasive
plant species (https://hawaii.gov/dlnr/
dofaw/nars/reserves/maui/west-maui).
(B) Waikapu: In 2000, researchers
collected Hylaeus assimulans in
lowland dry shrubland dominated by
the native shrub, Dodonaea viscosa
(aalii) at 400 ft (120 m) elevation in
Waikapu Valley, which is south of Iao
Valley on the east side of west Maui
(Daly and Magnacca 2003, p. 217). The
10,000-square ft (.09-square-ha) site is
privately owned and surrounded by a
fence to exclude nonnative axis deer
(Axis axis). The fence was built in the
mid-1980s by the Native Hawaiian Plant
Society, and is currently managed by
inspecting the fence for breaks;
removing nonnative, invasive weeds;
and collecting seeds of native plants for
propagation. There have been two major
fires in the past 5 years in the vicinity
of the fenced area, although neither fire
has burned within the enclosed area (H.
Oppenheimer, Plant Extinction
Prevention Program, pers. comm. 2008).
Between 1997 and 2007, Hylaeus
assimulans was not collected during
surveys of potentially suitable habitat at
other locations on Maui where other
rare Hylaeus species were collected,
including lowland dry forest habitat in
Kanaio NAR and coastal habitat at
Manawainui Gulch (Daly and Magnacca
2003, pp. 217–229; Magnacca, pers.
comm. 2008a).
Oahu
Perkins found Hylaeus assimulans to
be widespread but not relatively
abundant on Oahu (Magnacca 2005b, p.
2). His Oahu collection sites included
Honolulu (Magnacca, pers. comm.
2008a), the Kaala mountains, the
Waianae Mountains, and the Waianae
coast (Perkins 1899, p. 102; Daly and
Magnacca 2003, p. 58). There are also
specimens collected by Perkins from
unknown locations labeled ‘‘Oahu’’ and
‘‘w. coast, near sea level’’ (Daly and
Magnacca 2003, p. 58).
Hylaeus assimulans was not found
during surveys conducted between 1998
and 2008, including surveys at one
historical location (Daly and Magnacca
2003, pp. 58, 217). Although H.
anthracinus was recently found on
Mokuania (see Hylaeus anthracinus
Range and Distribution), H. assimulans
was not found during surveys of
potentially suitable habitat on this offshore islet (S. Plentovich, Service, pers.
comm. 2008). The absence of H.
assimulans from potentially suitable
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coastal habitat on Oahu suggests it has
likely been extirpated from this island
(Daly and Magnacca 2003, p. 58; H.
Sahli, unpublished data).
Summary of Hylaeus assimulans Range
and Distribution
Hylaeus assimulans was historically
known from numerous coastal and
lowland dry habitats up to 2,000 ft (610
m) in elevation, on the islands of Lanai,
Maui, and Oahu. Currently, this species
is found in a few small patches of
coastal and lowland dry forest habitat in
five locations on Kahoolawe, Lanai, and
Maui. The lands on which H.
assimulans occurs are under private and
State (DLNR and KIRC) ownership.
Specific Information on Hylaeus facilis
Taxonomy and Description
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 2003, 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 H. facilis.
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).
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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).
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The native host plants of adult
Hylaeus facilis are unknown, but it is
likely this species visits several plants
other Hylaeus species are known to
frequent, including Acacia koa,
Metrosideros polymorpha, Styphelia
tameiameiae, Scaevola spp., and
Chamaesyce spp. (Daly and Magnacca
2003, p. 11). Hylaeus facilis has also
been observed visiting the nonnative
Tourneforia argentea for nectar and
pollen (Magnacca 2007a, p. 181).
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 H. facilis was
among the most common and
widespread Hylaeus species on Oahu
and all of Maui Nui (Lanai, Maui, and
Molokai) (Magnacca 2007a, p. 183). 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 2007a, p. 183). Although the
species was widely collected within a
diverse range of habitats historically, 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).
Hylaeus facilis has almost entirely
disappeared from most of its historical
range (Daly and Magnacca 2003, p. 7;
Magnacca 2007a, p. 183). Between 1998
and 2006, 39 sites on Lanai, Maui,
Molokai, and Oahu were surveyed,
including 13 historical sites. Hylaeus
facilis was absent from each of the 13
historical localities (Magnacca 2007a, p.
183) and was also 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
2007a, p. 183). Likely extirpated from
Lanai, H. facilis is currently only known
from two locations, one each on the
islands of Molokai and Oahu (Daly and
Magnacca 2003, pp. 81–82; Magnacca
2005c, p. 2). In addition, in 1990, a
single individual was collected on Maui
in a residential area near Makawao at
1,500 ft (457 m) in elevation. However,
this site is an urbanized area devoid of
native plants, and it is likely this
collection was a single vagrant
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individual and not from an established
population on Maui.
Lanai
Perkins (1899) described Hylaeus
facilis as ‘‘common’’ at two Lanai
locations. He noted H. facilis was
collected from the Koele Mountains at
2,000 ft (610 m) in elevation.
Researchers believe the collection
locality was northwest of Puu Alii
where the ridges are at an elevation of
approximately 2,000 ft (600 m). The Puu
Alii summit itself is 2,800 ft (850 m) in
elevation, and less likely to be the site
of Perkins’ collection (Magnacca in litt.
2011, p. 36). Today this area contains
mixed native and nonnative vegetation.
Researchers collected three other
species of Hylaeus in the same general
area, along the Munro Trail and
Kaiholena ridge in 1999 and 2001 (Daly
and Magnacca 2003, pp. 217–229).
Perkins’ second collection site was in
montane habitat at 3,000 ft (900 m) in
elevation at Haalelepaakai in the
‘‘summit mountains on Lanai’’ (Daly
and Magnacca 2003, p. 83). Researchers
surveyed this area in 1999 and 2001,
and were unable to find H. facilis,
although they collected four other
Hylaeus species (Daly and Magnacca
2003, pp. 217–229). Hylaeus facilis is
likely extirpated from Lanai because it
has not been relocated in over 100 years,
and its potentially suitable habitat has
been extensively surveyed (Magnacca
2007a, pp. 177, 183).
Maui
Perkins collected Hylaeus facilis from
three different sites on Maui, including
coastal habitat at the Wailuku sand hills
(Waiehu dunes), montane mesic forest
habitat on Haleakala, and lowland wet
habitat in Iao Valley. Although other
species of Hylaeus were collected from
the Waiehu dunes in 1999 and 2001, H.
facilis, as well as several other species
collected by Perkins in the late 19th
century, were absent (Daly and
Magnacca 2003, pp. 217–229).
Perkins (1899) collected Hylaeus
facilis in montane mesic forest habitat
on Haleakala at an elevation of 5,000 ft
(1,524 m) on Haleakala, in the Olinda
area where he is known to have camped
while surveying for and collecting
insects (Evenhuis 2009, pp. 199–200).
These native forests were once abundant
in this area up to 6,000 ft (1,818 m) in
elevation across the west slope of
Haleakala, but have now been
completely converted by agriculture and
other land uses (Juvik and Juvik 1998,
pp. 123–124). Hylaeus facilis and other
species with similar habitat
requirements (e.g., H. difficilis, H.
volcanicus) are absent from the native,
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wetter forest across the eastern slope of
Haleakala (Daly and Magnacca 2003, pp.
219–221, 228–229).
Perkins also collected Hylaeus facilis
in lowland wet habitat at an elevation
of 2,000 ft (610 m) in Iao Valley in the
west Maui Mountains (H. V. Daly,
unpublished data). The terrain in Iao
Valley is especially rugged and wet, and
Perkins relied on assistants to collect
specimens from this area (Liebherr and
Polhemus 1997, p. 351). Even today the
vegetation in this area is predominantly
native (Liebherr and Polhemus 1997, p.
351).
Since the late 1960s, there have been
only two collections of Hylaeus facilis
on Maui, but neither is from a distinct
population that can be relocated. One
collection was made in 1967 (Daly and
Magnacca 2003, p. 221; Magnacca
2005c, p. 2), but the location is
unknown (Xerces Society 2009c, p. 7).
In 1990, a single individual was
collected at Kokomo at an elevation of
1,500 ft (457 m) near Makawao, in a
residential area devoid of native plants
(Daly and Magnacca 2003, p. 221). This
individual may have been a straggler
blown in from a different site altogether
(Magnacca 2005c, p. 2). Researchers
question whether any viable H. facilis
populations still remain on Maui
(Magnacca 2007a, pp. 183–184).
Molokai
Perkins collected Hylaeus facilis in
three locations within montane mesic
forest habitat in the east Molokai
Mountains (Daly and Magnacca 2003, p.
83). These locations were probably
between Makakupaia and the rim of
Pelekunu Valley, where Perkins did
most of his collecting (Liebherr and
Polhemus 1997, p. 347). Makakupaia is
located within TNC’s Kamakou
Preserve. Researchers have surveyed
extensively in similar, high-elevation
habitat near Perkins’ collecting area,
including Kamakou Road (3,200 ft (975
m)), Puu Kolekole (3,400 ft (1,040 m)),
and Kawela Gulch (3,600 ft (2,000 m)),
and found other Hylaeus species, but
were unable to locate H. facilis (Daly
and Magnacca 2003, pp. 217–229).
In 2005, researchers collected Hylaeus
facilis in coastal habitat at Kuololimu
Point, within Kalaupapa National
Historical Park (KNHP) on the southeast
coast of the Kalaupapa peninsula
(Magnacca 2007b, pp. 44–45). This area,
located on the east side of the
peninsula, is largely rocky and devoid
of vegetation, but contains scattered
patches of native coastal vegetation
similar to habitat at Ka Lae on the island
of Hawaii (Magnacca 2007a, p. 181). The
park is cooperatively managed by the
NPS, and the State of Hawaii’s DHHL,
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DLNR, DOH, and DOT (NPS 2006
(https://www.nps.gov/kala/index.htm)).
Oahu
Perkins collected Hylaeus facilis from
six sites on Oahu (Daly and Magnacca
2003, p. 83). One site described by
Perkins was coastal habitat in Honolulu.
Although the exact location is
unknown, Honolulu coastal habitat has
been completely developed for urban
land use and there is no potentially
suitable coastal habitat remaining in
Honolulu for Hylaeus species. Perkins
also described collecting Hylaeus
species from mountains in Honolulu,
and although the exact locations are
unknown, these sites are presumed to be
near known sites where he collected,
including Waiolani Ridge, Lanihuli
Ridge, Nuuanu Valley, and Konahuanui
(Liebherr and Polhemus 1997, p. 348).
While these mountain areas are largely
undeveloped, many are dominated by
nonnative vegetation. Researchers have
surveyed potentially suitable native
habitat near Perkins’ collection sites and
found other species of Hylaeus, but not
H. facilis (Daly and Magnacca 2003, pp.
217–229). Descriptions of the five
remaining suitable habitats follow:
(A) Makaha Valley: Perkins (1899)
collected H. facilis at an elevation of
3,000 ft (900 m) in the upper part of
Makaha Valley, on Oahu’s northwest
side. There have been no surveys for
Hylaeus in this area since Perkins’
collections, but researchers have
observed this area now lacks suitable
Hylaeus habitat due to development,
urbanization, and conversion of native
habitat to nonnative vegetation
(Magnacca, pers. comm. 2008c). Some of
the upper reaches of Makaha Valley
contain patches of native vegetation, but
much of the native vegetation has been
destroyed by brush fires (Liebherr and
Polhemus 1997, p. 347).
(B) Mount Kaala: Perkins (1899)
collected Hylaeus facilis at 2,000 ft (610
m) in elevation on Mt. Kaala, possibly
within what is now Mt. Kaala NAR.
This area is a mix of dry and mesic
forest communities (DLNR 1990, p. 3),
and is generally characterized as
predominantly native vegetation
(Liebherr and Polhemus 1997, p. 348).
This area has not been extensively
resurveyed for Hylaeus spp. because
much of it is either inaccessible (due to
either private or U.S. Army ownership),
or too rugged in general, requiring a
long and steep approach along the
Dupont Trail on the north slope of Mt.
Kaala.
(C) Waianae Mountains: Perkins
(1899) collected Hylaeus facilis in the
Waianae Mountains, ‘‘upland from
Waianae’’, likely in dry lowland forest,
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although the exact location is unknown.
In 2008, researchers surveyed
potentially suitable habitat in the
Waianae-Kaala Forest Reserve (FR), but
did not find H. facilis (Magnacca, pers.
comm. July 2008c).
(D) Tantalus: Perkins collected
Hylaeus facilis in lowland mesic habitat
on ‘‘Tantalus’’ (Liebherr and Polhemus
1997, p. 348), which today is in close
proximity to the urban core of
Honolulu. This area is a mix of
residential development and
undeveloped sites dominated by
nonnative plants, including various
species of Phyllostachys spp. (bamboo),
Acacia confusa (Formosa koa),
Eucalyptus robusta (swamp mahogany),
and Aleurites moluccana (kukui) (USDA
2001 https://soilseries.sc.egov.usda.gov/
OSD_Docs/T/TANTALUS.html). Habitat
dominated by nonnative plants does not
support viable populations of Hylaeus,
and no species have been reported from
this area since Perkins’ collections
despite more recent surveys in the few
small, widely separated areas containing
native plant habitat (Magnacca in litt.
2011, p. 41).
(E) Poamoho Trail: In 1975, Hylaeus
facilis was collected in lowland wet
forest at an unknown elevation along
the Poamoho Trail in Oahu’s Koolau
Mountains. Located in central Oahu, the
Poamoho Trail is part of the Na Ala Hele
trail and access system, and is within
the Ewa FR (DLNR 2008, p. 15). The
land adjacent to the trail, including the
access road to the forest reserve, is State
(DOFAW) and privately owned. The
Poamoho Trail traverses a public
hunting area, and some of the land
surrounding the access road is leased to
the Army for training purposes (DLNR
2011—https://hawaiitrails.ehawaii.gov/
trail.php?TrailID=OA+08+007). Access
is only allowed on weekends and
holidays, and by permit only. Dominant
vegetation in the summit area includes
the indigenous fern, Dicranopteris
linearis (uluhe), Acacia koa, and
Metrosideros polymorpha (DLNR
2011—https://hawaiitrails.ehawaii.gov/
trail.php?TrailID=OA+08+007).
Summary of Hylaeus facilis Range and
Distribution
At the end of the 19th century,
Hylaeus facilis was known from
numerous locations in coastal and
lowland habitats, including lowland
dry, mesic, and wet forest habitat on the
islands of Lanai, Maui, Molokai, and
Oahu. Currently, this species is only
known from two locations, one each on
the islands of Molokai and Oahu
(Magnacca 2007a, p. 177), under State
(DHHL, DLNR, DOFAW, DOH, DOT)
and private (TNC and others)
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ownership. Researchers question
whether viable populations of this
species remain on Maui because only
two single individuals have been
collected in the past 100 years.
Specific Information on Hylaeus hilaris
Taxonomy and Description
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). Hylaeus hilaris is
distinguished by its large size (male
wing length is 0.185 in (4.7 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 species (see Life
History below), 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 Hylaeus species to possess
apical (at the end or tip of a structure)
bands of fine white hairs on the
segments of the metasoma.
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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 hatching, the
cleptoparasitic larva kills the host egg,
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 other species possess on the front
legs. Cleptoparasitism is actually quite
common among bees, with
approximately 25 percent of known bee
species having evolved to become
cleptoparasites. Among the world’s
bees, other than the Hawaiian Hylaeus
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group, no cleptoparasites are known
from the family Colletidae (Daly and
Magnacca 2003, p. 9).
The larvae of Hylaeus 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 (Michener 2000, pp. 26–37, 126).
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 (Magnacca 2007a, p. 181).
Range and Distribution
Hylaeus hilaris was historically
known from coastal habitat on the
islands of Lanai, Maui, and Molokai. It
is believed to have occurred along much
of the coast of these islands’ as its
primary hosts, H. anthracinus, H.
assimulans, and H. longiceps, likely
extended throughout this habitat. The
majority of coastal habitat on these
islands has either been 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).
First collected on Maui in 1879,
Hylaeus hilaris has been collected only
twice in the last 100 years, but as noted
above, there is a gap of about 50 to 100
years between major collecting efforts.
Hylaeus hilaris has recently been
collected on two occasions: once in
1989 and again in 1999. On the islands
of Lanai and Maui, the species was
absent from each of its historical
Perkins-era localities revisited between
1998 and 2006 (Magnacca 2007a, pp.
177, 181–82). Currently, the only known
population of H. hilaris is located on
TNC’s Moomomi Preserve on Molokai
(Daly and Magnacca 2003, pp. 103, 106;
Magnacca 2005d, p. 2).
Lanai
Perkins (1899) collected Hylaeus
hilaris in coastal habitat at Manele, on
the southern coast of Lanai. This area is
now both the site of the ferry landing
from Lahaina, Maui, and a small boat
harbor, and is in close proximity to a
major resort. The area was surveyed in
1999, but researchers noted little native
vegetation aside from Scaevola sericea
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and an absence of Hylaeus species.
Additionally, the nonnative bee,
Lasioglossum impavidum (no common
name (NCN)), was found at the site.
Three other potentially suitable
locations were surveyed between 1999
and 2007 for Hylaeus species, but H.
hilaris was not observed at these sites,
despite the presence of H. assimulans
and H. longiceps, a recorded host
species (Daly and Magnacca 2003, p.
106; Magnacca 2007a, pp. 177, 181).
Most native coastal habitats are now
severely degraded across the entire
island, and it is believed Hylaeus hilaris
has likely been extirpated (Magnacca
2005d, p. 2; Magnacca 2007a, p. 181).
Although large areas of remote sandy
beach on the north and east coasts
remain to be thoroughly surveyed for
Hylaeus species, those that have been
inspected contain few native plants.
Two of the three known host species of
H. hilaris occur on Lanai, but all recent
(i.e., since 1999) collections have
primarily been made in lowland dry
forest habitat where H. hilaris has never
been collected.
Maui
Perkins collected Hylaeus hilaris from
three sites, including one now unknown
site possibly south of Wailuku and
simply labeled ‘‘Maui,’’ and two sites in
coastal habitat at the Wailuku sand hills
(an area noted as ‘‘the sandy isthmus’’)
(Daly and Magnacca 2003, p. 106). In
addition, in 1880, Reverend Thomas
Blackburn collected H. hilaris from an
unspecified location on the island (Daly
and Magnacca 2003, p. 106). Although
other rare Hylaeus species were
collected from the Waiehu dunes in
1999 and 2001 (See H. anthracinus
Range and Distribution), H. hilaris, as
well as several other species once
collected there by Perkins, was absent
(Daly and Magnacca 2003, pp. 217–229).
All three known host species of
Hylaeus hilaris occur on Maui.
However, H. anthracinus and H.
assimulans are currently known only
from dry forest or shrubland, which are
likely unsuitable habitat for H. hilaris.
The third known host species, H.
longiceps, occurs in the Wailuku sand
hills (Magnacca 2007a, p. 182). In
addition to its known historical sites,
several other potentially suitable sites
were surveyed between 1998 and 2006,
but H. hilaris was not found at any of
these sites, despite the presence of two
of its known host species (Daly and
Magnacca 2003, pp. 217–229; Magnacca
2007a, p. 177). Therefore, researchers
believe it is likely H. hilaris has been
extirpated from the island (Magnacca
2005d, p. 2).
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Molokai
Although Hylaeus hilaris was never
collected on Molokai by Perkins, in
1918, Fullaway (1918, p. 396) collected
the species at an unspecified site. As on
all of the Hawaiian Islands, most of the
coastal habitat on Molokai is now
dominated by nonnative vegetation.
Currently, the only known population of
H. hilaris occurs on the northwest coast
within TNC’s Moomomi Preserve. This
site is part of a large area of windswept
calcified dunes, some of which are
dominated by native plants while other
portions of the dunes are dominated by
nonnative plant species. Hylaeus
anthracinus and H. longiceps, both host
species of H. hilaris, are presently
known to occur in Moomomi Preserve
(Magnacca 2007a, p. 181). Only two
collections of H. hilaris have been made
at Moomomi since it was discovered at
this site in 1930. Both collections, 1989
and 1999, were of a single male. Dunes
to the west of Moomomi Preserve are
dominated by nonnative vegetation, and
no species of Hylaeus have been
collected from those areas. While H.
anthracinus, one of the host species of
H. hilaris, is currently known from the
Kalapapa peninsula, H. hilaris has never
been collected there.
Summary of Hylaeus hilaris Range and
Distribution
Hylaeus hilaris was historically
known from coastal habitat on the
islands of Lanai, Maui, and Molokai. It
is believed to have occurred along much
of the coast of these islands’ as its
known hosts, H. anthracinus, H.
assimulans, and H. longiceps, likely
also occurred throughout coastal habitat
on these three islands. Currently, H.
hilaris is only known from one site on
Molokai.
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Specific Information on Hylaeus kuakea
Taxonomy and Description
Hylaeus kuakea was first described by
Daly and Magnacca (2003, pp. 1, 125–
1,127) 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;
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Magnacca 2005e, p. 2). Only four adult
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 (Daly and
Magnacca 2003, p. 9). 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).
The native host plants of the adult
Hylaeus kuakea are unknown, but it is
likely this species visits several plants
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
In 1997, researchers collected 2 male
individuals of Hylaeus kuakea in
lowland mesic forest at an elevation of
about 1,900 ft (579 m) on Moho Gulch
Ridge at the northern end of the State’s
recently acquired Honouliuli Preserve
in the Waianae Mountains on Oahu.
Researchers surveyed the middle and
southern portions of the Preserve, but
they did not find H. kuakea, although
other species of Hylaeus are known
from these areas. In 2010, researchers
collected this species (two males), on
the endangered plant Chamaesyce
herbstii (akoko) in a remnant patch of
diverse lowland mesic forest in Makaha
Valley on Oahu’s west side (Magnacca,
in litt., 2010, p. 1). Phylogenetically, H.
kuakea belongs in a species-group
primarily including mesic forestinhabiting species (Magnacca &
Danforth 2006, p. 405).
Summary of Hylaeus kuakea Range and
Distribution
Because the first collection of Hylaeus
kuakea was not made until 1997, its
historical range is unknown (Magnacca
2005e, p. 2; Magnacca 2007a, p. 184).
Only four individuals (all males) of H.
kuakea have been collected at two
different sites in lowland mesic forest
habitat in the Waianae Mountains on
Oahu (Magnacca 2007a, p. 184;
Magnacca, in litt., 2010, p. 1), and the
species has never been collected in any
other habitat type or area, including
some that have been more thoroughly
surveyed (Magnacca in litt., 2011, p. 49).
Researchers have not exhaustively
surveyed all potentially suitable
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lowland mesic forest areas due their
remote and rugged locations, small size,
and distant spacing among large areas of
nonnative forest. Lowland mesic forest
habitat is becoming 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).
Specific Information on Hylaeus
longiceps
Taxonomy and Description
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 H. longiceps.
Hylaeus longiceps is a small to mediumsized, 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 (Daly and Magnacca 2003, p. 9).
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 native plants, including
Scaevola coriacea (dwarf naupaka), Sida
fallax, Scaevola spp., Sesbania
tomentosa, Myoporum sandwicense,
Santalum ellipticum, Chamaesyce
degeneri, and Vitex rotundifolia
(pohinahina) (Daly and Magnacca 2003,
p. 135). It is also likely H. longiceps
visits several plant species other
Hylaeus species are known to frequently
visit, including Scaevola spp.,
Chamaesyce spp., Tournefortia
argentea, Jacquemontia ovalifolia, and
Sida fallax (Magnacca 2005f, p. 2).
Range and Distribution
Hylaeus longiceps is historically
known from coastal and lowland dry
shrubland habitat up to 2,000 ft (610 m)
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in elevation in numerous locations on
the islands of Lanai, Maui, Molokai, and
Oahu. Perkins (1899, p. 98) noted H.
longiceps was locally abundant, and
probably occurred historically
throughout much of the leeward and
lowland areas on Lanai, Maui, Molokai,
and Oahu, as its host plants, Sida fallax,
Chamaesyce spp., Scaevola spp., and
Jaquemontia ovalifolia, occurred
throughout these areas (Magnacca 2005f,
p. 2). 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 2007a, pp. 186–
188).
Hylaeus longiceps is now restricted to
small populations in small patches of
coastal and lowland dry habitat on
Lanai, Maui, Molokai, and Oahu
(Magnacca 2005f, p. 2). Twenty-five
sites that were either historical
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 and one site
each on the islands of Maui, Molokai,
and Oahu. Only one historical location,
Waieu dunes on Maui, still supports a
population of H. longiceps (Daly and
Magnacca 2003, p. 135).
Lanai
Perkins (1899) collected Hylaeus
longiceps at Manele, and other
unspecified localities (labeled ‘‘Lanai’’).
Between 1999 and 2001, researchers
surveyed seven sites for Hylaeus
species, and were unable to find H.
longiceps at Manele Bay, although other
rare Hylaeus species were observed
there (Daly and Magnacca 2003, pp.
217–229). In addition, researchers did
not find H. longiceps at three other sites
within potentially suitable lowland dry
habitat, including the Kahue unit of the
privately owned Kanepuu Preserve,
Garden of the Gods, and the Munro
Trail/Kaiholena area of the Koele
mountains (Daly and Magnacca 2003,
pp. 217–229). Hylaeus longiceps is now
known only from very small pockets of
native vegetation in three locations on
private land, including lowland dry
forest habitat at Kahue and Polihua
Road, and coastal habitat at Shipwreck
Beach. Descriptions of these three
locations follow:
(A) Kahue and Polihua Road: In 1999,
Magnacca collected Hylaeus longiceps
in lowland dry forest at Kahue (south of
Kanepuu Preserve) at an elevation of
1,400 ft (427 m) (Daly and Magnacca
2003, p. 135). Researchers also surveyed
the Kanepuu Preserve for H. longiceps,
but were unable to find this species. In
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1999, researchers collected H. longiceps
in lowland dry forest at 1,000 ft (300 m)
in elevation, along Polihua Road in
central Lanai (Daly and Magnacca 2003,
p. 135).
(B) Shipwreck Beach: Although he
did not collect Hylaeus longiceps at
Shipwreck Beach, Perkins collected
other species of Hylaeus at Awalua,
about 4 miles to the west (Daly and
Magnacca 2003, p. 58). In 2001,
researchers collected H. longiceps in
native, coastal habitat at Shipwreck
Beach (Daly and Magnacca 2003, p.
135). Shipwreck Beach is a popular
tourist site on Lanai and accessible by
four-wheel drive vehicles.
Maui
Perkins (1899) collected Hylaeus
longiceps at the Wailuku sand hills
(Waiehu Dunes) and on Haleakala. In
addition, some of his specimens were
collected from unknown localities
labeled ‘‘Maui.’’ Perkins collected H.
longiceps in dry forest habitat at an
elevation of 2,000 ft (610 m) on
Haleakala, probably near the towns of
Pukalani or Makawao, where he stopped
on his way to Wailuku (Daly and
Magnacca 2003, p. 135). Native dry
forests that supported populations of
Hylaeus were common in lowland areas
when Perkins collected, but this habitat
has been greatly reduced and
fragmented.
Hylaeus longiceps is now known from
only one Maui location, at the Wailuku
sand hills (Waiehu dunes). Between
1999 and 2001, a total of seven
specimens were collected in native
habitat in the northern portion of the
dunes (Daly and Magnacca 2003, p.
224). Researchers surveyed for, but did
not find, H. longiceps in the southern
(Kahului) portion of the dunes (Daly
and Magnacca 2003, p. 224).
Hylaeus longiceps was not found in
five other sites on Maui surveyed
between 1999 and 2001 (Daly and
Magnacca, pp. 217–229). One historical
site, in dry forest habitat on the slopes
of Haleakala, has been developed and is
overgrown with nonnative, invasive
plants (Magnacca, pers. comm., 2008f).
Hylaeus longiceps was absent from four
sites (Kanaio NAR, Lahainaluna,
Manawainui Gulch, and Waikapu near
Kaohonua) with potentially suitable
habitat where other Hylaeus species
with similar habitat requirements were
recently collected (Daly and Magnacca
2003, pp. 217–229).
Molokai
Perkins (1899) collected Hylaeus
longiceps at Kaunakakai, and at
unknown locations labeled ‘‘Molokai
coast and plains,’’ the ‘‘west end’’ [of
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55181
the island], and the ‘‘Molokai
Mountains.’’ Although Kaunakakai is
the primary urban area on Molokai,
researchers surveyed this area, noting
any former Hylaeus habitat has been lost
to urban development and nonnative,
invasive plants (Magnacca, pers. comm.,
2008f). Most coastal habitat on the west
end of Molokai, with the exception of
TNC’s Moomomi Preserve, has been
degraded and converted to nonnative,
invasive plants (Magnacca, pers. comm.,
2008f).
Researchers surveyed a total of six
sites on Molokai over the last several
years for Hylaeus longiceps, and
observed 8 individuals at Moomomi
Preserve (in 1999 and in 2001) (Daly
and Magnacca 2003, p. 135). Hylaeus
longiceps was notably absent from three
sites on the Kalaupapa peninsula
(Kuololimu Point, Hoolehua Beach, and
Kaupikiawa), where other Hylaeus
species have been recently collected
(Daly and Magnacca 2003, pp. 217–229).
Researchers were unable to find H.
longiceps in sand dune habitat near the
Kaluakoi Resort on Molokai’s northwest
coastline (Magnacca, pers. comm.,
2008f).
Oahu
Perkins (1899) collected Hylaeus
longiceps from only one site, in a coastal
area of southwest Waianae. In 1999,
2000, and 2002, researchers found H.
longiceps in coastal habitat at the State’s
Kaena Point NAR (Daly and Magnacca
2003, p. 224). Researchers did not find
H. longiceps during surveys conducted
at other coastal sites with potentially
suitable habitat, including Makapuu in
1999, and Kalaeloa in 2002. Although
both areas contain vegetation similar to
the vegetation in the Kaena Point NAR,
albeit more degraded, no species of
Hylaeus were observed in these areas
(Daly and Magnacca 2003, pp. 217–229;
Magnacca, pers. comm., 2008f).
Summary of Hylaeus longiceps Range
and Distribution
Hylaeus longiceps was historically
known from numerous coastal and
lowland dry forest locations up to 2,000
ft (600 m) in elevation on the islands of
Lanai, Maui, Molokai, and Oahu.
Currently, H. longiceps is restricted to a
total of six populations in small patches
of coastal and lowland dry forest
habitat: three sites on Lanai and one site
each on the islands of Maui, Molokai,
and Oahu (Magnacca 2005f, p. 2). The
lands on which H. longiceps occurs are
under a variety of jurisdictions
including private (e.g., TNC) and State
(NARS).
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Specific Information on 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, species with overlapping
ranges, 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 (Daly and
Magnacca 2003, p. 9). 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 2005g, p. 2;
Magnacca and Danforth 2006, p. 403).
Adult specimens of Hylaeus mana
were collected while they visited
flowers of Santalum freycinetianum var.
freycinetianum (iliahi, sandalwood), a
native Hawaiian plant found only on
Oahu and Molokai (Wagner et al. 1999,
p. 1,221). It is likely 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 (430 m). Few Hylaeus bees have been
found in this type of Acacia koadominated, 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
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Juvik 1998, p. 124; Wagner et al. 1999,
pp. 66–67, 75).
The Manana Trail is part of the Na
Ala Hele Hawaii Statewide Trail and
Access System (DLNR 2007), and is
located within the State’s Ewa FR. Six
miles in length, the beginning of the
Manana Trail is dominated by
nonnative plant species, but leads into
an area of native forest where Acacia
koa, Metrosideros polymorpha, and
Scaevola spp. are common (DLNR
2011—https://hawaiitrails.ehawaii.gov/
trail.php?TrailID=OA+09+008).
Summary of Hylaeus mana Range and
Distribution
Because the first collection of Hylaeus
mana was made in 2002, its historical
range and current distribution, other
than the collection on Manana Trail, are
unknown at this time (Magnacca 2005g,
p. 2). Additional surveys in potentially
suitable habitat may reveal additional
populations elsewhere on Oahu
(Magnacca 2007a, p. 184). However, the
extreme rarity of this species, its
absence from nearby sites, and the fact
it was not discovered until very
recently, suggests very few populations
remain (Magnacca 2005g, p. 2).
Summary of Information Pertaining to
the Five Factors
Section 4 of the Act (16 U.S.C. 1533)
and its implementing regulations (50
CFR 424) set forth procedures for adding
species to the Federal Lists of
Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, a species may be determined to be
endangered or threatened based on any
of the following five factors:
(A) The present or threatened
destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial,
recreational, scientific, or educational
purposes;
(C) Disease or predation;
(D) The inadequacy of existing
regulatory mechanisms; or
(E) Other natural or manmade factors
affecting its continued existence.
In making this finding, information
pertaining to the seven species of
Hawaiian yellow-faced bees in relation
to the five factors provided in section
4(a)(1) of the Act is discussed below.
In considering what factors might
constitute threats, we must look beyond
the exposure of the species to the factor
to determine whether the species
responds to the factor in a way that
causes actual impacts to the species. If
there is exposure and the species
responds negatively, the factor may be
a threat and we then attempt to
determine how significant a threat it is.
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If the threat is significant, it may drive
or contribute to the risk of extinction of
the species such that the species
warrants listing as endangered or
threatened as those terms are defined by
the Act.
Factor A. Present or Threatened
Destruction, Modification, or
Curtailment of the Habitat or Range
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;
Magnacca, pers. comm. 2010). 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, 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
Destruction and modification of
Hylaeus bee habitat by urbanization and
land use conversion leads to the direct
fragmentation of foraging and nesting
habitat of these species. In particular,
because native host plant species are
known to be essential to the yellowfaced bees for foraging of nectar and
pollen, any further loss of this habitat
may endanger their long-term chances
for conservation and recovery.
Additionally, conversion and
modification of the seven yellow-faced
bees’ habitat is also likely to further
exacerbate the introduction and spread
of nonnative plants into and within
these areas (see Habitat Destruction and
Modification by Nonnative Plants
section below).
Coastal Habitat
Native coastal habitat is one of the
rarest habitats on the main Hawaiian
Islands (Hawaii, Kahoolawe, Kauai,
Lanai, Maui, Molokai, and Oahu)
(Wagner et al. 1999, pp. 45, 54; Cuddihy
and Stone 1990, pp. 94–95; Magnacca
2007, p. 180). Coastal habitat is highly
valued for development, popular for
recreation, typically dry on both the
windward and leeward sides of the
islands, vulnerable to fire, and
especially susceptible to invasion by
nonnative plants. Increased access to
coastal areas, and resulting habitat
disturbance, has been facilitated by
development, road-building, and past
agricultural activities (Cuddihy and
Stone 1990, pp. 94–95). The native
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coastal habitat that remains is in small
remnant patches, and most of these
remnants have been overtaken by
invasive plant species and have
relatively low diversity (Cuddihy and
Stone 1990, pp. 94–95) (see Habitat
Destruction and Modification by
Nonnative Plants section below). Most
of the coastal areas of the main
Hawaiian Islands now lack significant
amounts of native plants suitable for
foraging by Hylaeus, other than
Scaevola sericea, which alone cannot
support Hylaeus populations (Magnacca
2007a, p. 187). The restricted and
isolated nature of coastal habitat places
species that depend on these areas even
more at risk for a variety of reasons,
including but not limited to their
increased susceptibility to random
events (e.g., hurricanes and wildfire),
the reduced range of native plants
including host plants, and the reduced
number of suitable sites for species to
expand their range (Sakai et al. 2002, p.
291).
Five species of Hawaiian yellow-faced
bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, and H.
longiceps) were once widespread and
common in coastal habitat (Perkins
1912, p. 688) throughout the main
Hawaiian Islands (see Table 1 above),
with the exception of Kauai. These five
species are now absent from all of
Perkins’ coastal collection localities
(Kealakekua Bay and Keei and the urban
area near Kona on the island of Hawaii;
the Awalua area on Lanai; the Wailuku
sand hills area on Maui; the northwest
dunes and Kaunakakai areas on
Molokai; and Waikiki, the Waianae area,
and the Honolulu mountains on Oahu)
(Daly and Magnacca 2003, pp. 217–229),
although they have recently been
collected in disparate coastal habitat on
one or more of the islands of Hawaii,
Kahoolawe, Lanai, Maui, Molokai, and
Oahu (Daly and Magnacca 2003, pp.
217–229).
Lowland Dry Habitat
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). Less than 1 percent of lowland dry
forest and shrubland remains on Oahu,
Molokai, and Lanai; less than 2 percent
remains on Maui; and less than 17
percent remains on Hawaii Island (Sakai
et al. 2002, p. 296). Without greater
conservation and restoration efforts, we
believe the remaining lowland dry forest
and shrublands, which were once
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abundant and perhaps the most diverse
of all Hawaiian habitat types (Medeiros
2006, p. 1), could completely disappear
due to continued development and
other land use conversion, compounded
by the effects of nonnative species, wild
fire, and stochastic events (see following
sections on Habitat Destruction and
Modification by Nonnative Plants; by
Nonnative Ungulates; by Fire; by
Recreational Activities; by Hurricanes
and Drought; and by Climate Change)
(Cabin et al. 2000, p. 449).
Four species (Hylaeus anthracinus, H.
assimulans, H. facilis, and H. longiceps)
were once widespread (i.e., there were
several populations across two or more
islands) and found within lowland dry
habitat on several islands, including
Hawaii, Lanai, Maui, Molokai, and
Oahu. However, these species have not
been observed during recent surveys
from their historical population sites on
these islands (Magnacca 2005a, b, c, f,
pp. 1–2). Five of the seven Hylaeus bee
species (Hylaeus assimulans, H. facilis,
H. kuakea, H. longiceps, and H. mana)
are most often found in dry and mesic
forest (see discussion below) 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). According to Magnacca
(2007, pp. 186–187), lowland dry and
mesic forests now support less-diverse
Hylaeus communities because many
native plants used for foraging are
extirpated from these habitats.
Lowland Mesic Habitat
Hawaii’s lowland mesic forest habitat
was once abundant and considered the
most diverse (in terms of number of
species) of all Hawaiian forest types
(Rock 1913, p. 9). Lowland mesic forest
habitat is now very rare, and has been
converted to pasture, military use,
agricultural use, or lost to urbanization.
Development and land use conversion is
ongoing (Cuddihy and Stone 1990, p.
61; Magnacca 2007, p. 187; Wagner et
al. 1999, p. 75). Fire has also negatively
impacted this habitat type and remains
a significant threat (see Habitat
Destruction and Modification by Fire
section below).
Historically, Hylaeus facilis was
found in a wide variety of habitats
including lowland mesic forest on
Lanai, Maui, and Oahu and montane
mesic habitat on Molokai. However, this
species no longer occurs in these
habitats on any of these four islands.
Hylaeus kuakea and H. mana are known
from a total of three locations in
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lowland mesic forest habitat on the
island of Oahu. Because we lack
information on the historical range of H.
kuakea and H. mana (they were only
discovered relatively recently), we are
unable to determine the extent of habitat
loss these two species have experienced.
However, because the extent and the
quality of lowland mesic forest has been
reduced throughout the Hawaiian
Islands, it is reasonable to conclude H.
kuakea and H. mana now have less
habitat because of urbanization and land
use conversion.
Lowland Wet Habitat
Native lowland wet forests were once
one of the dominant ecosystem types in
lowland areas on the main Hawaiian
Islands (Wagner et al. 1999, p. 45). Most
of the original loss of this habitat type
was due to agricultural uses in the 18th
and 19th centuries, and many remaining
areas were overtaken by aggressive
nonnative plant species such as Psidium
cattleianum (strawberry guava),
nonnative grasses such as Brachiaria
mutica (California grass), and Rubus
spp. (e.g., prickly Florida blackberry,
thimbleberry). Remnants of native
lowland wet forest can be found in
rocky or steep terrain, such as on some
peaks and summit ridges on Oahu,
Molokai, and West Maui (Cuddihy and
Stone 1990, p. 105). Although these
remaining remote and remnant native
lowland areas are now less likely
threatened by land use conversion, they
remain very threatened by the impacts
of nonnative plants (see Habitat
Destruction and Modification by
Nonnative Plants section below).
Furthermore, the original loss of
lowland and montane wet forest habitat
on Oahu, Lanai, Maui, and Molokai was
likely a contributing factor to the
decline of H. facilis, a species now
known only from coastal habitat on
Molokai and wet forest habitat on
Oahu’s Poamoho Trail. Researchers
believe the site on Oahu likely once had
more open understory and the presence
of H. facilis in this wet forest habitat
represents an outlier or residual
population (Perkins 1899, p. 76;
Liebherr and Polhemus 1997, p. 347).
In summary, destruction and
modification by urbanization and land
use conversion of the coastal and
lowland habitat of the seven Hylaeus
bees is continuing, and is expected to
continue reducing and fragmenting the
remaining habitat available to the
yellow-faced bees in the future,
endangering the species’ long-term
chances for conservation and recovery.
Because of the decreased amount of
suitable native coastal and lowland
habitat remaining in the Hawaiian
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Islands and the continued conversion of
these native habitats by development,
road building, or agriculture, we
conclude the ongoing habitat loss and
land modification is a significant
ongoing threat to H. anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana.
Habitat Destruction and Modification by
Nonnative Plants
Native vegetation on all of the main
Hawaiian Islands has undergone
extreme alteration because of past and
present land management practices,
including ranching, agricultural
development, and the deliberate
introduction of nonnative plants and
animals (Cuddihy and Stone 1990, pp.
27, 58). The original native flora of
Hawaii (species that were present before
humans arrived) consisted of about
1,000 taxa, 89 percent of which were
endemic (species that occur only in the
Hawaiian Islands). Over 800 plant taxa
have been introduced from elsewhere,
and nearly 100 of these have become
pests (e.g., injurious plants) in Hawaii
(Smith 1985, p. 180; Cuddihy and Stone
1990, p. 73; Gagne and Cuddihy 1999,
p. 45). Some of these plants were
brought to Hawaii by various groups of
people, including the Polynesians, for
food or cultural reasons. Beginning in
the early 1900s, plantation owners (and
the territorial government of Hawaii),
alarmed at the reduction of water
resources for their crops caused by the
destruction of native forest cover by
grazing feral and domestic animals,
introduced nonnative trees for
reforestation and continued the practice
through the late 1930s (Nature
Conservancy of Hawaii 2003, p. 19).
Ranchers intentionally introduced
pasture grasses and other nonnative
plants for agriculture, and sometimes
inadvertently introduced weed seeds as
well. Other plants were brought to
Hawaii for their potential horticultural
value (Scott et al. 1986, pp. 361–363;
Cuddihy and Stone 1990, p. 73).
Nonnative plants adversely impact
native Hawaiian habitat, including that
of the seven yellow-faced bees
identified in this finding, by modifying
the availability of light, altering soilwater regimes, modifying nutrient
cycling, altering fire characteristics of
native plant communities (for example,
successive fires that burn farther and
farther into native habitat, destroy
native plants, and remove habitat for
native species by altering microclimatic
conditions to favor nonnative species),
and ultimately converting native
dominated plant communities to
nonnative plant communities (Smith
1985, pp. 180–181; Cuddihy and Stone
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1990, p. 74; D’Antonio and Vitousek
1992, p. 73; Vitousek et al. 1997, p. 6).
Nonnative plants directly and indirectly
affect the seven yellow-faced bees by
modifying or destroying their terrestrial
and riparian habitat and reducing food
sources.
The spread of nonnative plant species
is one of the primary causes of decline
of the seven Hylaeus bee species, and a
current threat to their existing
populations because these bees depend
closely on native vegetation for nectar
and pollen. 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). The native flora within most of
lowland habitat in the Hawaiian Islands
is being replaced by aggressive,
nonnative plant species (Cuddihy and
Stone 1990, pp. 73–74; Wagner et al.
1999, p. 52). Many native plant species
communities that have been replaced by
often monotypic communities of
nonnative plants were once foraging
resources for numerous species of
Hylaeus bees (Cox and Elmqvist 2000, p.
1238; Daly and Magnacca 2003, p. 11;
USFWS 1999, pp. 145, 163, 171, 180;
USFWS 2008b, pp. 7, 9).
Many of the native plants that
currently serve as foraging resources for
the adults of the seven Hylaeus bee
species are declining due to a lack of
pollinators and competition with
nonnative plants (Daly and Magnacca
2003, p. 11; USFWS 2008b, pp. 7, 9;
Smith 1985, pp. 180–181; Cuddihy and
Stone, 1990, p. 74; D’Antonio and
Vitousek 1992, p. 73; Vitousek et al.
1997, p. 6), and are found only in very
small populations (USFWS 1999, pp.
145, 163, 171, 180; Cox and Elmqvist
2000, p. 1,238). For example, H.
longiceps and H. anthracinus are known
to forage on the federally endangered
plant Sesbania tomentosa. Both H.
longiceps and H. anthracinus also visit
Chamaesyce celastroides var. kaenana,
a federally endangered plant endemic to
coastal dry shrubland on Oahu (Koutnik
1999, p. 606; Daly and Magnacca 2003,
pp. 55, 74). Hylaeus longiceps is also
known to forage on the endangered
Scaevola coriacea (USFWS 1999, p. 145;
Daly and Magnacca 2003, pp. 55, 135).
In addition, H. anthracinus has been
collected from inside the fruit capsule of
Hedyotis coriacea, a federally
endangered dry forest plant, known
from fewer than 200 individuals on the
island of Hawaii (Center for
Environmental Management of Military
Lands, 2010). Several other widespread
nonnative plant species threaten coastal
habitats of the five Hylaeus species
known from these areas. Understory and
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sub-canopy species include Asystasia
gangetica (Chinese violet), Atriplex
semibaccata (Australian saltbush),
Leucana leucocephala (koa haole),
Pluchea indica (Indian fleabane), P.
symphytifolia (sourbush), and Verbesina
encelioides (golden crown-beard)
(DOFAW 2007, pp. 20–22, 54–58;
HBMP 2008). Nonnative canopy species
include Prosopis pallida (kiawe)
(DOFAW 2007, pp. 20–22, 54–58;
HBMP 2008), an invasive, nonnative,
deciduous thorny tree (TNC 2009, p. 8).
For example, in Moomomi Preserve on
Molokai, which represents the only
known location for Hylaeus hilaris,
most of the sand dunes and areas
adjacent to the preserve are entirely
covered in Prosopis pallida. The narrow
coastal strip in the Preserve itself is the
only area that remains somewhat intact
with native plant species (TNC 2008, p.
8; Magnacca in litt. 2011, p. 65). In
addition, several nonnative grasses such
as Cenchrus ciliaris (buffelgrass),
Chloris barbata (swollen fingergrass),
Digitaria insularis (sourgrass), and
Panicum maximum (guinea grass)
threaten the coastal habitats in which
they are known to occur (DOFAW 2007,
pp. 20–22, 54–58; HBMP 2008).
As noted in the Life History section,
above, Hylaeus species almost
exclusively visit native plants to collect
nectar and pollen (Daly and Magnacca
2003, p. 11), pollinating those plants in
the process (Sakai et al. 1995, pp. 2,524–
2,528; Cox and Elmqvist 2000, p. 1,238;
Sahli et al. 2008, p. 1). Hylaeus bees are
very rarely found visiting nonnative
plants for nectar and pollen (Magnacca
2007a, pp. 186, 188). Unpublished data
on Hylaeus spp. pollen use (Magnacca
in litt. 2011, p. 65) suggest only
approximately 3 percent of pollen
collected by yellow-faced bees (although
not exclusively the seven Hylaeus
species addressed in this finding) is
from nonnative plant sources. These
data do not include observations
regarding yellow-faced bee use of
Tournefortia argentea, which is a
naturalized and relatively recent arrival
to the Hawaiian Islands, as a pollen
resource (Magnacca in litt. 2011, p. 65)
(see additional information on this
species below). Other than Scaevola
sericea, native vegetation is lacking
along most of the coastline of the main
Hawaiian Islands. As Hylaeus spp. have
not been observed at coastal sites where
Scaevola sericea represents the only
native plant species occurrence,
researchers believe the yellow-faced
bees are unable to survive on this
species alone (Magnacca 2007, p. 187;
Magnacca in litt. 2011, p. 65).
In summary, the spread of nonnative
plants throughout the coastal and
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lowland habitat of the seven Hylaeus
bees represents a serious and ongoing
threat to these species. Many of the
native plant species being replaced by
invasive, nonnative plants provide
foraging resources (e.g. pollen, nectar)
for Hylaeus bees, including these seven
species. The best available information
indicates these seven bee species do not
characteristically forage on nonnative
plants (Daly and Magnacca 2003, p. 13).
Only 14 of 820 recent (1998 to 2010)
Hylaeus spp. observations were on
flowers of nonnative plant species;
however, none of those observations
involved the seven Hylaeus species
addressed in this finding. We
acknowledge those observations do not
include records documenting Hylaeus
spp. using Tournefortia argentea
(another nonnative species). However,
there are only 13 observations of
Hylaeus spp. using this species,
including four records for H.
anthracinus and one record for H. facilis
(Magnacca in litt. 2011, p. 66).
Therefore, we conclude that the ongoing
spread of nonnative plants into the
habitats of the seven Hylaeus bees
remains a significant threat due to
manner in which nonnative plants alter
and fragment habitat, increase the
likelihood of fire, and attract nonnative
insect species. This threat further
endangers the species’ long-term
chances for conservation and recovery.
Habitat Destruction and Modification by
Nonnative Ungulates
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 and moving
across large areas) facilitates the spread
and establishment of competing,
invasive, nonnative plant species
(Cuddihy and Stone 1990, p. 65). Feral
pigs occur on all of the main Hawaiian
Islands except Kahoolawe and Lanai
(HEAR 1998; C. Kessler, USFWS, pers.
comm. 2011); goats are found on all of
the main Hawaiian Islands except Lanai
(HEAR 1998); feral cattle are found on
Hawaii and Maui (HEAR 1998);
Mouflon sheep and hybrids are found
on Hawaii and Lanai (Hawaii
Conservation Alliance (HCA) 2007); and
axis deer are found on Lanai, Maui,
Molokai, and Oahu (HCA 2007). At least
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one endangered coastal and lowland
plant species, Sesbania tomentosa,
threatened by the browsing, trampling,
and digging activities of nonnative
ungulates (e.g., axis deer, goats, and
cattle), is a foraging source for Hylaeus
anthracinus and H. longiceps (USFWS
1999, pp. 145, 163, 171, 180; Daly and
Magnacca 2003, pp. 11, 13).
The State of Hawaii provides game
mammal (e.g., feral pigs, goats, and
deer) hunting opportunities on Statedesignated public hunting areas on the
islands of Hawaii, Kauai, Lanai, Maui,
Molokai, and Oahu (Hawaii
Administrative Rules § 13–123–14–13–
123–20; DLNR 1999). The State’s
management objectives for game
animals ranges from maximizing public
hunting opportunities (e.g., ‘‘sustained
yield’’) in some areas to removal by
State staff, or their designees, in other
areas (Hawaii Administrative Rules
§ 13–123). Several of the seven Hylaeus
bees have populations in or adjacent to
areas where terrestrial habitat may be
manipulated for game enhancement and
where game populations are maintained
at certain levels for public hunting
(Hawaii Administrative Rules § 13–123).
Public hunting areas are predominantly
not fenced, and game mammals have
unrestricted access to most areas across
the landscape, regardless of underlying
land use designation. While fences are
sometimes built to provide protection
from game mammals to the natural
resources within the fenced area, the
current number and locations of fences
are not adequate to prevent habitat
destruction and degradation of the
terrestrial habitat of the seven species of
Hawaiian yellow-faced bees.
In summary, feral pigs, cattle, goats,
and axis deer continue to alter and
degrade native vegetation within
Hylaeus habitat in the Hawaiian Islands.
We believe these ungulates represent a
significant and ongoing threat to the
continued existence of the seven
Hylaeus bees, endangering the species’
long-term chances for conservation and
recovery. Ungulates directly trample
and consume native plants, including
plants used for foraging by H.
anthracinus and H. longiceps. The best
available information indicates that
other than the plant Tournefortia
argentea, none of the seven Hylaeus
bees use nonnative plants for foraging
(Daly and Magnacca 2003, p. 13). While
some specific areas throughout the
State, including some Hylaeus spp.
habitat sites, are managed to exclude the
presence of or control ungulates, we are
unaware of any plans to entirely
eradicate or eliminate ungulates from
the Hawaiian Islands. In addition,
public hunting areas maintain
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populations of nonnative ungulates and
often do not provide adequate fencing to
prevent nonnative ungulates from
negatively impacting the habitat of the
seven yellow-faced bees. Therefore, the
ongoing alteration and degradation of
many of the native coastal and lowland
habitat where these seven Hylaeus bees
occur by ungulates is expected to
further impact the bees’ foraging and
nesting habitat through the direct
consumption and trampling of native
plants, introduction and spread of
nonnative plants, and increased erosion.
Habitat Destruction and Modification by
Fire
Fire is a relatively new, humanexacerbated threat to native species and
natural vegetation in Hawaii. The
historical fire regime in Hawaii was
characterized by infrequent, low
severity fires, as few natural ignition
sources existed (Cuddihy and Stone
1990, p. 91; Smith and Tunison 1992,
pp. 395–397). Natural fuel beds were
often discontinuous, with moderate to
high rainfall in many areas on most
islands. Fires inadvertently or
intentionally ignited by the original
Polynesians in Hawaii probably
contributed to the initial decline of
native vegetation in the drier plains and
foothills. These early settlers practiced
slash-and-burn agriculture that created
open lowland areas suitable for the later
colonization of nonnative, fire-adapted
grasses (Kirch 1982, pp. 5–6, 8; Cuddihy
and Stone 1990, pp. 30–31). Beginning
in the late 18th century, Europeans and
Americans introduced plants and
animals that further degraded native
Hawaiian ecosystems. Pasture areas and
ranching, in particular, created highly
fire-prone areas of nonnative grasses
and shrubs (D’Antonio and Vitousek
1992, p. 67). Fires of all intensities,
seasons, and sources are destructive to
native Hawaiian ecosystems (Brown and
Smith 2000, p. 172), and a single grassfueled fire can kill most native trees and
shrubs in the burned area (D’Antonio
and Vitousek 1992, p. 74). Although
Vogl (1969) (in Cuddihy and Stone
1990, p. 91) suggests naturally occurring
fires, primarily from lightning strikes,
have been important in the development
of the original Hawaiian flora, and many
Hawaiian plants might be fire-adapted,
Mueller-Dombois (1981) (in Cuddihy
and Stone 1990, p. 91) points out most
natural vegetation types of Hawaii
would not carry fire before the
introduction of nonnative grasses. Smith
and Tunison (in Cuddihy and Stone
1990, p. 91) state native plant fuels
typically have low flammability.
Fire represents a threat to the seven
Hylaeus species in coastal, lowland dry,
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and lowland mesic habitat. In addition,
ordnance-induced fires have
periodically occurred on Hawaii’s
military installations, including the
Army’s PTA, and are considered an
ongoing threat to the montane dry forest
habitat that supports H. anthracinus
(The Center for Environmental
Management of Military Lands 2002,
Appendix 1 pp. 1–6; USFWS 2004, p.
110). Fire threatens the seven Hylaeus
species by destroying the native plant
species and communities on which the
bees depend and opening up habitat for
increased invasion by nonnative plants.
Fire can destroy dormant seeds of native
plants as well as the plants themselves.
Successive fires that burn farther and
farther into native habitat destroy native
plants and remove habitat for native
plant and animal species by altering
microclimate conditions favorable to
nonnative plants. Nonnative plant
species most likely to be spread as a
consequence of fire are those that (1)
produce a high fuel load; (2) are adapted
to survive and regenerate after fire; and
(3) establish rapidly in newly burned
areas. Grasses (particularly those that
produce mats of dry material or retain
a mass of standing dead leaves) that
invade native forests and shrublands
provide fuels that allow fire to burn
areas that would not otherwise easily
burn, including even the edges of wetter
forests (Fujioka and Fujii 1980, in
Cuddihy and Stone 1990, p. 93;
D’Antonio and Vitousek 1992, pp. 70,
73–74; Tunison et al. 2002, p. 122).
Native woody plants may recover from
fire to some degree, but fire tips the
competitive balance toward nonnative
species (National Park Service 1989, in
Cuddihy and Stone 1990, p. 93).
For example, on a post-burn survey at
Puuwaawaa on the island of Hawaii, an
area of native Diospyros forest with
undergrowth of the nonnative grass
Pennisetum setaceum, Takeuchi noted
‘‘no regeneration of native canopy is
occurring within the Puuwaawaa burn
area’’ (Takeuchi 1991, p. 2). Takeuchi
also stated, ‘‘burn events served to
accelerate a decline process already in
place, compressing into days a sequence
which would ordinarily have taken
decades’’ (Takeuchi 1991, p. 4). The
author concluded that in addition to
increasing the number of fires, the
nonnative Pennisetum acted to suppress
establishment of native plants after a
fire (Takeuchi 1991, p. 6).
There have been several recent fires
on Oahu that have impacted rare or
endangered species in coastal, lowland
dry, and mesic habitats. Between 2004
and 2005, wildfires burned more than
360 ac (146 ha) of mesic habitat in
Honouliuli Preserve, home to more than
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90 rare and endangered plants and
animals, and located along the
windward side of the Waianae
Mountains (The Nature Conservancy, in
litt. 2005). In 2006, a fire at Kaena Point
State Park burned 60 ac (24 ha) and
encroached on endangered plants in
Makua Military Training Area. The area
that burned in this fire is near the Kaena
Point NAR, where two of the yellowfaced bees (Hylaeus anthracinus and H.
longiceps) in this finding are still known
to occur. In 2007, there was a significant
fire in lowland dry and mesic habitat at
Kaukonahua that crossed 12 gulches,
eventually encompassing 5,655 ac
(2,289 ha), negatively impacting seven
endangered plant species. Occurrences
of three of the species were extirpated
as a result of the fire. The Kaukonahua
fire also provided pathways for
nonnative ungulates (cattle, goats, and
pigs) to access previously undisturbed
areas. This fire opened gaps in
previously densely vegetated areas
allowing the growth of the invasive
grass Panicum maximum (guinea grass),
which is also used as a food source by
cattle and goats. An area infested by
guinea grass burned, and the grass
resprouted blades over 2 feet in length
only 2 weeks after the fire (U.S. Army
Garrison 2007, p. 3). In 2009, there were
two smaller fires which burned 200 ac
(81 ha) at Manini Pali (Kaena Point State
Park), and 3.8 ac (1.5 ha) at Makua Cave
(at the mouth of Makua Valley). These
examples of recent fires illustrate
nonnative grass invasion leads to grass/
fire cycles that convert native vegetation
to grassland (D’Antonia and Vitousek
1992, p. 77)
Several areas in the State of Hawaii,
including some areas containing
Hylaeus spp. habitat sites, are currently
loosely addressed under fire
management plans. For example, in
2003, the Army completed an Integrated
Wildland Fire Management Plan
(WFMP) for all of its Oahu training
installations. This plan is currently
being updated (U.S. Army 2009, pp. 4–
73). The goal of the WFMP is to reduce
the threat of wildfire that adversely
affects listed and other rare species.
Although none of the Oahu yellowfaced bees are known from military
lands, at least one species, H. kuakea,
occurs on lands roughly adjacent to
military lands and which could be
impacted by fires caused by military
activities, or conversely, could benefit
from activities to suppress and control
origination of fires either on or adjacent
to military lands.
Additionally, DOFAW maintains a
fire management program tasked with
fire suppression activities targeted
toward the protection of watershed
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areas, forest reserves, public hunting
areas, wildlife and plant sanctuaries,
and NARS. Their activities include the
maintenance of fire break roads, signage,
and helicopter dip tanks; active fire
control during fire outbreak; controlled
burns when and where deemed
necessary; fire training efforts, including
education; and maintenance of a State
fire management program Web site
(https://www.state.hi.us/dlnr/dofaw/
fmp). According to their Web site,
DOFAW is involved in the protection of
3,360,000 acres Statewide, which is
approximately 81percent of the State’s
land area.
In summary, while we are aware of
fire management in some areas of the
State, including some Hylaeus spp.
habitat sites, there is evidence that the
repeated outbreak of fire within
Hawaii’s native coastal, lowland dry,
and lowland mesic forests often leads to
the irrevocable conversion of native to
nonnative habitat (i.e., nonnative plant
species). These nonnative habitats are
unsuitable for nesting and foraging by
the seven Hylaeus bees. Therefore, we
conclude fire is a significant ongoing
threat to the habitat of all seven species
of Hylaeus bees in coastal, lowland dry,
and lowland mesic habitat.
Habitat Destruction and Modification by
Recreational Activities
Some of the best habitat areas for
Hylaeus species are also popular
recreational sites, particularly those
areas located within coastal habitat
(Magnacca 2007a, p. 180). Suitable
remaining habitat for H. anthracinus
and H. longiceps are also popular hiking
areas, including coastal sites such as
Kaena Point (on Oahu); the Mahaiula
section of Kekaha Kai State Park,
Makalawena, Mokuauia, and Kalauna
Bay (on the island of Hawaii); and Kahu,
Polihua Road, and Shipwreck Beach on
Lanai. Human impacts at recreational
sites can include removal or trampling
of vegetation on or near trails and the
compaction of vegetation by off-road
vehicles (Magnacca 2007a, p. 180). None
of these areas, however, are known to be
currently impacted by recreational
activities (Magnacca pers. comm. 2010).
In summary, while trampling and
compaction of vegetation from human
activities may negatively impact the
habitat of some populations of the seven
Hylaeus bees, we have no basis to
conclude these impacts would be at a
scale that represents a threat to the
seven Hawaiian yellow-faced bees.
While some areas, particularly coastal
sites, are undoubtedly popular
recreational sites, we believe this is a
local rather a rangewide problem for
each of the seven species. Therefore, we
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conclude that recreational activities are
not a threat to the seven yellow-faced
bees at this time.
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Habitat Destruction and Modification by
Hurricanes and Drought
Stochastic (random, naturally
occurring) events, such as hurricanes
and drought, can alter or degrade the
habitat of Hawaiian Hylaeus bees
directly by modifying and destroying
native coastal and lowland dry and
mesic habitats (e.g., by mechanical
damage to vegetation). Indirect effects
include creating disturbed areas
conducive to invasion by nonnative
plants, which out-compete the native
plants used by the bees for foraging of
nectar and pollen. We presume these
events also alter microclimatic
conditions (e.g., opening the tree canopy
leading to an increase in habitat
temperature, soil erosion, and
decreasing soil moisture) so that the
habitat no longer supports the native
host plants necessary to the Hylaeus
bees for nectar and pollen foraging, as
well as nesting.
Hurricanes affecting Hawaii were only
rarely reported from ships in the area
from the 1800s until 1949. Between
1950 and 1997, 22 hurricanes passed
near or over the Hawaiian Islands, 5 of
which caused serious damage (Businger
1998, pp. 1–2). In November 1982,
Hurricane Iwa struck the Hawaiian
Islands, with wind gusts exceeding 100
miles per hour (mph) (161 kilometers
per hour (kph)), causing extensive
damage, especially on the islands of
Niihau, Kauai, and Oahu (Businger
1998, pp. 2, 6). Many forest trees were
destroyed (Perlman 1992, pp. 1–9),
which opened the canopy and
facilitated the invasion of nonnative
plants (Kitayama and Mueller-Dombois
1995, p. 671). Habitat alteration and
degradation by nonnative plants is a
threat to the habitat of each of the seven
yellow-faced bees addressed in this
finding, as described in the Habitat
Destruction and Modification by
Nonnative Plants section above. In
September 1992, Hurricane Iniki, a
category 4 hurricane with maximum
sustained wind speeds recorded at 140
mph (225 kph), passed directly over the
island of Kauai and close to the island
of Oahu, causing significant damage to
areas along Oahu’s southwestern coast
(Barber’s Point or Kalaeloa, through
Kaena) (Blake et al. 2007, p. 20), where
populations of two of the seven bee
species (H. anthracinus and H.
longiceps) are found. Damage by future
hurricanes could further decrease the
remaining native-plant-dominated
habitat areas that support the yellow-
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faced bees (Bellingham et al. 2005, p.
681).
All seven of the Hylaeus bees may
also be affected by temporary habitat
loss (e.g., desiccation of habitats, die-off
of host plants) associated with droughts,
which are not uncommon on the
Hawaiian Islands. Between 1860 and
2002, the Hawaiian Islands were
affected by approximately 49 periods of
drought (Giambelluca et al. 1991, pp. 3–
4; Hawaii Commission on Water
Resource Management 2009a and
2009b). These drought events lead to an
increase in the number of forest and
brush fires (Giambelluca et al. 1991, p.
v), causing a reduction of native plant
cover and habitat (D’Antonio and
Vitousek 1992, pp. 77–79). With
populations that have already been
severely reduced in both abundance and
geographic distribution, and particularly
in the case of H. hilaris, with only one
known population, even such a
temporary loss of habitat can have a
severe negative impact on the species if,
for example, the host plants for nectar
and pollen foraging are lost for one or
more seasons. Because small
populations are demographically
vulnerable to extinction caused by
random fluctuations in population size
and sex ratio, stochastic events such as
hurricanes pose the threat of immediate
extinction of a species with a very small
and geographically restricted
distribution such as the seven species of
Hawaiian yellow-faced bees (Lande
1988).
In summary, natural disasters, such as
hurricanes and drought, represent a
significant threat to coastal and lowland
dry and mesic habitats and the seven
Hylaeus species addressed in this
finding, endangering their chances for
conservation and recovery. These types
of events are known to cause significant
habitat damage, and because the species
addressed in this finding now persist in
low numbers or occur in restricted
ranges, they are more vulnerable to
these events and less resilient to such
habitat disturbances. Hurricanes and
drought, even though unpredictable,
have been and are expected to continue
to be threats to the Hawaiian yellowfaced bees, and they therefore pose
immediate and ongoing threats to the
seven Hylaeus species and their habitat.
Habitat Destruction and Modification by
Climate Change
Climate change will be a particular
challenge for biodiversity because the
interaction of additional stressors may
push species beyond their ability to
survive (Lovejoy et al. 2005, pp. 325–
326). The synergistic implications of
climate change and habitat
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fragmentation are the most threatening
facet of climate change for biodiversity
(Lovejoy et al. 2005, p. 4). The
magnitude and intensity of the impacts
of global climate change and increasing
temperatures on native Hawaiian
ecosystems are unknown; we are not
aware of climate change studies
specifically related to the coastal and
lowland habitat areas occupied by the
seven Hylaeus bees, or to other Hylaeus
bee species. Based on the best available
information, climate change impacts
could include the loss of native plant
species that comprise the habitats in
which the seven Hylaeus bees occur
(Pounds et al. 1999, pp. 611–612; Still
et al. 1999, p. 610; Benning et al. 2002,
pp. 14,246 and 14,248); however,
because there have been no climate
change studies looking at effects to
coastal and lowland habitat, we have no
way of predicting the amount or extent
of any such possible habitat loss.
Because the host plant habitat of the five
coastal species in this finding are
outside of the tidal and immediate near
shore zone, we do not expect any direct
effects to their habitat from sea level rise
itself.
In addition, the seven yellow-faced
bees may be vulnerable to changes in
precipitation caused by global climate
change. However, future changes in
precipitation are uncertain because they
˜
depend in part on how El Nino (a
disruption of the ocean atmospheric
system in the tropical Pacific having
important global consequences for
weather and climate) might change, and
reliable projections of changes in El
˜
Nino have yet to be made (Benning et
al. 2002, pp. 14,248–14,249). Oki (2004,
p. 4) has noted long-term evidence of
decreased precipitation and stream flow
in the Hawaiian Islands, based upon
evidence collected by stream gauging
stations. This long-term drying trend,
˜
coupled with periodic El Nino-caused
drying events, has created a pattern of
severe and persistent stream dewatering
events (D. Polhemus, in litt 2008, p. 26).
Future changes in precipitation and the
forecast of those changes are highly
uncertain because they depend, in part,
˜
˜
on how the El Nino-La Nina (a different
disruptive extreme weather and climate
˜
pattern that can alternate with El Nino)
weather cycle might change (Hawaii
Climate Change Action Plan 1998, pp.
2–10).
If precipitation is significantly
reduced, the seven yellow-faced bees
may be among the species most
vulnerable to extinction, with possible
impacts expected to include habitat loss
and alteration or changes in disturbance
regimes (e.g., storms and hurricanes), in
addition to possible direct physiological
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stress of an unknown nature, which
could potentially cause the species to
seek out less suitable habitats as their
preferred habitats become degraded.
The probability of a species going
extinct as a result of these factors
increases when ranges are restricted,
habitat decreases, and population
numbers decline (Intergovernmental
Panel on Climate Change 2007, p. 8).
Such is the case for each of the seven
yellow-faced bees, which are
characterized by limited climatic ranges
and restricted habitat requirements,
small population size, and low number
of individuals. However, without
reliable predictions of the amount and
extent of anticipated precipitation
change, we are unable to determine
whether precipitation changes would
result in negative impacts to any of the
seven yellow-faced bees at this time.
In summary, the seven Hylaeus bees,
like most insects, are presumed to have
limited environmental tolerances. They
also have limited ranges and restricted
habitat requirements (Daly and
Magnacca 2003, p. 11). Four species (H.
facilis, H. hilaris, H. kuakea, and H.
mana) have small population sizes (i.e.,
a limited number of populations
restricted to relatively small habitat
sites), and low numbers of individuals.
The projected effects of global climate
change and increasing temperatures on
the seven Hawaiian yellow-faced bees
would likely be related to changes in
microclimatic conditions in their
habitats. These changes may also lead to
the loss of native plant species due to
direct physiological stress, the loss or
alteration of habitat, increased
competition from nonnative bee species,
and changes in disturbance regimes
(e.g., fire, storms, and hurricanes).
Therefore, we believe all seven species
will be exposed to projected
environmental impacts that may result
from changes in climate, and
subsequent impacts to their habitats
(Pounds et al. 1999, pp. 611–612; Still
et al. 1999, p. 610; Benning et al. 2002,
pp. 14,246 and 14,248), and we do not
anticipate a reduction in this ongoing
threat any time in the near future.
However, because the specific and
cumulative effects of climate change on
these seven species are presently
unknown, we are not able to determine
the magnitude of this potential threat
with confidence or precision.
Summary of Factor A
The seven species of Hawaiian
yellow-faced bees are dependent upon
the persistence of native Hawaiian
plants and their increasingly rare
associated habitat types, particularly
coastal, lowland dry, and lowland mesic
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areas. As identified above in our Factor
A analysis, the native habitats on which
the Hylaeus bees depend have been
drastically directly altered during the
last century, with many areas either
converted for development or
agriculture, or indirectly altered due to
the effects of nonnative ungulates,
nonnative plants, and fire. Habitat
conversion and loss of host plants, and
other stochastic events (e.g., hurricanes
and drought), are all contributing factors
to the present and threatened
destruction, modification, and
curtailment of the habitat and range of
the seven Hawaiian yellow-faced bees.
Land conversion and fragmentation of
remaining coastal, lowland dry, and
lowland mesic habitat is continuing
throughout these species’ known ranges,
particularly due to the effects of feral
ungulates, fire, and nonnative plants.
We anticipate habitat conversion and
fragmentation to continue, and likely
increase, throughout their known
ranges. As discussed above, at least five
of the seven bees have experienced
significant habitat losses. It is
reasonable to presume the substantial
reduction in lowland mesic habitat has
similarly impacted the populations of
Hylaeus kuakea and H. mana
(Magnacca in litt. 2011, p. 78). As more
habitats become unsuitable, we expect
their population declines to continue or
accelerate.
We have evaluated the best scientific
and commercial information available
regarding the present or threatened
destruction, modification, or
curtailment of the seven Hawaiian
yellow-faced bees’ habitat or range.
Based on the current and ongoing
habitat issues identified, their
synergistic effects, and their likely
continuation, we have determined this
factor poses a significant threat to
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana.
Available Conservation Measures
Some historic and current collection
localities are protected from
development, urbanization, and
conversion to agriculture by Federal,
State, or private agencies: one of two
known populations of H. facilis and two
of three known populations of H.
anthracinus occur at Kalaupapa NHP on
Molokai; three species (H. anthracinus,
H. assimulans, and H. kuakea) occur in
the State’s Kaena Point NAR (Oahu),
Kanaio NAR (Maui), West Maui NAR,
and the recently acquired Honouliuli
Preserve (Oahu); and three species (H.
anthracinus, H. hilaris, and H.
longiceps) are found on TNC’s
Moomomi Preserve. These areas are
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actively managed to restore native
habitat and to reduce or eliminate many
of the common threats to the native
plant communities found there,
including feral ungulates and wildfire.
However, existing regulatory
mechanisms are inadequate to provide
the necessary active management
needed to protect the habitat of the
populations outside of these protected
TNC, NHP or NAR areas (see discussion
under Factor D, below). Conservation of
the seven Hylaeus bees will 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).
Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
We are unaware of any collections of
the seven yellow-faced bees by
recreational or insect enthusiast
collectors. However, insect collecting is
a valuable component of research,
including taxonomic work, and is often
necessary to document the existence of
populations and population trends.
Based on comments received in
response to the 90-day finding, six of
the yellow-faced bees are not believed to
be particularly vulnerable to overcollection; however, one species (H.
hilaris) may be vulnerable (Magnacca, in
litt. 2010, p. 2). This species is a
cleptoparasite on other rare bees, and
has an inherently smaller population
size and lower reproductive rate than
most Hylaeus species, including the
other six species in this finding.
However, as both sexes of H. hilaris are
readily recognizable to Hylaeus
researchers, experts believe there will be
little need to retain individuals
collected during field surveys in the
future (Magnacca, in litt. 2010, p. 2).
Additionally, while this species is
known from only one population site,
the area where this population is found
occurs within the Moomomi Preserve
and is actively managed by TNC for
common habitat threats such as feral
ungulates, wild fire, and nonnative
plant species.
Therefore, we find that overutilization
for commercial, recreational, scientific,
or educational purposes is not a threat
to Hylaeus anthracinus, H. assimulans,
H. facilis, H. kuakea, H. longiceps, and
H. mana because we could find no
evidence they are being collected by
insect collection enthusiasts or overcollected by researchers for scientific
purposes. We examined whether H.
hilaris was directly or indirectly
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vulnerable to over-collection due to its
small population size (one known
location), low reproductive rate, and
biological dependence upon other rare
Hylaeus host species. However, as both
sexes are easily recognizable in the field
and it does not collect pollen (which
differentiates it from all other species),
researchers believe there is little reason
to retain individuals observed during
surveys (Magnacca, in litt. 2010, p. 2).
Therefore, we find over-collection of H.
hilaris is not a threat to this species.
Factor C. Disease or Predation
Disease
We are not aware of any information
indicating disease presents a threat to
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, or H. mana. Therefore, based
on the best available information, we do
not find that disease is a threat to the
seven Hawaiian yellow-faced bees.
Predation
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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. In
this study, nests of Nesoprosopis sp., an
endemic ground-nesting bee, could not
be found in ant-infested plots but were
commonly encountered in ant-free sites
of the same habitat. Nesoprosopis was
reduced to a subgenus of Hylaeus in
1923 (Meade-Waldo 1923, p. 1). Ants
are not a natural component of Hawaii’s
arthropod fauna, and the native Hylaeus
species of the islands evolved in the
absence of predation pressure from ants.
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 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,
these attributes allow some ants to
destroy otherwise geographically
isolated populations of native
arthropods (Nafus 1993, pp. 19, 22–23).
Ants have not been observed preying
upon any of the seven species addressed
in this finding. However, at least one or
more of the most aggressive and
widespread species (discussed below)
occur in every known population site of
the seven Hylaeus species and are
presumed to be a serious threat due to
the impact of predation.
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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 (NCN),
and Solenopsis geminata (NCN).
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). Due to their preference for
drier habitat sites, ants are more likely
to occur in high densities in the coastal,
dry, and mesic habitat currently
occupied by the seven bees (Reimer
1994, p. 12).
The big-headed ant originated in
central Africa (Krushelnycky et al. 2005,
p. 24) and was first reported in Hawaii
in 1879 (Krushelnycky et al. 2005, p.
24). This species is considered one of
the most invasive and widely
distributed ants in the world
(Krushelnycky et al. 2005, p. 5). In
Hawaii, this species is the most
ubiquitous ant species found, from
coastal to mesic habitat up to 4,000 ft
(1,219 m) in elevation, including within
the habitat areas of the seven Hylaeus
species addressed in this finding. With
few exceptions, native insects have been
eliminated in habitats where the bigheaded ant is present (Perkins 1913, p.
xxxix; Gagne 1979, p. 81; Gillespie and
Reimer 1993, p. 22). Consequently, bigheaded ants represent a threat to
populations of all seven Hylaeus bee
species in coastal to dry and mesic areas
Hawaii, Lanai, Maui, and Oahu (Reimer
1993, p. 14; Reimer 1994, p. 17; Daly
and Magnacca 2003, pp. 9–10).
The long-legged ant appeared in
Hawaii in 1952, and now occurs on
Hawaii, Kauai, Maui, and Oahu (Reimer
et al. 1990, p. 42; https://www.antweb.org
2011). It 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). Although surveys have not been
conducted to ascertain this species’
presence in each of the known habitat
sites occupied by the seven Hylaeus
species addressed in this finding, we
may presume that the long-legged ant
likely occurs within some of the
identified population sites based upon
anecdotal evidence of their expanding
range and their preference (as indicated
where the species is most commonly
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collected) for coastal and dry forest
habitats (antweb.org 2011). Direct
observations indicate Hawaiian
arthropods are susceptible to predation
by this species; Gillespie and Reimer
(1993, p. 21) and Hardy (1979, pp. 37–
38) documented the complete
extirpation of several native insects
within the Kipahulu area on Maui after
this area was invaded by the long-legged
ant. Lester and Tavite (2004, p. 391),
found that long-legged ants in the
Tokelau Atolls (New Zealand) can form
very high densities in a relatively short
period of time with locally serious
consequences for invertebrate diversity.
Densities of 3,600 individuals collected
in pitfall traps within a 24-hour period
were observed, as well as predation
upon invertebrates ranging from crabs to
other ant species. On Christmas Island
in the Indian Ocean, numerous studies
have documented the range of impacts
to native invertebrates, including the
red land crab (Gecarcoidea natalis), as
a result of predation by supercolonies of
the long-legged ant (Abbott 2006, p.
102). Long-legged ants have the
potential as predators to profoundly
affect the endemic insect fauna in
territories they occupy. Studies
comparing insect populations at
otherwise similar ant-infested and antfree sites found extremely low numbers
of large endemic noctuid moth larvae
(Agrostis spp. and Peridroma spp.) in
ant-infested areas. Nests of groundnesting cottelid bees (Nesoprosopis
spp.) were eliminated from ant-infested
sites (Reimer et al. 1990, p. 42).
Although only cursory observations
exist in Hawaii (Reimer et al. 1990, p.
42), we believe long-legged ants are a
threat to populations of all seven
yellow-faced bees, in dry to mesic areas
within their elevation ranges.
Solenopsis papuana is the only
abundant, aggressive ant that has
invaded intact mesic to wet forest, as
well as coastal and lowland dry
habitats. This species occurs 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). Although surveys have not been
conducted to ascertain this species’
presence in each of the known habitat
sites occupied by the seven Hylaeus
species addressed in this finding,
because of this species’ expanding range
and its widespread occurrence in
coastal, dry lowland, and mesic
habitats, it may threaten populations of
all seven Hylaeus bees with predation
pressure on the islands of Hawaii,
Kahoolawe, Lanai, Maui, and Oahu over
2,000 ft (600 m) in elevation (Reimer et
al. 1990, p. 42; Reimer 1993, p. 14).
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Like Solenopsis papuana, S. geminata
is also considered a significant threat to
native invertebrates (Gillespie and
Reimer 1993) and occurs on all the main
Hawaiian Islands (Reimer et al. 1990;
Nishida 1997). Found in drier areas of
the Hawaiian Islands, it has displaced
Pheidole megacephala as the dominant
ant in some areas (Wong and Wong
1988, p. 175). Known to be a voracious
nonnative predator in many areas to
where it has spread, the species was
documented to significantly increase
fruit fly mortality in field studies in
Hawaii (Wong and Wong 1988, p. 175).
In addition to predation, S. geminata
workers tend honeydew-producing
members of the Homoptera suborder,
especially mealybugs, which can impact
plants directly and indirectly through
the spread of disease (Manaaki
Whenua—Landcare Research 2011:
https://www.landcareresearch.co.nz/
research/biocons/invertebrates/Ants/
invasive_ants/solgem_info.asp).
Solenopsis geminata was included
among the eight species ranked as
having the highest potential risk to New
Zealand in a detailed pest risk
assessment for the country (Global
Invasive Species Database 2011: https://
www.issg.org/database/species/
ecology.asp?si=169&fr=1&
sts=&lang=EN), and is included as one
of five ant species listed among the ‘‘100
of the World’s Worst invaders’’
(Manaaki Whenua—Landcare Research
2011: https://www.landcareresearch.
co.nz/research/biocons/invertebrates/
Ants/invasive_ants/solgem_info.asp).
Although surveys have not been
conducted to ascertain this species’
presence in each of the known habitat
sites occupied by the seven Hylaeus
species addressed in this finding,
because of this species’ expanding range
and its widespread occurrence in
coastal, dry lowland, and mesic
habitats, it may threaten populations of
all seven Hylaeus bees with predation
pressure on the islands of Hawaii,
Kahoolawe, Lanai, Maui, and Oahu from
sea level up to 1,000 ft (300 m) in
elevation (Wong and Wong 1988, p.
175).
The Hylaeus egg, larvae, and pupal
stages are more vulnerable to attack by
ants than the mobile adult bees (Daly
and Magnacca 2003, p. 10). Invasive
ants have severely impacted groundnesting Hylaeus species in particular
(Cole et al. 1992, pp. 1317, 1320;
Medeiros et al. 1986, pp. 45–46),
because their nests are easily accessible
and in or near the ground. Because
Hylaeus anthracinus, H. facilis, H.
hilaris, and H. longiceps are believed to
be ground-nesting species, they may
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also be more susceptible to ant
predation (Magnacca 2005g, p. 2).
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). 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 (Reimer
1994, pp. 17–18; Daly and Magnacca
2003, pp. 9–10). Although the primary
impact of ants 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
bees indirectly as well, by preying on
seeds of native plants, thereby reducing
the plant’s recruitment and fecundity
(Bond and Slingsby 1984, p. 1,031).
Several studies (Krushelnycky 2005, p.
9; Lach 2008, p. 155) suggest a serious
ecosystem-level effect of invasive ants
on pollination. Where ranges overlap,
ants compete with native pollinators
such as Hylaeus bees 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 bees that regularly collect
pollen from flowers of Metrosideros
polymorpha were entirely absent from
trees with flowers exposed to foraging
by big-headed ants.
The rarity or disappearance of native
Hylaeus species from historically
documented localities over the past 100
years (including the seven Hawaiian
yellow-faced bee species) is due to a
variety of factors. Although we have no
direct information that conclusively
correlates the decrease in populations of
these seven Hylaeus bees due to the
establishment of nonnative ants, severe
predation of other Hylaeus species by
ants has been documented, resulting in
clear reductions in populations. We
expect similar predation impacts to
these seven Hylaeus bees to continue as
a result of the widespread presence of
ants throughout the Hawaiian Islands,
their highly efficient and non-specific
predatory behavior, and their ability to
quickly disperse and establish new
colonies. Therefore, we conclude that
predation by nonnative ants represents
a serious threat to the continued
existence of H. anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
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kuakea, H. longiceps, and H. mana now
and into the future.
Predation by Nonnative Western Yellow
Jacket Wasps
The western yellow jacket wasp
(Vespula pensylvanica) is a potentially
serious threat to the seven Hylaeus bees
(Gambino et al. 1987, p. 170; Wilson et
al. 2009, pp. 1–5). The western yellow
jacket 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 yellow
jacket 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 approximately 2,000 and
3,500 ft (approximately 600 and 1,050
m) in elevation (Gambino et al. 1990, p.
1,088), although they can also occur at
sea level. The western yellow jacket
wasp is known to be an aggressive,
generalist predator (Gambino et al.
1987, p. 170), and has been documented
preying upon Hawaiian Hylaeus species
(although not specifically upon any of
the seven species addressed in this
finding) (Wilson et al. 2009, p. 2).
However, predation by the western
yellow jacket wasp is a potentially
significant threat to all seven of the
yellow-faced bees because of the wasp’s
presence in habitat occupied by the
seven Hylaeus bees combined with their
small population sizes. This may
present a particular threat to H. facilis,
H. hilaris, H. kuakea, and H. mana,
because each species is known from
only two or fewer sites. It has been
suggested the western yellow jacket
wasp may compete for nectar with
Hylaeus species, but we have no
information to suggest this represents a
threat to the seven Hylaeus bees.
Predation by Nonnative Parasitoid
Wasps
Native and nonnative parasitoid
wasps are known to parasitize some
Hylaeus species on Oahu (although not
upon any of the seven species addressed
in this finding), and may pose a threat
to five of the seven yellow-faced bees
(H. anthracinus, H. facilis, H. kuakea,
H. longiceps, and H. mana) (Daly and
Magnacca 2003, p. 10) because they
occur on Oahu as well. While the
available information indicates some
Oahu Hylaeus larvae have been
parasitized (and subsequently killed) by
parasitoid wasps from the Encyrtidae
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and Eupelmidae families, it is unknown
whether these wasps also utilize H.
anthracinus, H. facilis, H. kuakea, H.
longiceps, and H. mana as nutritional
hosts for their larvae (Daly and
Magnacca 2003, p. 98). We are
concerned that H. anthracinus, H.
facilis, H. kuakea, H. longiceps, and H.
mana may be exposed to wasp
parasitism, but we are unaware of any
information to indicate this is a threat
to these five Hylaeus bees.
Summary of Factor C
We do not find evidence that disease
is currently impacting the seven
Hawaiian yellow-faced bees, nor do we
have information to indicate disease
outbreaks will occur in the future.
Although we have no direct information
that conclusively correlates the decrease
in populations of these seven Hylaeus
bees due to the establishment of western
yellow jacket wasps, severe predation of
other Hylaeus species by yellow jacket
wasps has been documented, resulting
in clear reductions in populations. We
expect similar predation impacts to
these seven Hylaeus bees to continue as
a result of the widespread presence of
yellow jacket wasps in many areas
throughout the Hawaiian Islands, their
highly efficient and non-specific
predatory behavior, and their ability to
quickly disperse and establish new
colonies.
While we are concerned that Hylaeus
anthracinus, H. facilis, H. kuakea, H.
longiceps, and H. mana may be
threatened by wasp parasitism on Oahu,
we are unaware of any information to
indicate this is a threat to these five
Hylaeus bees at this time, or that it is
likely to become so in the future. The
presence of nonnative ants in nearly all
lowland habitat historically and
currently occupied by the seven
Hylaeus bees, combined with the near
extirpation of native insects in these
areas, suggest predation by nonnative
ants is a serious threat to the seven
Hawaiian yellow-faced bees.
Observations and reports have
documented that ants are particularly
destructive predators because of their
high densities, broad ranges of diet, and
ability to establish new colonies in
otherwise geographically isolated
locations because the reproductive adult
ants are able to fly. Because the ranges
of the big-headed ant, long-legged ant,
Solenopsis geminata, and Solenopsis
papuana overlap the ranges of the seven
Hylaeus bees, and based on their
observed predatory behavior at other
locations where they occur, these
nonnative predators represent an
imminent and serious threat to H.
anthracinus, H. assimulans, H. facilis,
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H. hilaris, H. kuakea, H. longiceps, and
H. mana. Unless these aggressive,
nonnative ant predators are eliminated
or controlled, we expect this threat to
continue or increase. Furthermore, a
decrease in the amount and distribution
of suitable host plants for foraging could
indirectly impact these seven species by
forcing them to seek less optimal, but
predator-free, foraging sites.
Factor D. The Inadequacy of Existing
Regulatory Mechanisms
Currently, there are no Federal, State,
or local laws, treaties, or regulations that
specifically conserve or protect the
seven Hylaeus bee species from the
threats described in this finding. There
are some regulations that potentially
address the threats posed by introduced,
nonnative species; these are discussed
below.
Inadequate Protection from Nonnative
Ungulates
Nonnative ungulates pose a major
ongoing threat to the seven Hylaeus bees
through destruction and degradation of
their habitat. Although some public
hunting areas are fenced to prevent the
incursion of nonnative ungulates, there
are currently no Federal, State, or local
laws, treaties, or regulations that
adequately address the threats from
nonnative ungulates to the seven
yellow-faced bees’ terrestrial habitat.
The existing regulatory mechanisms do
not address the threats from nonnative
ungulates to the seven yellow-faced bee
species or their habitat. The absence of
regulatory mechanisms exacerbates the
threats discussed under Factor A.
Inadequate Protection from Introduction
of Nonnative Species
The Hawaii Department of
Agriculture (HDOA) is the lead State
agency in protecting Hawaii’s
agricultural and horticultural industries,
animal and public health, natural
resources, and environment from the
introduction of nonnative, invasive
species (HDLNR 2003, p. 3–10). While
there are several State agencies (Hawaii
Department of Agriculture (HDOA),
Hawaii Department of Lands and
Natural Resources (HDLNR), Hawaii
Department of Health (HDOH))
authorized to prevent the entry of pest
species into the State, the existing
regulations are inadequate for the
reasons discussed in the sections below.
In 1995, a partnership, Coordinating
Group on Alien Pest Species (CGAPS),
comprised primarily of managers from
every major Federal, State, county, and
private agency and organization
involved in invasive species work in
Hawaii, was formed in an effort to
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influence policy and funding decisions,
improve communication, increase
collaboration, and promote public
awareness (CGAPS 2009). This group
facilitated the formation of the Hawaii
Invasive Species Council (HISC), which
was created by gubernatorial executive
order in 2002 to coordinate local
initiatives for the prevention and
control of invasive species by providing
policy-level direction and planning for
the State departments responsible for
invasive species issues. In 2003, the
governor signed into law Act 85, which
conveys statutory authority to the HISC
to coordinate approaches among the
various State and Federal agencies, and
international and local initiatives, for
the prevention and control of invasive
species (HDLNR 2003, p. 3–15; HISC
2009a; Haw. Rev. Stat. section 194–2(a)).
Some of the recent priorities for the
HISC include interagency efforts to
control nonnative species such as the
plants Miconia calvescens (miconia) and
Cortaderia sp. (pampas grass), coqui
frogs (Eleutherodactylus coqui), and
ants (HISC 2009). However, in October
2009, HISC approved a 2010 budget
that, due to a tighter economy in Hawaii
and anticipated budget cuts in State
funding support, resulted in a 50
percent reduction in funding with an
anticipated setback in conservation
achievements and the loss of
experienced, highly trained staff (HISC
2009b).
Inadequate Regulatory Control of
Nonnative Invertebrate Species
As noted above (see Factor C, Disease
and Predation), predation by nonnative
ants and the nonnative yellow jacket
wasp is a potentially significant threat
to the seven species. Commercial
shipping and air cargo, as well as
biological introductions to Hawaii, have
resulted in the establishment of over
3,372 species of nonnative insects
(Howarth 1990, p. 18; Staples and
Cowie 2001, p. 52), with an estimated
continuing establishment rate of 20 to
30 new species per year (Beardsley
1962, p. 101; Beardsley 1979, p. 36;
Staples and Cowie 2001, p. 52). The
prevention and control of introduced
pest species in Hawaii is the
responsibility of Hawaii State
government and Federal agencies, along
with a few private organizations. Even
though these agencies have regulations
and some controls in place, complete
control of introduced pest species is
difficult to achieve. Consequently, the
introduction and movement of
nonnative invertebrate pest species,
including nonnative ants and yellow
jacket wasps, between islands and from
one watershed to the next, continues.
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Inadequate Regulatory Control of
Nonnative Plant Species
Nonnative plants destroy and modify
habitat throughout the ranges of each of
the seven Hylaeus species addressed in
this 12-month finding. As such, they
represent a significant and immediate
threat to each of these species. In
addition, nonnative plants have been
shown to out-compete native plants and
convert native-dominated plant
communities to nonnative plant
communities (see Factor A—Habitat
Destruction and Modification by
Nonnative Plants). The HDOA regulates
the import of plants into the State from
domestic origins under Hawaii State law
(Haw. Rev. Stat. Ch. 150A). While all
plants require inspection upon entry
into the State and must be ‘‘apparently
free’’ of insects and diseases, not all
plants require import permits. Parcels
brought into the State by mail or cargo
must be clearly labeled as ‘‘Plant
Materials’’ or ‘‘Agricultural
Commodities,’’ but, given budget
constraints and an insufficient number
of personnel, it is unlikely that all of
these parcels are inspected or monitored
prior to delivery in Hawaii. Shipments
of plant material into Hawaii must be
accompanied by an invoice or packing
manifest listing the contents and
quantities of the items imported,
although it is unclear if all of these
shipments are inspected or monitored
prior to delivery (HDOA 2009). There
are only 12 plant crops regulated
(H.A.R. chapter 4–70) to some degree:
sugarcane and grasses, pineapple and
other bromeliads, coffee, cruciferous
vegetables, orchids, banana, passion
fruit, pine, coconut, hosts of European
corn borer, palms, and hosts of
Caribbean fruit fly (HDLNR 2003, p. 3–
11). The HDOA also maintains the State
list of noxious weeds, and these plants
are restricted from entry into the State
except by permit from the HDOA’s Plant
Quarantine Branch.
Although the State has general
guidelines for the importation of plants,
and regulations are in place regarding
the plant crops mentioned above, the
intentional or inadvertent introduction
of nonnative plants outside the
regulatory process and movement of
species between islands and from one
watershed to the next continues, which
represents a threat to native flora and
fauna for the reasons described above.
In addition, government funding is
inadequate to provide for sufficient
inspection services and monitoring. One
study concluded plant importation laws
virtually ensure new invasive plants
will be introduced via the nursery and
ornamental trade, and outreach efforts
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cannot keep up with the multitude of
new invasive plants being distributed.
The author states the only thing widescale public outreach can do in this
regard is to let the public know new
invasive plants are still being sold, and
suggest that people should ask for
noninvasive or native plants instead (C.
Martin, in litt. 2007, p. 9).
On the basis of the above information,
existing regulatory mechanisms do not
adequately protect the seven Hylaeus
species from the threat of new
introductions of nonnative species, and
the continued expansion of nonnative
species populations on and between
islands and watersheds. Nonnative
species may directly compete with, prey
upon, or modify or destroy the habitat
of one or more of the seven yellow-faced
bees for food, space, and other necessary
resources. Because current Federal,
State, and local laws, treaties, and
regulations are inadequate to prevent
the introduction and spread of
nonnative species from outside the State
of Hawaii, as well as between islands
and watersheds, the threats from these
introduced species remain immediate
and significant due to an inadequacy of
existing regulatory mechanisms.
Summary of Inadequacy of Existing
Regulatory Mechanisms
We found that existing regulatory
mechanisms and agency policies do not
address the primary threats to the seven
yellow-faced bee species and their
habitat from nonnative ungulates. The
State’s current management of
nonnative game mammals does not
prevent the degradation and destruction
of habitat of Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana (see
discussion under Factor A).
We consider the threat from
inadequate regulatory mechanisms to be
immediate and significant for the
following reasons:
(1) Existing State and Federal
regulatory mechanisms are not
preventing the introduction and spread
of nonnative species between islands
and watersheds. Habitat-altering
nonnative plant species (Factor A) and
predation by nonnative animal species
(Factor C) pose major ongoing threats to
the seven Hylaeus species.
Because existing regulatory
mechanisms are inadequate to maintain
habitat for the seven species of Hylaeus
and to prevent the spread of nonnative
species, the inadequacy of existing
regulatory mechanisms is considered to
be a significant and immediate threat to
Hylaeus anthracinus, H. assimulans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana.
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Factor E. Other Natural or Manmade
Factors Affecting the Species’ Continued
Existence
Small Number of Populations and
Individuals
Species 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. 1,455; 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 seven Hylaeus bee species 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). According to
Magnacca (2007, p. 3), five species have
not been collected recently from one or
more islands from which they were
historically known, all seven species are
restricted to rare habitat, and two are
particularly rare and potentially
endangered. Hylaeus facilis and H.
hilaris have not been recently observed
at some historical collection sites; H.
facilis is currently known from two
populations, and H. hilaris is known
from only a single population. In
addition, H. kuakea, first collected in
1997, is only known from two
populations, and H. mana, just collected
in 2002, is known from a single
population. Although H. kuakea and H.
mana were only discovered relatively
recently, researchers believe these two
species were once more widespread
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when their lowland mesic habitat was
not highly fragmented and degraded by
invasive species, as is currently the case
(Magnacca in litt. 2011, p. 95). The
small number of populations known for
each of these four Hylaeus species
increases their risk of extinction due to
stochastic events such as hurricanes,
wildfires, or prolonged drought (Jones et
al. 1984, p. 209; Smith and Tunison
1992, p. 398).
The recurrence intervals for stochastic
events, for example, wildfires,
prolonged drought, and hurricanes,
cannot be predicted, which introduces
some uncertainty regarding potential
effects to H. facilis, H. hilaris, H.
kuakea, and H. mana (the four species
most at risk of the seven Hylaeus bees).
However, because Hylaeus hilaris is
cleptoparasitic and restricted to one
known population, it is at particularly
high risk of extinction because of the
rarity of its hosts and the fact it is the
most habitat-specific of all Hawaiian
bees (Magnacca 2007a, p. 181). The fact
that a species is potentially vulnerable
to stochastic processes does not
necessarily mean it is reasonably likely
to experience or have its status affected
by a given stochastic process within
timescales meaningful under the Act.
Because of their small number of
populations, negative impacts to H.
facilis, H. hilaris, H. kuakea, and H.
mana from hurricanes, wildfires, and
drought would be likely if these events
occur. Because these events have been
documented on Oahu and other
Hawaiian islands in the past, we believe
that they represent an ongoing threat to
these four species, although the specific
timing, location, or magnitude is
unknown. The threat from fire is
unpredictable, but omnipresent in
habitats that have been invaded by
nonnative, fire-prone grasses.
Hurricanes and drought conditions
present an ongoing and ever-present
threat, because they can occur at any
time, although the incidence and
magnitude of specific events is not
predictable.
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). Most nonnative bees inhabit areas
dominated by nonnative vegetation and
do not compete with native Hawaiian
bees for foraging resources (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
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(Hopper et al. 1996, p. 9; Daly and
Magnacca 2003, p. 13; Snelling 2003, p.
345).
The European honey bee was first
introduced to the Hawaiian Islands in
1875, and currently inhabits areas from
sea level to the upper tree line boundary
(Howarth 1985, p. 156). European honey
bees have been observed foraging on
Hylaeus host plants such as Scaevola
spp. and Sesbania tomentosa (Hopper et
al. 1996, p. 9; Daly and Magnacca 2003,
p. 13; Snelling 2003, p. 345). Although
we lack information indicating
Hawaiian Hylaeus populations have
declined because of competition with
European honey bees for nectar and
pollen, the European honey bee does
forage in Hylaeus spp. habitat and may
exclude Hylaeus spp. (Magnacca 2007,
p. 188; Lach 2008, p. 155). Hylaeus
species do not occur in native habitat
where there are large numbers of honey
bees, although the impact of moderate
populations of honey bees is not known
(Magnacca 2007, p. 188). Nonnative,
invasive bees are widely documented to
decrease nectar volumes and usurp
native pollinators (Lach 2008, p. 155).
There are also indications that
populations of the European honey bee
are not as vulnerable as Hylaeus bees to
predation by nonnative ant species (see
Factor C. Disease and Predation). Lach
(2008, p. 155) observed that Hylaeus
bees that regularly collect pollen from
the flowers of Metrosideros polymorpha
trees were entirely absent from trees
with flowers visited by the big-headed
ant, while visits by the European honey
bee were not affected. As a result, the
European honey bee may have a
competitive advantage over Hylaeus
spp., as it is not excluded by the bigheaded ant (Lach 2008, p. 155).
Other nonnative bees found in areas
of native vegetation include carpenter
bees (Ceratina species), Australian
colletid bees (Hylaeus albonitens), and
Lasioglossum impavidum (NCN)
(Magnacca 2007, p. 188). While it has
been suggested these nonnative bees
may impact native Hylaeus bees through
competition for pollen based on their
similar size and flower preferences,
there is no information that
demonstrates these nonnative bees
forage on Hylaeus host plants
(Magnacca 2007, p. 188). It has also
been suggested parasitoid wasps may
compete for nectar with native Hylaeus
species (Daly and Magnacca 2003, p.
10); however, information
demonstrating nonnative parasitoid
wasps forage on the same host plants as
the seven Hawaiian yellow-faced bees is
unavailable.
We acknowledge the potential for
negative impacts on Hylaeus
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anthracinus, H. assimulans, H. facilis,
H. hilaris, H. kuakea, H. longiceps, and
H. mana from competition with the
European honey bee for nectar and
pollen (Magnacca 2007, p. 188). In
addition, one study in Hawaii suggests
the European honey bee may have an
additional advantage for collecting
pollen and nectar because it may not be
negatively affected by the presence of
predatory big-headed ants on native
vegetation (Lach 2008, p. 155).
Competition with the European honey
bee may be a potential threat to the
seven Hylaeus species, because (1)
Honey bees forage on Hylaeus host plant
species; (2) they may exclude Hylaeus
spp. from those resources (Hylaeus spp.
are never found foraging in the presence
of European honey bees); and (3) honey
bees may have a competitive advantage
over Hawaiian Hylaeus ssp., as one
study suggests honey bees are not
negatively affected by the presence of
big-headed ants on native vegetation to
the extent the Hylaeus species may be.
Honey bees have been known to exclude
other Hylaeus species, and it is welldocumented that they forage in native
plant areas. However, the best available
scientific information indicates that
competition with the European honey
bee may represent a threat to these
seven Hylaeus species, but the threat is
of unknown magnitude, and additional
research would be helpful to better
understand this interaction.
We have no information indicating
other species of nonnative bees or
parasitoid wasps negatively impact
populations of the seven species of
Hylaeus bees due to competition for
nectar and pollen. Therefore, we have
determined that competition with other
species of nonnative bees or parasitoid
wasps is not a threat.
Summary of Factor E
The small number of populations of
Hylaeus facilis, H. hilaris, H. kuakea,
and H. mana increase their risk of
extinction due to stochastic events such
as hurricanes, wildfires, and drought,
which, although unpredictable,
represent an ongoing and significant
threat to H. facilis, H. hilaris, H. kuakea,
and H. mana. We have no information
indicating other nonnative bees or
parasitoid wasps compete for nectar and
pollen on Hylaeus host plants.
Therefore, we have determined that
competition with these species does not
present a significant threat to the seven
Hylaeus species. Honey bees forage in
native plant areas and have been known
to exclude other Hylaeus species.
However, the best available information
does not indicate competition between
honey bees and the seven Hylaeus
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species addressed in this finding is a
significantly quantifiable threat.
Finding
As required by the Act, we conducted
a review of the status of the species and
considered the five factors in assessing
whether Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana are
endangered or threatened throughout
their ranges. We examined the best
scientific and commercial information
available regarding the past, present,
and future threats faced by these seven
Hylaeus species. We reviewed the
petitions, information available in our
files, information submitted to us
following publication of our 90-day
petition finding (75 FR 34077; June 16,
2010), and other available published
and unpublished information, and we
consulted with Hylaeus bee experts and
other Federal and State resource
agencies. In considering what factors
might constitute a threat, we must look
beyond the mere exposure of the species
to the factor to determine whether the
species responds to the factor in a way
that causes actual impacts to the
species. If there is exposure to a factor,
but no response, or only a positive
response, that factor is not a threat. If
there is exposure and the species
responds negatively, the factor may be
a threat and we then attempt to
determine how significant a threat it is.
If the threat is significant, it may drive
or contribute to the risk of extinction of
the species such that the species
warrants listing as endangered or
threatened as those terms are defined by
the Act. This does not necessarily
require empirical proof of a threat. The
combination of exposure and some
corroborating evidence of how the
species is likely impacted could suffice.
However, the mere identification of
factors that could impact a species
negatively is not sufficient to compel a
finding that listing is appropriate; we
require evidence that these factors are
operative threats that act on the species
to the point the species meets the
definition of endangered or threatened
under the Act.
In this review of the status of the
seven Hylaeus species, we identified a
number of threats under the five-factor
analysis including: destruction or
modification of coastal and lowland
habitats from urbanization and land
conversion, nonnative plants, nonnative
ungulates, and wildfire (Factor A);
predation by nonnative ants and the
western yellow jacket wasp (Factor C);
inadequate protection from threats by
existing regulatory mechanisms (Factor
D); and other natural or manmade
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factors, such as small population size
(Factor E).
Under Factor A (‘‘Present or
Threatened Destruction, Modification,
or Curtailment of the Habitat or
Range’’), we evaluated the effects of: (1)
Urbanization and land use conversion;
(2) nonnative plant species; (3)
nonnative ungulates; (4) fire; (5)
recreational activities; (6) stochastic
events, such as hurricanes and droughts;
and (7) climate change.
Hylaeus anthracinus, H. assimilans,
H. facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana are known from
native coastal, lowland dry, and
lowland mesic habitats. These habitats
have been severely altered and degraded
over the past 200 years due to land
management practices such as
agriculture and urban development, and
from the impacts of nonnative species,
fire, recreational activities, and
stochastic events (e.g., hurricanes and
drought). The loss of native coastal and
lowland dry habitats in the main
Hawaiian Islands is estimated to be
more than 75 percent and 90 percent,
respectively (Bruegmann 1996, p. 26;
Juvik and Juvik 1998, p. 124; Xerces
Society 2009, p. 23). Additionally,
native coastal and lowland habitats
continue to become increasingly
fragmented due to a variety of factors,
thereby reducing the ability of the seven
Hylaeus species to locate host plants to
forage for nectar and pollen and to
locate suitable nesting sites. In
particular, coastal and lowland dry
habitats remain popular for land use
and development. During surveys
conducted between 1998 and 2007, the
five Hylaeus species collected by
Perkins over 100 years ago (Hylaeus
anthracinus, H. assimilans, H. facilis, H.
hilaris, and H. longiceps), were largely
absent from almost all of their
historically known locations. Hylaeus
kuakea and H. mana were discovered
relatively recently, and we lack
information that would conclusively
establish their historical range. Based on
our assessment of the best available
information, we believe degradation and
destruction of native coastal and
lowland habitats due to past and present
land management practices, such as
agriculture and urban development,
pose a significant threat to the seven
Hylaeus species throughout their ranges
now and will likely continue for the
foreseeable future.
The spread of nonnative plants and
the conversion of coastal and lowland
native habitat to nonnative habitat are
believed to be primary causes of the
decline of, and current threats to, the
known populations of each of the seven
Hylaeus species. The seven Hylaeus
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species depend on native plants for
nectar and pollen and are almost
entirely absent from habitat dominated
by nonnative plants. Many of the native
plants used as foraging resources by the
adults of the seven Hylaeus species are
declining due to competition with
nonnative plants and a lack of native
pollinators that actually pollinate while
collecting nectar. For example, H.
anthracinus and H. longiceps forage on
three federally endangered plants
(Chamaesyce celastroides var. kaenana,
Hedyotis coriacea, and Sesbania
tomentosa). To compound our concerns,
inadequate regulatory control (see
Factor D. The Inadequacy of Existing
Regulatory Mechanisms) has led to and
continues to contribute to an ever
increasing number of nonnative plant
species introductions to the Hawaiian
Islands. Once established, nonnative
plant species are quickly spread by
intrastate commerce, birds, people, feral
ungulates, and on their own, and result
in the rapid alteration and degradation
of the native plant communities upon
which these seven Hawaiian yellowfaced bees depend. Therefore, based on
our assessment of the best available
information, we believe degradation and
destruction of native coastal and
lowland habitat due to nonnative plants
poses a significant threat to the seven
Hylaeus species throughout their ranges
now and will likely continue for the
foreseeable future.
Nonnative ungulates (e.g., pigs, goats,
axis deer, and cattle) are one of the
primary causes of the alteration and
degradation of native vegetation and
habitat in the Hawaiian Islands. Because
feral ungulate populations are managed
by the State for the enhancement of
State Game Management Units and
because there is no regulatory
mechanism for their control or
elimination (see Factor A. Habitat
Destruction and Modification by
Nonnative Ungulates), it is expected
that this threat will continue to impact
the habitat of the seven yellow-faced
bees addressed in this finding. Habitat
degradation and destruction, due to
their direct effects of trampling and
consuming native plants and indirect
effects of rooting, erosion, and spreading
seeds and fruits of nonnative plants,
pose a significant threat to the seven
Hylaeus species throughout their ranges
now and will likely continue for the
foreseeable future.
Fire is a human-exacerbated threat to
native species and natural vegetation in
Hawaii. Fire can kill most native trees
and shrubs, and in a burned area native
plants are usually replaced by nonnative
plants adapted to survive and regenerate
after fire. The seven Hylaeus bees
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primarily occur in coastal, lowland dry,
and lowland mesic habitat areas that are
particularly prone to the impacts of fire.
Repeated fires in these areas often result
in the conversion of native-dominated
vegetation to nonnative-dominated
vegetation. Fires enable fire-adapted,
nonnative plants to gain a competitive
edge over native plants, resulting in the
replacement of native plants used for
foraging by Hylaeus bees with nonnative
plants that are not used by the bees for
foraging. Although there are
management plans in place to address
the threat of fire in many areas of the
State, fires continue to occur annually
across the State and threaten the future
existence of known yellow-faced bee
habitat and population sites (see Factor
A. Habitat Destruction and Modification
by Fire). For these reasons, we conclude
fire remains a significant threat to the
seven Hylaeus species throughout their
ranges in coastal, lowland dry, and
lowland mesic habitats, and will likely
continue for the foreseeable future.
While trampling and compaction of
vegetation from human activities may
negatively impact the habitat of some
populations of the seven Hylaeus bees,
we conclude recreational activities are
not a threat to Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana
throughout their ranges.
We are concerned about the effects of
projected climate change, particularly
rising temperatures and their impact on
Hylaeus spp. host plants; however, we
recognize there is limited information
on the exact nature of impacts from
climate change. Because the specific
and cumulative effects of climate
change on the seven Hylaeus bees are
presently unknown, any conclusion
regarding the immediacy and
significance of the threat from climate
change would be speculative. However,
the effects of climate change are
expected to exacerbate and compound
the many ongoing threats facing these
species and their habitat (e.g., frequency
of wildfire, reduced precipitation, etc.).
Based on our evaluation of Factor A,
using the best available scientific and
commercial information as summarized
above, we conclude the present or
threatened destruction, modification, or
curtailment of the habitat or range of
Hylaeus anthracinus, H. assimilans, H.
facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana presents a
significant threat to these seven Hylaeus
species across their ranges.
Under Factor B (‘‘Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes’’), we determined
six of the seven Hylaeus species are not
threatened by over-collection. We
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examined whether H. hilaris was
potentially vulnerable to over-collection
because it is inherently rare, known
from only one location, and has a
cleptoparasitic life history. However,
because this species is easily
recognizable, we see little reason for
scientists to retain specimens observed
in the field during future collections. In
addition, because it occurs in habitat
that is protected and managed by TNC,
we find overutilization is not a threat to
H. hilaris throughout its range.
Furthermore, recreational or insect
enthusiast collection of the seven
Hylaeus bees does not appear to be a
threat to any of these species.
Under Factor C (‘‘Disease or
Predation’’), we found no evidence that
disease is currently impacting the seven
Hawaiian yellow-faced bees, or that
disease outbreaks will increase in the
future. Ants are found in habitats
throughout the Hawaiian Islands, are
known to prey upon Hylaeus bees, and
are reported to have eliminated Hylaeus
species from specific areas where their
ranges overlap. Because ants are easily
able to widely disperse and are efficient
predators, and because Hylaeus species
are not adapted to avoid ant predation,
we believe this threat will continue to
threaten all populations of all seven
yellow-faced bees. Therefore, we
conclude predation by ants is an
ongoing and significant threat to the
seven Hylaeus bees across their entire
ranges, and this threat is likely to
continue into the future.
Yellow jacket wasps are aggressive,
generalist predators found in the same
types of habitats as these seven Hylaeus
species, and have been documented
preying upon other Hawaiian Hylaeus
bees. Therefore, we conclude yellow
jacket wasp predation is a significant
threat to the seven Hylaeus bees across
their entire ranges and particularly to
those species known from two or fewer
population sites. The best available
information does not suggest predation
by native and nonnative parasitoid
wasps is a significant threat to the seven
Hylaeus bees.
Under Factor D (‘‘Inadequacy of
Existing Regulatory Mechanisms’’), we
consider the threat from inadequate
regulatory mechanisms to be immediate
and significant. The State of Hawaii’s
current management of nonnative game
mammals does not adequately address
the primary threats to Hylaeus
anthracinus, H. assimulans, H. facilis,
H. hilaris, H. kuakea, H. longiceps, and
H. mana or their habitat (Factor A).
Existing State and Federal regulatory
mechanisms are not adequately
preventing the introduction and spread
of nonnative animal and habitat-altering
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plant species between islands and
watersheds (Factor A), and predation by
nonnative animal species (Factor C)
poses a major ongoing threat to the
seven Hylaeus species. In addition,
existing regulatory mechanisms are
inadequate to prevent the introduction
and spread of nonnative insect
predators, or competitors that directly
compete with one or more of the seven
bee species for food, space, and other
necessary resources (see Factors C and
E). Based on our evaluation of Factor D,
we conclude that the seven Hylaeus bee
species are threatened by inadequate
existing regulatory mechanisms across
their ranges.
Under Factor E (‘‘Other Natural or
Manmade Factors Affecting the Species’
Continued Existence’’), we determined
that small population size is a
significant threat to Hylaeus facilis, H.
hilaris, H. kuakea, and H. mana. These
species are each only known from one
or two populations, and the risk of
extinction from stochastic events (e.g.,
hurricanes, wildfires, and drought) is
high. We have also determined that
competition with the European honey
bee is a potentially significant threat to
all seven species. While we lack
information indicating Hawaiian
Hylaeus populations have declined
because of competition with the
European honey bee for nectar and
pollen, the native Hylaeus and the
European honey bee are competing for
the same pollen and nectar resources.
However, we have no information
indicating that competition is at a level
that represents a threat to the seven
Hylaeus species addressed in this
finding.
We found that competition for nectar
and pollen with other species of
nonnative bees or parasitoid wasps is
not a threat to the seven Hylaeus bees
at this time. Based on our evaluation
under Factor E as summarized above,
we conclude Hylaeus facilis, H. hilaris,
H. kuakea, and H. mana are threatened
because of their small population size
across their ranges.
On the basis of the best scientific and
commercial information available, we
find that the petitioned action, listing
the seven species of Hawaiian yellowfaced bees (Hylaeus anthracinus, H.
assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana) as
endangered or threatened is warranted.
We will make a determination on the
status of these species as endangered or
threatened when we prepare a proposed
listing determination. However, as
explained in more detail below, an
immediate proposal of a regulation
implementing this action is precluded
by higher priority listing actions, and
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progress is being made to add or remove
qualified species from the Lists of
Endangered and Threatened Wildlife
and Plants.
We reviewed the available
information to determine if the existing
and foreseeable threats render any of the
seven Hawaiian yellow-faced bee
species at risk of extinction now such
that issuing an emergency regulation
temporarily listing the species under
section 4(b)(7) of the Act is warranted.
We determined that issuing an
emergency regulation temporarily
listing these species is not warranted at
this time for the following reasons.
Although populations are small, five of
the seven species occur in several
discrete localities, and we do not
believe there are any potential threats of
such great immediacy, severity, or scope
that would simultaneously threaten all
of the known populations of these five
species with the imminent risk of
extinction. Although Hylaeus hilaris is
known from one population on the
northwest coast within TNC’s Moomomi
Preserve on Molokai, and H. mana is
known from one population along the
Manana Trail in the Koolau Mountains
on Oahu, within the State’s Ewa Forest
Reserve, we are unaware of any
potential threats in either of these areas
that would threaten these populations
with the imminent risk of extinction.
However, if at any time we determine
that issuing an emergency regulation
temporarily listing any of these seven
species of Hawaiian yellow-faced bees is
warranted, we will initiate this action at
that time.
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Listing Priority Number
The Service adopted guidelines on
September 21, 1983 (48 FR 43098), to
establish a rational system for utilizing
available resources for the highest
priority species when adding species to
the Lists of Endangered or Threatened
Wildlife and Plants or reclassifying
species listed as threatened to
endangered status. These guidelines,
titled ‘‘Endangered and Threatened
Species Listing and Recovery Priority
Guidelines,’’ address the immediacy
and magnitude of threats, and the level
of taxonomic distinctiveness by
assigning priority in descending order to
monotypic genera (genus with one
species), full species, and subspecies (or
equivalently, distinct population
segments of vertebrates). We assigned
the seven species of Hawaiian yellowfaced bees a Listing Priority Number
(LPN) of 2, based on our finding that the
seven species face threats that are of
high magnitude and are imminent. This
is the highest priority that can be
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provided to a species under our
guidance.
Threats to the seven species of
Hawaiian yellow-faced bees include the
present or threatened destruction,
modification, or curtailment of their
habitat, predation, the inadequacy of
existing regulatory mechanisms, and
other natural or manmade factors. One
or more of the threats are occurring in
each of the seven species’ known
populations in the Hawaiian Islands.
These threats are ongoing and, in some
cases (such as nonnative species), are
considered irreversible. Our rationale
for assigning each of the seven species
of Hawaiian yellow-faced bees an LPN
2 is outlined below.
Under the Service’s LPN Guidance,
the magnitude of threat is the first
criterion we look at when establishing a
listing priority. The guidance indicates
that species with the highest magnitude
of threat are those species facing the
greatest threats to their continued
existence. These species receive the
highest listing priority. The threats
facing the seven species of Hawaiian
yellow-faced bees are high in magnitude
because the major threats (destruction or
modification of their habitat, predation,
inadequate protection from threats by
existing regulatory mechanisms, and
other natural or manmade factors) occur
throughout all of the ranges of each of
the seven species.
Based on an evaluation of the effects
of urbanization and land use
conversion, nonnative plants and
ungulates, fire, and stochastic events on
the coastal and lowland habitat of each
of the seven Hylaeus bees, we
determined these effects occur
throughout the range of each species
and will continue to occur into the
future. While habitat degradation and
destruction continues to reduce the
amount of potentially suitable habitat
available for foraging, predation by
nonnative ants and likely predation by
yellow jacket wasps are a significant
threat to the seven species throughout
their ranges, and, lacking any viable
means of their control, will continue to
occur into the future. Regulations are
not in place at the local, State, or
Federal level to adequately minimize
the threat of habitat degradation and
destruction from nonnative plants and
ungulates. In addition, existing
regulatory mechanisms are inadequate
to prevent the introduction and spread
of nonnative insect predators or
competitors. We determined these
threats occur throughout the range of
each of the seven species of Hylaeus
bees and will continue to occur into the
future unless restriction on the
introduction and the control of,
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nonnative plants and animals, are put in
place. We believe the ability of the
populations of the seven Hylaeus bees
to stabilize or increase over the long
term is highly diminished given the
widespread landscape-level changes
and the threats from predation and
competition that are occurring. Thus,
we believe the available information
indicates the magnitude of threats is
high.
Under our LPN Guidance, the second
criterion we consider in assigning a
listing priority is the immediacy of
threats. This criterion is intended to
ensure species that face actual,
identifiable threats are given priority
over those for which threats are only
potential or that are intrinsically
vulnerable but are not known to be
presently facing such threats. The
threats to the seven Hawaiian yellowfaced bees are imminent because we
have factual information that the threats
are identifiable, and that all of the seven
species are currently facing these threats
throughout all portions of their ranges.
The identifiable threats are covered in
detail under the discussion of Factors A
and E of this finding and include
destruction or modification of their
habitat, predation, inadequate existing
regulatory mechanisms, and other
natural or manmade factors such as
small population size. In addition to
their current existence, we expect these
threats to continue and likely intensify
into the foreseeable future.
The third criterion in our LPN
guidance is intended to devote
resources to those species representing
highly distinctive or isolated gene pools
as reflected by taxonomy. The seven
Hawaiian yellow-faced bees are valid
taxa at the species level, and therefore
receive a higher priority than subspecies
or distinct population segments, but a
lower priority than species in a
monotypic genus.
The seven Hawaiian yellow-faced
bees face high magnitude, imminent
threats, and are valid taxa at the species
level. Thus, in accordance with our LPN
guidance, we have assigned each of the
seven Hawaiian yellow-faced bees an
LPN of 2. We will continue to monitor
the threats to the seven Hylaeus bees
and the species’ status on an annual
basis; should the magnitude or the
imminence of the threats change, we
will revisit our assessment of the LPN.
Work on a proposed listing
determination for Hylaeus anthracinus,
H. assimulans, H. facilis, H. hilaris, H.
kuakea, H. longiceps, and H. mana is
precluded by work on higher priority
listing actions with absolute statutory,
court-ordered, or court-approved
deadlines and final listing
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determinations for those species that
were proposed for listing with funds
from Fiscal Year 2011. This work
includes all the actions listed in the
tables below under expeditious
progress.
Preclusion and Expeditious Progress
Preclusion is a function of the listing
priority of a species in relation to the
resources that are available and the cost
and relative priority of competing
demands for those resources. Thus, in
any given fiscal year (FY), multiple
factors dictate whether it will be
possible to undertake work on a listing
proposal regulation or whether
promulgation of such a proposal is
precluded by higher priority listing
actions.
The resources available for listing
actions are determined through the
annual Congressional appropriations
process. The appropriation for the
Listing Program is available to support
work involving the following listing
actions: Proposed and final listing rules;
90-day and 12-month findings on
petitions to add species to the Lists of
Endangered and Threatened Wildlife
and Plants (Lists) or to change the status
of a species from threatened to
endangered; annual ‘‘resubmitted’’
petition findings on prior warrantedbut-precluded petition findings as
required under section 4(b)(3)(C)(i) of
the Act; critical habitat petition
findings; proposed and final rules
designating critical habitat; and
litigation-related, administrative, and
program-management functions
(including preparing and allocating
budgets, responding to Congressional
and public inquiries, and conducting
public outreach regarding listing and
critical habitat). The work involved in
preparing various listing documents can
be extensive and may include, but is not
limited to: Gathering and assessing the
best scientific and commercial data
available and conducting analyses used
as the basis for our decisions; writing
and publishing documents; and
obtaining, reviewing, and evaluating
public comments and peer review
comments on proposed rules and
incorporating relevant information into
final rules. The number of listing
actions that we can undertake in a given
year also is influenced by the
complexity of those listing actions; that
is, more complex actions generally are
more costly. The median cost for
preparing and publishing a 90-day
finding is $39,276; for a 12-month
finding, $100,690; for a proposed rule
with critical habitat, $345,000; and for
a final listing rule with critical habitat,
$305,000.
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We cannot spend more than is
appropriated for the Listing Program
without violating the Anti-Deficiency
Act (see 31 U.S.C. 1341(a)(1)(A)). In
addition, in FY 1998 and for each fiscal
year since then, Congress has placed a
statutory cap on funds that may be
expended for the Listing Program, equal
to the amount expressly appropriated
for that purpose in that fiscal year. This
cap was designed to prevent funds
appropriated for other functions under
the Act (for example, recovery funds for
removing species from the Lists), or for
other Service programs, from being used
for Listing Program actions (see House
Report 105–163, 105th Congress, 1st
Session, July 1, 1997).
Since FY 2002, the Service’s budget
has included a critical habitat subcap to
ensure that some funds are available for
other work in the Listing Program (‘‘The
critical habitat designation subcap will
ensure that some funding is available to
address other listing activities’’ (House
Report No. 107–103, 107th Congress, 1st
Session, June 19, 2001)). In FY 2002 and
each year until FY 2006, the Service has
had to use virtually the entire critical
habitat subcap to address courtmandated designations of critical
habitat, and consequently none of the
critical habitat subcap funds have been
available for other listing activities. In
some FYs since 2006, we have been able
to use some of the critical habitat
subcap funds to fund proposed listing
determinations for high-priority
candidate species. In other FYs, while
we were unable to use any of the critical
habitat subcap funds to fund proposed
listing determinations, we did use some
of this money to fund the critical habitat
portion of some proposed listing
determinations so that the proposed
listing determination and proposed
critical habitat designation could be
combined into one rule, thereby being
more efficient in our work. At this time,
for FY 2011, we plan to use some of the
critical habitat subcap funds to fund
proposed listing determinations.
We make our determinations of
preclusion on a nationwide basis to
ensure that the species most in need of
listing will be addressed first and also
because we allocate our listing budget
on a nationwide basis. Through the
listing cap, the critical habitat subcap,
and the amount of funds needed to
address court-mandated critical habitat
designations, Congress and the courts
have in effect determined the amount of
money available for other listing
activities nationwide. Therefore, the
funds in the listing cap, other than those
needed to address court-mandated
critical habitat for already listed species,
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set the limits on our determinations of
preclusion and expeditious progress.
Congress identified the availability of
resources as the only basis for deferring
the initiation of a rulemaking that is
warranted. The Conference Report
accompanying Pub. L. 97–304
(Endangered Species Act Amendments
of 1982), which established the current
statutory deadlines and the warrantedbut-precluded finding, states that the
amendments were ‘‘not intended to
allow the Secretary to delay
commencing the rulemaking process for
any reason other than that the existence
of pending or imminent proposals to list
species subject to a greater degree of
threat would make allocation of
resources to such a petition [that is, for
a lower-ranking species] unwise.’’
Although that statement appeared to
refer specifically to the ‘‘to the
maximum extent practicable’’ limitation
on the 90-day deadline for making a
‘‘substantial information’’ finding, that
finding is made at the point when the
Service is deciding whether or not to
commence a status review that will
determine the degree of threats facing
the species, and therefore the analysis
underlying the statement is more
relevant to the use of the warranted-butprecluded finding, which is made when
the Service has already determined the
degree of threats facing the species and
is deciding whether or not to commence
a rulemaking.
In FY 2011, on April 15, 2011,
Congress passed the Full-Year
Continuing Appropriations Act (Pub. L.
112–10), which provides funding
through September 30, 2011. The
Service has $20,902,000 for the listing
program. Of that, $9,472,000 is being
used for determinations of critical
habitat for already listed species. Also
$500,000 is appropriated for foreign
species listings under the Act. The
Service thus has $10,930,000 available
to fund work in the following categories:
Compliance with court orders and
court-approved settlement agreements
requiring that petition findings or listing
determinations be completed by a
specific date; section 4 (of the Act)
listing actions with absolute statutory
deadlines; essential litigation-related,
administrative, and listing programmanagement functions; and highpriority listing actions for some of our
candidate species. In FY 2010, the
Service received many new petitions
and a single petition to list 404 species.
The receipt of petitions for a large
number of species is consuming the
Service’s listing funding that is not
dedicated to meeting court-ordered
commitments. Absent some ability to
balance effort among listing duties
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under existing funding levels, the
Service is only able to initiate a few new
listing determinations for candidate
species in FY 2011.
In 2009, the responsibility for listing
foreign species under the Act was
transferred from the Division of
Scientific Authority, International
Affairs Program, to the Endangered
Species Program. Therefore, starting in
FY 2010, we used a portion of our
funding to work on the actions
described above for listing actions
related to foreign species. In FY 2011,
we anticipate using $1,500,000 for work
on listing actions for foreign species,
which reduces funding available for
domestic listing actions; however,
currently only $500,000 has been
allocated for this function. Although
there are no foreign species issues
included in our high-priority listing
actions at this time, many actions have
statutory or court-approved settlement
deadlines, thus increasing their priority.
The budget allocations for each specific
listing action are identified in the
Service’s FY 2011 Allocation Table (part
of our record).
We assigned each of the seven species
of Hawaiian yellow-faced bees an LPN
of 2, based on our finding that each
species faces immediate and high
magnitude threats from the present or
threatened destruction, modification, or
curtailment of its habitat, the threat of
predation from and competition with
nonnative species, and from the
inadequacy of existing regulatory
mechanisms. In addition, H. facilis, H.
hilaris, H. kuakea, and H. mana are
each significantly threatened by small
population size. Under our 1983
Guidelines, a ‘‘species’’ facing imminent
high-magnitude threats is assigned an
LPN of 1, 2, or 3 depending on its
taxonomic status. Because H.
anthracinus, H. assimulans, H. facilis,
H. hilaris, H. kuakea, H. longiceps, and
H. mana are species, we assigned each
an LPN of 2 (the highest category
available for a species). For the above
reasons, funding a proposed listing
determination for the seven species of
Hawaiian yellow-faced bees is
precluded by court-ordered and court-
approved settlement agreements, listing
actions with absolute statutory
deadlines, and work on proposed listing
determinations for those candidate
species with a higher listing priority.
Based on our September 21, 1983,
guidelines for assigning an LPN for each
candidate species (48 FR 43098), we
have a significant number of species
with a LPN of 2. Using these guidelines,
we assign each candidate an LPN of 1
to 12, depending on the magnitude of
threats (high or moderate to low),
immediacy of threats (imminent or
nonimminent), and taxonomic status of
the species (in order of priority:
monotypic genus (a species that is the
sole member of a genus); species; or part
of a species (subspecies, or distinct
population segment)). The lower the
listing priority number, the higher the
listing priority (that is, a species with an
LPN of 1 would have the highest listing
priority).
Because of the large number of highpriority species, we have further ranked
the candidate species with an LPN of 2
by using the following extinction-risk
type criteria: International Union for the
Conservation of Nature and Natural
Resources (IUCN) Red list status/rank,
Heritage rank (provided by
NatureServe), Heritage threat rank
(provided by NatureServe), and species
currently with fewer than 50
individuals, or 4 or fewer populations.
Those species with the highest IUCN
rank (critically endangered), the highest
Heritage rank (G1), the highest Heritage
threat rank (substantial, imminent
threats), and currently with fewer than
50 individuals, or fewer than 4
populations, originally comprised a
group of approximately 40 candidate
species (‘‘Top 40’’). These 40 candidate
species have had the highest priority to
receive funding to work on a proposed
listing determination. As we work on
proposed and final listing rules for those
40 candidates, we apply the ranking
criteria to the next group of candidates
with an LPN of 2 and 3 to determine the
next set of highest priority candidate
species. Finally, proposed rules for
reclassification of threatened species to
endangered species are lower priority,
because as listed species, they are
already afforded the protections of the
Act and implementing regulations.
However, for efficiency reasons, we may
choose to work on a proposed rule to
reclassify a species to endangered if we
can combine this with work that is
subject to a court-determined deadline.
With our workload so much bigger
than the amount of funds we have to
accomplish it, it is important that we be
as efficient as possible in our listing
process. Therefore, as we work on
proposed rules for the highest priority
species in the next several years, we are
preparing multi-species proposals when
appropriate, and these may include
species with lower priority if they
overlap geographically or have the same
threats as a species with an LPN of 2.
In addition, we take into consideration
the availability of staff resources when
we determine which high-priority
species will receive funding to
minimize the amount of time and
resources required to complete each
listing action.
As explained above, a determination
that listing is warranted but precluded
must also demonstrate that expeditious
progress is being made to add and
remove qualified species to and from
the Lists of Endangered and Threatened
Wildlife and Plants. As with our
‘‘precluded’’ finding, the evaluation of
whether progress in adding qualified
species to the Lists has been expeditious
is a function of the resources available
for listing and the competing demands
for those funds. (Although we do not
discuss it in detail here, we are also
making expeditious progress in
removing species from the list under the
Recovery program in light of the
resource available for delisting, which is
funded by a separate line item in the
budget of the Endangered Species
Program. So far during FY 2011, we
have completed delisting rules for three
species.) Given the limited resources
available for listing, we find that we are
making expeditious progress in FY 2011
in the Listing Program. This progress
included preparing and publishing the
following determinations:
FY 2011 COMPLETED LISTING ACTIONS
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Publication date
Title
Actions
10/6/2010 ..........
Endangered Status for the Altamaha Spinymussel and Designation of Critical Habitat.
12-month Finding on a Petition To list the Sacramento Splittail as
Endangered or Threatened.
Endangered Status and Designation of Critical Habitat for
Spikedace and Loach Minnow.
90-Day Finding on a Petition To List the Bay Springs Salamander
as Endangered.
Proposed Listing .........................
Endangered ................................
Notice of 12-month petition finding, Not warranted.
Proposed Listing Endangered
(uplisting).
Notice of 90-day Petition Finding, Not substantial.
10/7/2010 ..........
10/28/2010 ........
11/2/2010 ..........
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FR pages
75 FR 61664–61690
75 FR 62070–62095
75 FR 66481–66552
75 FR 67341–67343
Federal Register / Vol. 76, No. 172 / Tuesday, September 6, 2011 / Proposed Rules
55199
FY 2011 COMPLETED LISTING ACTIONS—Continued
Publication date
Title
Actions
11/2/2010 ..........
Determination of Endangered Status for the Georgia Pigtoe Mussel, Interrupted Rocksnail, and Rough Hornsnail and Designation of Critical Habitat.
Listing the Rayed Bean and Snuffbox as Endangered ..................
12-Month Finding on a Petition To List Cirsium wrightii (Wright’s
Marsh Thistle) as Endangered or Threatened.
Endangered Status for Dunes Sagebrush Lizard ...........................
12-Month Finding on a Petition To List the North American Wolverine as Endangered or Threatened.
12-Month Finding on a Petition To List the Sonoran Population of
the Desert Tortoise as Endangered or Threatened.
12-Month Finding on a Petition To List Astragalus microcymbus
and Astragalus schmolliae as Endangered or Threatened.
Listing Seven Brazilian Bird Species as Endangered Throughout
Their Range.
90-Day Finding on a Petition To List the Red Knot subspecies
Calidris canutus roselaari as Endangered.
Endangered Status for the Sheepnose and Spectaclecase Mussels.
12-Month Finding on a Petition To List the Pacific Walrus as Endangered or Threatened.
90-Day Finding on a Petition To List the Sand Verbena Moth as
Endangered or Threatened.
Determination of Threatened Status for the New Zealand-Australia Distinct Population Segment of the Southern Rockhopper
Penguin.
12-Month Finding on a Petition To List Solanum conocarpum
(marron bacora) as Endangered.
12-Month Finding on a Petition To List Thorne’s Hairstreak Butterfly as Endangered.
12-Month Finding on a Petition To List Astragalus hamiltonii,
Penstemon flowersii, Eriogonum soredium, Lepidium ostleri,
and Trifolium friscanum as Endangered or Threatened.
90-Day Finding on a Petition To List the Wild Plains Bison or
Each of Four Distinct Population Segments as Threatened.
90-Day Finding on a Petition To List the Unsilvered Fritillary Butterfly as Threatened or Endangered.
12-Month Finding on a Petition To List the Mt. Charleston Blue
Butterfly as Endangered or Threatened.
90-Day Finding on a Petition To List the Texas Kangaroo Rat as
Endangered or Threatened.
Initiation of Status Review for Longfin Smelt ..................................
Withdrawal of Proposed Rule To List the Flat-Tailed Horned Lizard as Threatened.
Proposed Threatened Status for the Chiricahua Leopard Frog
and Proposed Designation of Critical Habitat.
Final Listing Endangered ...........
75 FR 67511–67550
Proposed Listing Endangered ....
Notice of 12-month petition finding, Warranted but precluded.
Proposed Listing Endangered ....
Notice of 12-month petition finding, Warranted but precluded.
Notice of 12-Month petition finding, Warranted but precluded.
Notice of 12-month petition finding, Warranted but precluded.
Final Listing Endangered ...........
75 FR 67551–67583
75 FR 67925–67944
11/2/2010 ..........
11/4/2010 ..........
12/14/2010 ........
12/14/2010 ........
12/14/2010 ........
12/15/2010 ........
12/28/2010 ........
1/4/2011 ............
1/19/2011 ..........
2/10/2011 ..........
2/17/2011 ..........
2/22/2011 ..........
2/22/2011 ..........
2/23/2011 ..........
2/23/2011 ..........
2/24/2011 ..........
2/24/2011 ..........
3/8/2011 ............
3/8/2011 ............
3/10/2011 ..........
3/15/2011 ..........
3/15/2011 ..........
3/22/2011 ..........
4/1/2011 ............
4/5/2011 ............
4/5/2011 ............
4/12/2011 ..........
4/13/2011 ..........
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4/14/2011 ..........
4/14/2011 ..........
4/26/2011 ..........
4/26/2011 ..........
5/12/2011 ..........
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12-Month Finding on a Petition To List the Berry Cave Salamander as Endangered.
90-Day Finding on a Petition To List the Spring Pygmy Sunfish
as Endangered.
12-Month Finding on a Petition To List the Bearmouth
Mountainsnail, Byrne Resort Mountainsnail, and Meltwater
Lednian Stonefly as Endangered or Threatened.
90-Day Finding on a Petition To List the Peary Caribou and Dolphin and Union Population of the Barren-Ground Caribou as
Endangered or Threatened.
Proposed Endangered Status for the Three Forks Springsnail and
San Bernardino Springsnail, and Proposed Designation of Critical Habitat.
90-Day Finding on a Petition To List Spring Mountains Acastus
Checkerspot Butterfly as Endangered.
90-Day Finding on a Petition To List the Prairie Chub as Threatened or Endangered.
12-Month Finding on a Petition To List Hermes Copper Butterfly
as Endangered or Threatened.
90-Day Finding on a Petition To List the Arapahoe Snowfly as
Endangered or Threatened.
90-Day Finding on a Petition To List the Smooth-Billed Ani as
Threatened or Endangered.
Withdrawal of the Proposed Rule To List the Mountain Plover as
Threatened.
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Notice of 90-day Petition Finding, Not substantial.
Proposed Listing Endangered ....
Notice of 12-month petition finding, Warranted but precluded.
Notice of 90-day Petition Finding, Substantial.
Final Listing Threatened .............
Notice of 12-month petition finding, Warranted but precluded.
Notice of 12-month petition finding, Not warranted.
Notice of 12-month petition finding, Warranted but precluded
& Not Warranted.
Notice of 90-day Petition Finding, Not substantial.
Notice of 90-day Petition Finding, Not substantial.
Notice of 12-month petition finding, Warranted but precluded.
Notice of 90-day Petition Finding, Substantial.
Notice of Status Review .............
Proposed rule withdrawal ...........
FR pages
75 FR 77801–77817
75 FR 78029–78061
75 FR 78093–78146
75 FR 78513–78556
75 FR 81793–81815
76 FR 304–311
76 FR 3392–3420
76 FR 7634–7679
76 FR 9309–9318
76 FR 9681–9692
76 FR 9722–9733
76 FR 9991–10003
76 FR 10166–10203
76 FR 10299–10310
76 FR 10310–10319
76 FR 12667–12683
76 FR 12683–12690
76 FR 13121–13122
76 FR 14210–14268
Proposed Listing Threatened;
Proposed Designation of Critical Habitat.
Notice of 12-month petition finding, Warranted but precluded.
Notice of 90-day Petition Finding, Substantial.
Notice of 12-month petition finding, Not Warranted and Warranted but precluded.
Notice of 90-day Petition Finding, Substantial.
76 FR 14126–14207
Proposed Listing Endangered;
Proposed Designation of Critical Habitat.
Notice of 90-day Petition Finding, Substantial.
Notice of 90-day Petition Finding, Substantial.
Notice of 12-month petition finding, Warranted but precluded.
Notice of 90-day Petition Finding, Substantial.
Notice of 90-day Petition Finding, Not substantial.
Proposed Rule, Withdrawal ........
76 FR 20464–20488
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76 FR 15919–15932
76 FR 18138–18143
76 FR 18684–18701
76 FR 18701–18706
76 FR 20613–20622
76 FR 20911–20918
76 FR 20918–20939
76 FR 23256–23265
76 FR 23265–23271
76 FR 27756–27799
55200
Federal Register / Vol. 76, No. 172 / Tuesday, September 6, 2011 / Proposed Rules
FY 2011 COMPLETED LISTING ACTIONS—Continued
Publication date
Title
Actions
5/25/2011 ..........
90-Day Finding on a Petition To List the Spot-Tailed Earless Lizard as Endangered or Threatened.
Listing the Salmon-Crested Cockatoo as Threatened Throughout
Its Range With Special Rule.
12-Month Finding on a Petition To List Puerto Rican Harlequin
Butterfly as Endangered.
90-Day Finding on a Petition To Reclassify the Straight-Horned
Markhor (Capra falconeri jerdoni) of Torghar Hills as Threatened.
90-Day Finding on a Petition To List the Golden-Winged Warbler
as Endangered or Threatened.
12-Month Finding on a Petition To List the Striped Newt as
Threatened.
12-Month Finding on a Petition To List Abronia ammophila,
Agrostis rossiae, Astragalus proimanthus, Boechera (Arabis)
pusilla, and Penstemon gibbensii as Threatened or Endangered.
90-Day Finding on a Petition To List the Utah Population of the
Gila Monster as an Endangered or a Threatened Distinct Population Segment.
Revised 90-Day Finding on a Petition To Reclassify the Utah
Prairie Dog From Threatened to Endangered.
12-Month Finding on a Petition To List Castanea pumila var.
ozarkensis as Threatened or Endangered.
90-Day Finding on a Petition To List the Eastern Small-Footed
Bat and the Northern Long-Eared Bat as Threatened or Endangered.
12-Month Finding on a Petition To List a Distinct Population Segment of the Fisher in Its United States Northern Rocky Mountain Range as Endangered or Threatened With Critical Habitat.
90-Day Finding on a Petition To List the Bay Skipper as Threatened or Endangered.
12-Month Finding on a Petition To List Pinus albicaulis as Endangered or Threatened With Critical Habitat.
Petition To List Grand Canyon Cave Pseudoscorpion ...................
Notice of 90-day Petition Finding, Substantial.
Final Listing Threatened .............
5/26/2011 ..........
5/31/2011 ..........
6/2/2011 ............
6/2/2011 ............
6/7/2011 ............
6/9/2011 ............
6/21/2011 ..........
6/21/2011 ..........
6/28/2011 ..........
6/29/2011 ..........
6/30/2011 ..........
7/12/2011 ..........
7/19/2011 ..........
7/19/2011 ..........
7/26/2011 ..........
7/26/2011 ..........
7/27/2011 ..........
7/27/2011 ..........
8/2/2011 ............
8/2/2011 ............
8/2/2011 ............
8/2/2011 ............
8/4/2011 ............
8/9/2011 ............
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8/9/2011 ............
8/9/2011 ............
8/10/2011 ..........
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12-Month Finding on a Petition To List the Giant Palouse Earthworm (Drilolerius americanus) as Threatened or Endangered.
12-month Finding on a Petition To List the Frigid Ambersnail as
Endangered.
Determination of Endangered Status for Ipomopsis polyantha
(Pagosa Skyrocket) and Threatened Status for Penstemon
debilis (Parachute Beardtongue) and Phacelia submutica
(DeBeque Phacelia).
12-Month Finding on a Petition To List the Gopher Tortoise as
Threatened in the Eastern Portion of Its Range.
Proposed Endangered Status for the Chupadera Springsnail
(Pyrgulopsis chupaderae) and Proposed Designation of Critical
Habitat.
90-Day Finding on a Petition To List the Straight Snowfly and
Idaho Snowfly as Endangered.
12-Month Finding on a Petition To List the Redrock Stonefly as
Endangered or Threatened.
Listing 23 Species on Oahu as Endangered and Designating Critical Habitat for 124 Species.
90-Day Finding on a Petition To List Six Sand Dune Beetles as
Endangered or Threatened.
Endangered Status for the Cumberland Darter, Rush Darter,
Yellowcheek Darter, Chucky Madtom, and Laurel Dace.
12-Month Finding on a Petition To List the Nueces River and Plateau Shiners as Threatened or Endangered.
Four Foreign Parrot Species [crimson shining parrot, white
cockatoo, Philippine cockatoo, yellow-crested cockatoo].
Proposed Listing of the Miami Blue Butterfly as Endangered, and
Proposed Listing of the Cassius Blue, Ceraunus Blue, and
Nickerbean Blue Butterflies as Threatened Due to Similarity of
Appearance to the Miami Blue Butterfly.
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76 FR 30082–30087
76 FR 30758–30780
Notice of 12-month petition finding, Warranted but precluded.
Notice of 90-day Petition Finding, Substantial.
76 FR 31282–31294
Notice of 90-day Petition Finding, Substantial.
Notice of 12-month petition finding, Warranted but precluded.
Notice of 12-month petition finding, Not Warranted and Warranted but precluded.
76 FR 31920–31926
Notice of 90-day Petition Finding, Not substantial.
76 FR 36049–36053
Notice of 90-day Petition Finding, Not substantial.
Notice of 12-month petition finding, Not warranted.
Notice of 90-day Petition Finding, Substantial.
76 FR 36053–36068
Notice of 12-month petition finding, Not warranted.
76 FR 38504–38532
Notice of 90-day Petition Finding, Substantial.
Notice of 12-month petition finding, Warranted but precluded.
Notice of 12-month petition finding, Not warranted.
Notice of 12-month petition finding, Not warranted.
Notice of 12-month petition finding, Not warranted.
Final
Listing
Endangered,
Threatened.
76 FR 40868–40871
Notice of 12-month petition finding, Warranted but precluded.
Proposed Listing Endangered ....
76 FR 45130–45162
Notice of 90-day Petition Finding, Not substantial.
Notice of 12-month petition finding, Not warranted.
Proposed Listing Endangered ....
Notice of 90-day Petition Finding, Not substantial and substantial.
Final Listing Endangered ...........
Notice of 12-month petition finding, Not warranted.
Proposed Listing Endangered
and Threatened; Notice of 12month petition finding, Not
warranted.
Proposed Listing Endangered
Similarity of Appearance.
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76 FR 31903–31906
76 FR 32911–32929
76 FR 33924–33965
76 FR 37706–37716
76 FR 38095–38106
76 FR 42631–42654
76 FR 42654–42658
76 FR 44547–44564
76 FR 44566–44569
76 FR 45054–45075
76 FR 46218–46234
76 FR 46238–46251
76 FR 46251–46266
76 FR 46362–46594
76 FR 47123–47133
76 FR 48722–48741
76 FR 48777–48788
76 FR 49202–49236
76 FR 49408–49412
55201
Federal Register / Vol. 76, No. 172 / Tuesday, September 6, 2011 / Proposed Rules
FY 2011 COMPLETED LISTING ACTIONS—Continued
Publication date
Title
Actions
8/10/2011 ..........
90-Day Finding on a Petition To List the Saltmarsh Topminnow
as Threatened or Endangered Under the Endangered Species
Act.
Emergency Listing of the Miami Blue Butterfly as Endangered,
and Emergency Listing of the Cassius Blue, Ceraunus Blue,
and Nickerbean Blue Butterflies as Threatened Due to Similarity of Appearance to the Miami Blue Butterfly.
Notice of 90-day Petition Finding, Substantial.
76 FR 49412–49417
Emergency Listing Endangered
Similarity of Appearance.
76 FR 49542–49567
8/10/2011 ..........
Our expeditious progress also
includes work on listing actions that we
funded in FY 2010 and FY 2011 but
have not yet been completed to date.
These actions are listed below. Actions
in the top section of the table are being
conducted under a deadline set by a
court. Actions in the middle section of
the table are being conducted to meet
statutory timelines, that is, timelines
required under the Act. Actions in the
bottom section of the table are highpriority listing actions. These actions
include work primarily on species with
an LPN of 2, and, as discussed above,
selection of these species is partially
based on available staff resources, and
when appropriate, include species with
FR pages
a lower priority if they overlap
geographically or have the same threats
as the species with the high priority.
Including these species together in the
same proposed rule results in
considerable savings in time and
funding, when compared to preparing
separate proposed rules for each of them
in the future.
ACTIONS FUNDED IN FY 2010 AND FY 2011 BUT NOT YET COMPLETED
Species
Action
Actions Subject to Court Order/Settlement Agreement
4 parrot species (military macaw, yellow-billed parrot, red-crowned parrot, scarlet macaw) 5 ......................................
4 parrot species (blue-headed macaw, great green macaw, grey-cheeked parakeet, hyacinth macaw) 5 ...................
Longfin smelt ..................................................................................................................................................................
12-month petition finding.
12-month petition finding.
12-month petition finding.
Actions With Statutory Deadlines
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Casey’s june beetle ........................................................................................................................................................
6 Birds from Eurasia .......................................................................................................................................................
5 Bird species from Colombia and Ecuador ..................................................................................................................
Queen Charlotte goshawk ..............................................................................................................................................
Ozark hellbender 4 ..........................................................................................................................................................
Altamaha spinymussel 3 ..................................................................................................................................................
6 Birds from Peru and Bolivia ........................................................................................................................................
Loggerhead sea turtle (assist National Marine Fisheries Service) 5 ..............................................................................
2 mussels (rayed bean (LPN = 2), snuffbox No LPN) 5 .................................................................................................
CA golden trout 4 ............................................................................................................................................................
Black-footed albatross ....................................................................................................................................................
Mojave fringe-toed lizard 1 ..............................................................................................................................................
Kokanee—Lake Sammamish population 1 .....................................................................................................................
Cactus ferruginous pygmy-owl 1 .....................................................................................................................................
Northern leopard frog .....................................................................................................................................................
Tehachapi slender salamander ......................................................................................................................................
Coqui Llanero .................................................................................................................................................................
Dusky tree vole ...............................................................................................................................................................
Leatherside chub (from 206 species petition) ................................................................................................................
Platte River caddisfly (from 206 species petition) 5 ........................................................................................................
3 Texas moths (Ursia furtiva, Sphingicampa blanchardi, Agapema galbina) (from 475 species petition) ....................
3 South Arizona plants (Erigeron piscaticus, Astragalus hypoxylus, Amoreuxia gonzalezii) (from 475 species petition).
5 Central Texas mussel species (3 from 475 species petition) .....................................................................................
14 parrots (foreign species) ............................................................................................................................................
Mohave Ground Squirrel 1 ..............................................................................................................................................
Western gull-billed tern ...................................................................................................................................................
HI yellow-faced bees ......................................................................................................................................................
OK grass pink (Calopogon oklahomensis) 1 ...................................................................................................................
Ashy storm-petrel 5 .........................................................................................................................................................
Honduran emerald ..........................................................................................................................................................
Southeastern pop. snowy plover & wintering pop. of piping plover 1 ............................................................................
Eagle Lake trout 1 ...........................................................................................................................................................
32 Pacific Northwest mollusk species (snails and slugs) 1 ............................................................................................
42 snail species (Nevada and Utah) ..............................................................................................................................
Spring Mountains checkerspot butterfly .........................................................................................................................
10 species of Great Basin butterfly ................................................................................................................................
404 Southeast species ...................................................................................................................................................
Franklin’s bumble bee 4 ..................................................................................................................................................
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Final listing determination.
Final listing determination.
Final listing determination.
Final listing determination.
Final listing determination.
Final listing determination.
Final listing determination.
Final listing determination.
Final listing determination.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding/
Proposed listing.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
12-month petition finding.
90-day petition finding.
90-day petition finding.
90-day petition finding.
90-day petition finding.
90-day petition finding.
90-day petition finding.
90-day petition finding.
90-day petition finding.
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Federal Register / Vol. 76, No. 172 / Tuesday, September 6, 2011 / Proposed Rules
ACTIONS FUNDED IN FY 2010 AND FY 2011 BUT NOT YET COMPLETED—Continued
Species
Action
American eel 4 .................................................................................................................................................................
Leona’s little blue 4 ..........................................................................................................................................................
Aztec gilia 5 .....................................................................................................................................................................
White-tailed ptarmigan 5 ..................................................................................................................................................
San Bernardino flying squirrel 5 ......................................................................................................................................
Bicknell’s thrush 5 ............................................................................................................................................................
Chimpanzee ....................................................................................................................................................................
Sonoran talussnail 5 ........................................................................................................................................................
2 AZ Sky Island plants (Graptopetalum bartrami and Pectis imberbis) 5 ......................................................................
I’iwi 5 ................................................................................................................................................................................
Humboldt marten ............................................................................................................................................................
Desert massasauga ........................................................................................................................................................
Western glacier stonefly (Zapada glacier) ......................................................................................................................
Thermophilic ostracod (Potamocypris hunteri) ...............................................................................................................
Sierra Nevada red fox 5 ..................................................................................................................................................
Boreal toad (eastern or southern Rocky Mtn population) 5 ............................................................................................
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
90-day
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
petition
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
finding.
High-Priority Listing Actions
species 2
20 Maui-Nui candidate
(17 plants, 3 tree snails) (14 with LPN = 2, 2 with LPN = 3, 3 with LPN = 8) .........
8 Gulf Coast mussels (southern kidneyshell (LPN = 2), round ebonyshell (LPN = 2), Alabama pearlshell (LPN = 2),
southern sandshell (LPN = 5), fuzzy pigtoe (LPN = 5), Choctaw bean (LPN = 5), narrow pigtoe (LPN = 5), and
tapered pigtoe (LPN = 11)) 4.
Umtanum buckwheat (LPN = 2) and white bluffs bladderpod (LPN = 9) 4 ....................................................................
Grotto sculpin (LPN = 2) 4 ..............................................................................................................................................
2 Arkansas mussels (Neosho mucket (LPN = 2) & Rabbitsfoot (LPN = 9)) 4 ...............................................................
Diamond darter (LPN = 2) 4 ............................................................................................................................................
Gunnison sage-grouse (LPN = 2) 4 ................................................................................................................................
Coral Pink Sand Dunes Tiger Beetle (LPN = 2) 5 ..........................................................................................................
Lesser prairie chicken (LPN = 2) ...................................................................................................................................
4 Texas salamanders (Austin blind salamander (LPN = 2), Salado salamander (LPN = 2), Georgetown salamander
(LPN = 8), Jollyville Plateau (LPN = 8)) 3.
5 SW aquatics (Gonzales Spring Snail (LPN = 2), Diamond Y springsnail (LPN = 2), Phantom springsnail (LPN =
2), Phantom Cave snail (LPN = 2), Diminutive amphipod (LPN = 2)) 3.
2 Texas plants (Texas golden gladecress (Leavenworthia texana) (LPN = 2), Neches River rose-mallow (Hibiscus
dasycalyx) (LPN = 2)) 3.
4 AZ plants (Acuna cactus (Echinomastus erectocentrus var. acunensis) (LPN = 3), Fickeisen plains cactus
(Pediocactus peeblesianus fickeiseniae) (LPN = 3), Lemmon fleabane (Erigeron lemmonii) (LPN = 8), Gierisch
mallow (Sphaeralcea gierischii) (LPN = 2)) 5.
FL bonneted bat (LPN = 2) 3 ..........................................................................................................................................
3 Southern FL plants (Florida semaphore cactus (Consolea corallicola) (LPN = 2), shellmound applecactus
(Harrisia (= Cereus) aboriginum (= gracilis)) (LPN = 2), Cape Sable thoroughwort (Chromolaena frustrata) (LPN
= 2)) 5.
21 Big Island (HI) species 5 (includes 8 candidate species—6 plants and 2 animals; 4 with LPN = 2, 1 with LPN =
3, 1 with LPN = 4, 2 with LPN = 8).
12 Puget Sound prairie species (9 subspecies of pocket gopher (Thomomys mazama ssp.) (LPN = 3), streaked
horned lark (LPN = 3), Taylor’s checkerspot (LPN = 3), Mardon skipper (LPN = 8)) 3.
2 TN River mussels (fluted kidneyshell (LPN = 2), slabside pearlymussel (LPN = 2)) 5 ..............................................
Jemez Mountain salamander (LPN = 2) 5 ......................................................................................................................
Proposed listing.
Proposed listing.
Proposed
Proposed
Proposed
Proposed
Proposed
Proposed
Proposed
Proposed
listing.
listing.
listing.
listing.
listing.
listing.
listing.
listing.
Proposed listing.
Proposed listing.
Proposed listing.
Proposed listing.
Proposed listing.
Proposed listing.
Proposed listing.
Proposed listing.
Proposed listing.
1 Funds
for listing actions for these species were provided in previous FYs.
funds for these high-priority listing actions were provided in FY 2008 or 2009, due to the complexity of these actions and competing
priorities, these actions are still being developed.
3 Partially funded with FY 2010 funds and FY 2011 funds.
4 Funded with FY 2010 funds.
5 Funded with FY 2011 funds.
erowe on DSK5CLS3C1PROD with PROPOSALS2
2 Although
We have endeavored to make our
listing actions as efficient and timely as
possible, given the requirements of the
relevant law and regulations, and
constraints relating to workload and
personnel. We are continually
considering ways to streamline
processes or achieve economies of scale,
such as by batching related actions
together. Given our limited budget for
implementing section 4 of the Act, these
actions described above collectively
constitute expeditious progress.
VerDate Mar<15>2010
15:35 Sep 02, 2011
Jkt 223001
Hylaeus anthracinus, H. assimulans,
H. facilis, H. hilaris, H. kuakea, H.
longiceps, and H. mana will be added
to the list of candidate species upon
publication of this 12-month finding.
We will continue to monitor the status
of these species as new information
becomes available. This review will
determine if a change in status is
warranted, including the need to make
prompt use of emergency listing
procedures.
PO 00000
Frm 00034
Fmt 4701
Sfmt 4702
We intend that any proposed listing
action for the seven species of Hawaiian
yellow-faced bees will be as accurate as
possible. Therefore, we will continue to
accept additional information and
comments from all concerned
governmental agencies, the scientific
community, industry, or any other
interested party concerning this finding.
References Cited
A complete list of all references cited
in this document is available on the
E:\FR\FM\06SEP2.SGM
06SEP2
Federal Register / Vol. 76, No. 172 / Tuesday, September 6, 2011 / Proposed Rules
Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
Authors
The primary authors of this notice are
the staff members of the Pacific Islands
erowe on DSK5CLS3C1PROD with PROPOSALS2
Internet at https://www.regulations.gov
and upon request from the Pacific
Islands Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
VerDate Mar<15>2010
15:35 Sep 02, 2011
Jkt 223001
Authority
PO 00000
Frm 00035
55203
Dated: August 22, 2011.
Daniel M. Ashe,
Director, U.S. Fish and Wildlife Service.
[FR Doc. 2011–22433 Filed 9–2–11; 8:45 am]
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BILLING CODE 4310–55–P
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Agencies
[Federal Register Volume 76, Number 172 (Tuesday, September 6, 2011)]
[Proposed Rules]
[Pages 55170-55203]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2011-22433]
[[Page 55169]]
Vol. 76
Tuesday,
No. 172
September 6, 2011
Part II
Department of the Interior
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Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; 12-Month Finding on Five
Petitions To List Seven Species of Hawaiian Yellow-faced Bees as
Endangered; Proposed Rule
Federal Register / Vol. 76 , No. 172 / Tuesday, September 6, 2011 /
Proposed Rules
[[Page 55170]]
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R1-ES-2010-0012; MO 92210-0-008]
Endangered and Threatened Wildlife and Plants; 12-Month Finding
on Five Petitions To List Seven Species of Hawaiian Yellow-faced Bees
as Endangered
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), announce a
12-month 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 to
designate critical habitat under the Endangered Species Act of 1973, as
amended (Act). After review of all available scientific and commercial
information, we find that listing these seven species of Hawaiian
yellow-faced bees is warranted. Currently, however, listing these seven
species of Hawaiian yellow-faced bees is precluded by higher priority
actions to amend the Lists of Endangered and Threatened Wildlife and
Plants. Upon publication of this 12-month petition finding, we will add
these seven species of Hawaiian yellow-faced bees to our candidate
species list. We will develop a proposed rule to list these seven
species of Hawaiian yellow-faced bees as our priorities allow. We will
make any determinations on critical habitat during development of the
proposed listing rule. In any interim period we will address the status
of the candidate taxa through our annual Candidate Notice of Review
(CNOR).
DATES: The finding announced in this document was made on September 6,
2011.
ADDRESSES: This finding is available on the Internet at https://www.regulations.gov at Docket Number FWS-R1-ES-2010-0012. Supporting
documentation we used in preparing this finding is available for public
inspection, by appointment, during normal business hours at the U.S.
Fish and Wildlife Service, Pacific Islands Fish and Wildlife Office,
300 Ala Moana Boulevard, Room 3-122, Honolulu, HI 96850. Please submit
any new information, materials, comments, or questions concerning this
finding to the above street address.
FOR FURTHER INFORMATION CONTACT: Loyal Mehrhoff, Field Supervisor,
Pacific Islands Fish and Wildlife Office (see ADDRESSES); by telephone
at 808-792-9400; or by facsimile at 808-792-9581. If you use a
telecommunications device for the deaf (TTD) please call the Federal
Information Relay Service (FIRS) at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Act (16 U.S.C. 1531 et seq.) requires
that, for any petition to revise the Federal Lists of Endangered and
Threatened Wildlife and Plants that contains substantial scientific or
commercial information that listing a species may be warranted, we make
a finding within 12 months of the date of receipt of the petition. In
this finding, we determine whether the petitioned action is: (a) Not
warranted, (b) warranted, or (c) warranted, but immediate proposal of a
regulation implementing the petitioned action is precluded by other
pending proposals to determine whether species are endangered or
threatened, and expeditious progress is being made to add or remove
qualified species from the Federal Lists of Endangered and Threatened
Wildlife and Plants. Section 4(b)(3)(C) of the Act requires that we
treat a petition for which the requested action is found to be
warranted but precluded as though resubmitted on the date of such
finding, that is, requiring a subsequent finding to be made within 12
months. We must publish these 12-month findings in the Federal
Register.
Previous Federal Actions
On March 23, 2009, we received five petitions dated March 23, 2009,
from Scott Hoffman Black, Executive Director of the Xerces Society,
requesting that seven species of Hawaiian yellow-faced bees be listed
as endangered under the Act and critical habitat be designated.
Each petition contained information regarding the species' taxonomy
and ecology, historical and current distribution, present status, and
current and potential threats. We acknowledged the receipt of the
petitions in a letter to Mr. Black, dated May 8, 2009. In that letter
we also stated that issuing an emergency regulation temporarily listing
the species under section 4(b)(7) of the Act was not warranted at that
time. We published the 90-day finding in the Federal Register on June
16, 2010 (75 FR 34077). This notice constitutes the 12-month finding on
the March 23, 2009, petitions to list the seven species of Hawaiian
yellow-faced bees as endangered.
Species Information
Overview of the Genus Hylaeus
The seven species of bees described in this finding 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 endemic to single islands (Magnacca 2007a, p. 174). All 60
Hawaiian species are in the subgenus Nesoprosopis (Magnacca and
Danforth 2006, p. 393). The Hawaiian 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 family is
comprised of over 2,000 species, all of which are solitary nesting
(unlike social wasps and bees), although a few do nest in close
vicinity to each other.
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).
A great deal of our knowledge on Hawaiian Hylaeus bees is based
upon surveys by Robert Cyril Layton Perkins, a distinguished British
entomologist and naturalist renowned for his pioneering work on the
insects of the Hawaiian Islands, particularly the Hymenoptera
(sawflies, wasps, bees, and ants), in the early 20th Century. His
surveys were conducted between 1892 and 1906, and form the basis for
most of the historical records of Hylaeus in the Hawaiian Islands.
According to Perkins, Hylaeus species were ``almost the most ubiquitous
of any Hawaiian insects'' (Perkins 1913, p. lxxxi). However, there are
about 90 years between Perkins' surveys and the most recent surveys
conducted in the late 1990s for Hylaeus bees in the Hawaiian Islands.
Surveys in more recent years (1998-2010) for Hylaeus spp. in the
Hawaiian Islands have largely involved targeted collecting on specific
flowering plants (Daly and Magnacca 2003, pp. 217-233; Magnacca in
litt. 2011, p. 5), rather than survey methods such as pan trapping or
Pollard walks (see below). While this means the numbers of individuals
and species observed are not strictly quantifiable by effort, the
probability of collecting species actually present is
[[Page 55171]]
higher (Magnacca in litt. 2011, p. 5). Because the number and diversity
of Hylaeus spp. tend to be locally concentrated rather than widely
distributed, randomized and more quantifiable surveys such as pan
trapping and Pollard walks are actually less effective means of
locating Hylaeus spp. (Magnacca in litt. 2011, p. 5). Pan trapping
involves the use of shallow pans of fluid, and relies on the organism
falling or flying into the fluid preservative. Pollard walks involve
observers walking along a fixed transect route and recording the
insects observed.
The recent Hylaeus spp. survey efforts are not easily comparable to
Perkins' collections, which are considered now to have been conducted
opportunistically. For example, Perkins collected higher numbers of
individuals and species in certain areas, including coastal areas that
were much less disturbed at the time, and some species, such as H.
facilis, were formerly very common but have almost entirely disappeared
(Magnacca in litt. 2011, p. 5).
Life History of Genus Hylaeus
The following discussion includes all Hawaiian Hylaeus species, and
specific information about the seven petitioned Hylaeus species.
Hawaiian Hylaeus species are grouped within two categories: Ground-
nesting species that require relatively dry conditions, and wood-
nesting species that are often found within wetter areas (Zimmerman
1972, p. 533; Daly and Magnacca 2003, p. 11). Nests of Hylaeus species
are usually constructed opportunistically within dead twigs or plant
stems, or other similarly small natural cavities under bark or rocks
(i.e., they seek out existing cavities that they suit to their own
needs). This is unlike the nests of many other bee species, which are
purposefully excavated or constructed underground. Like other Hylaeus,
Hawaiian Hylaeus 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 female Hylaeus bee lays eggs in
brood cells 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 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 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).
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. 2,524-2,528; Cox and Elmqvist
2000, p. 1,238; Sahli et al. 2008, p. 1) have demonstrated Hawaiian
Hylaeus species almost exclusively visit native plants to collect
nectar and pollen, pollinating those plants in the process. Hylaeus
bees are very rarely found visiting nonnative plants for nectar and
pollen (Magnacca 2007a, 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 nonnative species changes with
elevation. In that study, eight flowering plants were observed at six
sites, which ranged in elevation from approximately 2,900 to 7,900 feet
(ft) (approximately 880 to 2,400 meters (m)). The study also found the
proportion of native pollinators changed along the elevation gradient;
at least 40 to 50 percent of visits were from nonnative pollinators at
low elevation, as opposed to 4 to 20 percent of visits by nonnative
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 suggest Hylaeus
may not be easily replaceable by nonnative pollinators (Sahli et al.
2008, p. 1).
For some of the seven Hawaiian yellow-faced bees addressed in this
finding, we have information about the specific host plants they visit
for nectar and pollen. For some species, we have also identified
primary host plants visited (see description of the species where
noted). However, for others, we lack detailed information on the
specific host plants visited for foraging. Nonetheless, researchers
believe native plants both endemic and indigenous to the Hawaiian
Islands are essential to the survival of the Hylaeus species (Hopper et
al. 1996, pp. 8-9; Daly and Magnacca 2003, pp. 217-229; Magnacca 2007a,
pp. 185-186).
Hawaiian Island Ecosystems
The five Hawaiian Island ecosystems that support the seven Hawaiian
yellow-faced bees addressed in this 12-month finding are described in
the following section. See Table 1 below for a list of the ecosystems
from which each species is reported. Because Hawaiian Hylaeus spp.,
including these seven, are believed to be essential pollinators of the
native Hawaiian plant fauna, we are providing this background
information on the different ecosystems in which they occur to better
elaborate upon the specific threats found in the five ecosystem types.
Table 1--Current (and Historical) Distribution of the Seven Yellow-Faced Bees by Ecosystem Type and Island
--------------------------------------------------------------------------------------------------------------------------------------------------------
Ecosystems
Species and number of current -----------------------------------------------------------------------------------------------------------------------
populations Coastal Lowland dry Lowland mesic Lowland wet Montane mesic Montane dry
--------------------------------------------------------------------------------------------------------------------------------------------------------
H. anthracinus, 13 populations.. HI, MA, MO, OA.... HI, KAH, (*LA), N/A............... N/A............... .................. HI.
MA, (*MO), (*OA).
H. assimulans, 5 populations.... KAH, (*MA), (*OA). LA, MA, (*OA)..... N/A............... N/A............... .................. N/A.
H. facilis, 2 populations....... (*MA), MO, (*OA).. (*LA), (*OA)...... (*LA), (*MA), (*MA), OA......... (*MO)............. N/A.
(*OA).
H. hilaris, 1 population........ (*LA), (*MA), MO.. (*MA)............. N/A............... N/A............... .................. N/A.
H. kuakea, 2 populations........ N/A............... N/A............... OA................ N/A............... .................. N/A.
H. longiceps, 6 populations..... LA, MA, MO, OA.... LA, (*MA), (*MO).. N/A............... N/A............... .................. N/A.
[[Page 55172]]
H. mana, 1 population........... N/A............... N/A............... OA................ N/A............... .................. N/A.
--------------------------------------------------------------------------------------------------------------------------------------------------------
HI = Hawaii (Island); KAH = Kahoolawe; LA = Lanai; MA = Maui; MO = Molokai; OA = Oahu;
(*XX) denotes a historical population; N/A means no population records
Coastal Ecosystem
The coastal ecosystem is found on all of the main Hawaiian Islands,
with the highest species diversity found in the least populated coastal
areas of Hawaii, Maui, Molokai, Kahoolawe, Oahu, and Kauai, and their
associated islets, and extends from sea level to approximately 1,000 ft
(approximately 300 m) in elevation. The coastal vegetation zone is
typically dry, with annual rainfall of less than approximately 20
inches (in) (50.8 centimeters (cm)); however windward rainfall may be
high enough (up to approximately 40 in (1,000 mm)) to support mesic-
associated and sometimes wet-associated vegetation (Gagne and Cuddihy
1999, pp. 54-66). Compared to dry and mesic ecosystems, biological
diversity (number of species) is low to moderate in the coastal
ecosystem, but may include some specialized plants and animals such as
nesting seabirds and the rare native plant Sesbania tomentosa (ohai)
(The Nature Conservancy (TNC) 2006a). Sesbania tomentosa formerly
occurred widely in lower elevation dry habitat on all of the main
islands and at least on Necker and Nihoa of the Northwestern Hawaiian
Islands. The species is now scattered throughout its former range, and
is restricted to relic populations on sandy beaches, on dunes, on soil
pockets on lava, and along pond margins (Wagner et al. 1990, p. 705).
The dominant native vegetation in coastal ecosystems is the shrub
Scaevola sericea (naupaka kahakai) (Alpha et al. 1996, p. 86). Other
common native plant species include Ipomoea pes-caprae (beach morning-
glory), Sporobolus virginicus (beach dropseed), Jacquemontia ovata (pau
o Hiiaka), and Sesuvium portulacastrum (akulikuli or sea purslane)
(Wagner et al. 1999, pp. 57-59). Among the Hylaeus species addressed in
this finding, five are known from coastal ecosystems, including H.
anthracinus, H. assimulans, H. facilis, H. hilaris, and H. longiceps.
Lowland Dry Ecosystem
The lowland dry ecosystem includes shrublands and forests below
approximately 3,300 ft (1,000 m) in elevation that receive less than 50
in (127 cm) annual rainfall, or are in otherwise prevailingly dry
substrate conditions. Areas consisting of predominantly native species
in the lowland dry ecosystem are now rare. This ecosystem is found on
the islands of Hawaii, Maui, Molokai, Lanai, Kahoolawe, Oahu, and
Kauai, and is best represented on the leeward sides of the islands
(Gagne and Cuddihy 1999, p. 67). Biological diversity is low to
moderate in this ecosystem, and includes specialized animals and plants
such as the Hawaiian owl or pueo (Asio flammeus sandwichensis) and
Santalum ellipticum (iliahialoe) (Wagner et al. 1999, pp. 1,220-1,221;
TNC 2006b).
Hylaeus anthracinus, H. assimulans, H. facilis, and H. longiceps
are known from lowland dry forests. These forests are typically
dominated by Diospyros sandwicensis (lama), Erythrina sandwicensis
(wiliwili), Nestegis sandwicensis (olopua), or Metrosideros polymorpha
(ohia) and a diversity of native shrubs growing within the understory
(Gagne and Cuddihy 1999, pp. 72-74).
Lowland Mesic Ecosystem
The lowland mesic ecosystem includes a variety of grasslands,
shrublands, and forests, below approximately 3,300 ft (1,000 m) in
elevation, that receive between 50 and 75 in (127 and 191 cm) annual
rainfall, or are in otherwise mesic substrate conditions (TNC 2006c).
In the Hawaiian Islands, this ecosystem is found on Hawaii, Maui,
Molokai, Lanai, Oahu, and Kauai, on both windward and leeward sides of
the islands. Biological diversity is high in this system (TNC 2006c).
Lowland mesic forests are typically dominated by Acacia koa (koa),
Diospyros sandwicensis, Metrosideros polymorpha, or Nestegis
sandwicensis, and a diversity of understory trees and native shrubs
growing below the canopy species (Gagne and Cuddihy 1999, p. 80-82).
Historically, Hylaeus facilis was known from lowland mesic forest, but
currently only H. kuakea and H. mana are found in this habitat.
Lowland Wet Ecosystem
The lowland wet ecosystem is generally found below approximately
3,300 ft (1,000 m) in elevation on the windward sides of the main
Hawaiian Islands, except Kahoolawe (Gagne and Cuddihy 1999, p. 85; TNC
2006d). These areas include a variety of wet grasslands, shrublands,
and forests that receive greater than 75 in (191 cm) annual
precipitation, or are in otherwise wet substrate conditions (TNC
2006d). Biological diversity is high in this system (TNC 2006d). The
majority of lowland wet forests are dominated by Metrosideros
polymorpha, with understory trees such as Psychotria spp. (kopiko) and
Antidesma platyphyllum (hame) (Gagne and Cuddihy 1999, p. 87).
Currently, Hylaeus facilis is known from lowland wet forest (Daly and
Magnacca 2003, p. 81).
Montane Dry Ecosystem
The montane dry ecosystem is composed of natural communities
(shrublands, grasslands, forest) found at elevations between
approximately 3,300 and 6,600 ft (1,000 and 2,000 m), in areas where
annual precipitation is less than 50 in (127 cm), or otherwise in dry
substrate conditions (TNC 2006g). Montane dry forests occur on the
leeward sides of the islands of Maui and Hawaii, and biological
diversity is moderate (Gagne and Cuddihy 1999, p. 93; TNC 2006g).
Montane dry forests are dominated by some combination of Acacia koa,
Sophora chrysophylla) (mamame), Metrosideros polymorpha, and rarely,
Chamaesyce olowaluana (akoko) (Gagne and Cuddihy, p. 95). In 2004, a
single individual of H. anthracinus was collected in montane dry forest
on Hawaii Island.
Specific Information on Hylaeus anthracinus
Taxonomy and Description
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). Nesoprosopis was
reduced to a subgenus of Hylaeus in
[[Page 55173]]
1923 (Meade-Waldo 1923, p. 1). Although the distinctness of this
species remains unquestioned, recent genetic evidence (Magnacca and
Brown 2010, pp. 5-7) suggests H. anthracinus may be composed of three
cryptic (not recognized) species or subspecies that represent the
populations on Hawaii, Maui and Kahoolawe, and Molokai and Oahu.
However, this has not been established scientifically; therefore, we
treat H. anthracinus as a single species in this finding.
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 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, Scaevola sericea, 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).
Tournefortia argentea was first collected on Oahu in 1864-1865, and is
naturalized and documented from all of the main islands except
Kahoolawe (Wagner et al. 1999, p. 398). 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
and lowland dry forest habitats up to 2,000 ft (610 m) in elevation on
the islands of Hawaii, Lanai, Maui, Molokai, and Oahu. Between 1997 and
2008, surveys for Hawaiian Hylaeus were conducted at 43 sites
throughout the Hawaiian Islands that were either historical collecting
localities for H. anthracinus, or potentially suitable habitat for this
species. Hylaeus anthracinus was observed at 13 of the 43 survey sites,
but had disappeared from each of the 9 historically occupied sites
surveyed (Daly and Magnacca 2003, p. 217; Magnacca 2007b, p. 44).
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 Magnacca 2003, p. 55; Magnacca 2007a, pp. 186-188).
Hylaeus anthracinus is currently known from 13 small patches of
coastal and lowland dry forest habitat (Magnacca 2005a, p. 2): five
locations on the island of Hawaii; one location on Kahoolawe; two
locations on Maui; three locations on Molokai; and two locations on
Oahu (Daly and Magnacca 2003, p. 217; Magnacca 2005a, p. 2; Magnacca
2007b, p. 44). These 13 locations supported small populations of H.
anthracinus, but the number of individual bees is unknown. In 2004, a
single individual was collected in montane dry forest on the island of
Hawaii; however, the presence of additional individuals has not been
confirmed at this site (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). The species is believed to be extirpated from Lanai (Daly and
Magnacca 2003, p. 55). Additionally, during surveys between 1997 and
2008, H. anthracinus was absent from 17 other sites on Hawaii, Maui,
Molokai, and Oahu with potentially suitable habitat from which other
species of Hylaeus were collected (Daly and Magnacca 2003; Magnacca,
University of Hawaii at Hilo, pers. comm. 2008a).
Hawaii Island
Hylaeus anthracinus was first described by Perkins (1899, p. 100)
from specimens collected by F. Smith on the Kona (west) coast at
Kealakekua Bay. In the intervening 99 years, H. anthracinus appears to
have declined significantly throughout its historical range on this
coastline. Between 1997 and 2008, researchers thoroughly surveyed the
area around Kealakekua Bay and Keei to the south, but found no species
of Hylaeus and observed that most of these areas are either dominated
by invasive, nonnative plants, such as Leucaena leucephala (koa haole),
or lack vegetation entirely (Magnacca, pers. comm. 2008a). Hylaeus
anthracinus is currently found in five locations in coastal and lowland
dry forest on the leeward (west) side of the island, including
Kohanaiki; Kaloko-Honokohau National Historic Park (NHP); Makalawena
Beach; the Mahaiula section of Kekaha Kai (Kona Coast) State Park; and
Kaulana Bay near Ka Lae (South Point). In addition, there is one recent
collection from montane dry forest in the U.S. Army's Pohakuloa
Training Area, in the northern part of the island. Collection reports
from these six areas follow:
(A) Kohanaiki: Hylaeus anthracinus was collected in coastal habitat
on Tournefortia argentea at this location near Puhili Point by Magnacca
(2007b, p. 44). Kohanaiki is an area of land granted to indigenous
Hawaiians in 1995 for cultural and recreational preservation and
pursuits (Kohanaiki Ohana 1995 (https://www.kohanaiki.org/)). There is
some possibility for increased recreational impact to the area, if and
when adjacent privately owned parcels are developed, as is currently
planned (Kohanaiki Ohana 1995 (https://www.kohanaiki.org/)).
(B) Kaloko-Honokohau NHP: In 2007, researchers collected Hylaeus
anthracinus in coastal habitat in Kaloko-Honokohau NHP, which is just
south of Kohanaiki, and managed by the National Park Service (NPS) (P.
Aldrich, University of Hawaii at Manoa, pers. comm. 2008a; Magnacca,
pers. comm. 2008c).
(C) Makalawena Beach: Researchers collected Hylaeus anthracinus in
coastal habitat in south Kona at Makalawena Beach in 2007 (P. Aldrich,
pers. comm., July 2008a). Inaccessible by motor vehicle, visitors must
hike to the beach on a trail that begins in nearby Kekaha Kai State
Park. Makalawena Beach is located on private land owned by Kamehameha
Schools.
(D) Mahaiula Section of Kekaha Kai State Park: Researchers
collected Hylaeus anthracinus in coastal habitat in the Mahaiula
section of Kekaha Kai State Park in 2007 (P. Aldrich, unpublished
data). The park is managed by the Hawaii Department of Land and Natural
Resources' (DLNR) Division of State Parks, and is open to the public
daily. This section of the park is accessed by a 1.5-mile (mi) (1.6-
kilometer (km)) unpaved road from the main highway (Queen Kaahumanu
Highway (Hwy 19)), and offers public
[[Page 55174]]
recreational opportunities for swimming and beach-related activities,
such as hiking, picnicking, and boating (https://www.hawaiistateparks.org/hawaiistateparks/parks/hawaii/index.cfm?park_id=47).
(E) Kaulana Bay: Hylaeus anthracinus appears to be restricted to an
area of 5,000-10,000 year-old lava flows east of Ka Lae at Kaulana Bay,
where it and other species of Hylaeus were collected in 1999 and 2002
(Magnacca 2007a, p. 181). The substrate of these lava flows is distinct
from the surrounding areas covered by Pahala ash (Magnacca, pers. comm.
2010b). The area near Ka Lae, at the southernmost tip of the island of
Hawaii, is believed to be the best coastal habitat for Hylaeus on the
island. However, H. anthracinus was absent from several sites with
potentially suitable vegetation near Ka Lae and other sites to the east
along the coast, including Kalu, Kaalualu, and Mahana, where other
Hylaeus species were collected. The population of H. anthracinus at
Kaulana Bay appears highly localized, and may have more stringent
habitat requirements related to localized substrate type than other
species of Hawaiian Hylaeus found in nearby areas (e.g., H. difficilis
and H. flavipes). The Ka Lae area, including Kaulana Bay, is registered
as a National Historic Landmark District and a large portion of the
area is primarily owned by the State's Department of Hawaiian Home
Lands (DHHL), although a smaller portion is privately owned. Public
access to Kaulana Bay is not restricted, and the area is used for
recreational activities such as off-road vehicle use (Magnacca, pers.
comm. 2008a).
(F) U.S. Army's Pohakuloa Training Area (PTA): In 2004, one male
Hylaeus anthracinus was collected on the southern slopes of Mauna Kea
in montane dry forest habitat in the U.S. Army's PTA at approximately
5,200-5,400 ft (1,590-1,650 m) in elevation (Magnacca 2007b, p. 44).
The specimen was found inside the fruit capsule of the federally
endangered plant, Hedyotis coriacea (kioele). Hylaeus anthracinus has
not been observed at the PTA since the collection made in 2004
(Magnacca 2007b, p. 44). It is unknown if this collection was a single
vagrant individual or from an established population at the PTA
(Magnacca 2007b, p. 44).
Kahoolawe Island
Previously unknown on Kahoolawe, a population of Hylaeus
anthracinus was discovered in 2002 in coastal habitat at Pali o
Kalapakea, where four specimens were collected at an elevation of 1,000
ft (300 m) (Daly and Magnacca 2003; Magnacca, pers. comm. 2008a).
However, this species was absent from potentially suitable habitat
located at Kamohio on the southeastern coast of the island where other
Hylaeus species were collected. Overgrazing by introduced cattle and
goats, and bombing and target practice by the U.S. military, have led
to soil erosion resulting in the loss of almost all of the coastal and
lowland dry forest habitat on this island (Warren 2004, p. 461). In
1993, Congress ended military use on Kahoolawe, and the Kahoolawe
Island Reserve Commission (KIRC) was created to manage land use and
restore Kahoolawe's natural resources (Dept. of Defense, p. 1). Access
to the island is limited and controlled by KIRC, and activities
conducted on the island include fishing, habitat restoration,
historical preservation, and education. Commercial enterprises are
currently prohibited on the island (Warren 2004, p. 1).
Maui
Perkins (1899, p. 100) originally described Hylaeus anthracinus as
abundant in coastal and lowland habitat on the island of Maui, where it
was known from four sites. Perkins' primary collection site for coastal
bees on Maui was the Wailuku sandhills, which once supported a diverse
bee fauna. Lacking adequate descriptions, researchers were unable to
relocate two of the Perkins collection sites during recent surveys, but
two sites were relocated and surveyed in 1999 and 2001 (Magnacca 2007a,
p. 173). Hylaeus anthracinus has also been collected at Kanaio on the
lower southern slopes of Haleakala, an unusual location for this
otherwise exclusively coastal species. The species was also collected
at the coast nearby, at Manawainui. Descriptions of these three sites
follows:
(A) Wailuku Sand Hills: Formerly a large expanse of coastal dune
habitat, the Wailuku sand hills remain as small remnant dunes and only
one, at Waiehu, contains intact native vegetation potentially suitable
for Hylaeus bees. This remnant coastal sand dune covers less than 2.5
acres (ac) (1 hectare (ha)) on State lands near a golf course. Hylaeus
anthracinus was not observed during the 1999 and 2001 surveys in this
location (Daly and Magnacca 2003, p. 217). The rest of the dunes have
been destroyed by development or are overgrown with the nonnative plant
Prosopis pallida (kiawe). Researchers observed that the Kahului section
of the dunes, located south of the native remnant dune, no longer
contains potentially suitable habitat for species of Hylaeus (Magnacca
2007a, p. 182).
(B) Kanaio Natural Area Reserve: Hylaeus anthracinus was collected
in 1999 in remnant native lowland dry forest in the State's Kanaio
Natural Area Reserve (NAR) on the southern slopes of Haleakala at 2,000
ft (600 m) in elevation (Daly and Magnacca 2003, p. 217). Kanaio NAR is
a State-protected area of approximately 876 ac (355 ha), and contains
patches of lowland dry forest and shrub lands. The State plans to
rehabilitate habitat in the Kanaio NAR by excluding feral ungulates
with fencing, managing weeds, and planting native species (https://hawaii.gov/dlnr/dofaw/rpc/projects-on-maui).
(C) Manawainui Gulch: In 1999, Hylaeus anthracinus was collected at
this coastal site on land owned by the State's DHHL (Magnacca, pers.
comm. 2008a). The site is east of Kahikinui, and should not be confused
with the Manawainui Valley, which is east of Kaupo, or Manawainui Gulch
at Ukumehame on west Maui.
Molokai
Perkins collected Hylaeus anthracinus at Kaulawai [Kauluwai] and
two unknown sites: the lower slopes of the north Molokai mountains and
the ``Molokai plains'' (Perkins 1899; Daly and Magnacca 2003, p. 55).
Hylaeus anthracinus occurred in three of five sites surveyed between
1999 and 2005. These locations include TNC's Moomomi Preserve on
Molokai's northwest coast, and Hoolehua Beach and Kaupikiawa, both
located on the Kalaupapa peninsula (Magnacca, pers. comm. 2008a). This
species was not observed at several other sites with potentially
suitable habitat, including sand dune habitat near the Kaluakoi resort
on Molokai's west coast (Magnacca, pers. comm. 2008a). Collection
reports of these sites follow:
(A) Moomomi Preserve: Between 1999 and 2001, researchers collected
H. anthracinus and H. longiceps from an area of native vegetation in
coastal dune habitat within Moomomi Preserve (Magnacca 2007a, p. 181).
Moomomi Preserve contains intact coastal dunes dominated by native
vegetation, as well as dune and inland areas dominated by nonnative
vegetation.
(B) Hoolehua Beach and Kaupikiawa: In 2005, Hylaeus anthracinus was
collected at a coastal site above Hoolehua Beach near the tip of the
Kalaupapa peninsula, and at Kaupikiawa, just to the east (Magnacca
2007b, p. 181). Both sites are located within Kalaupapa NHP, which is
cooperatively managed by the NPS, DHHL, and the State's DLNR and
Departments of Health (DOH) and Transportation (DOT). The areas on the
[[Page 55175]]
east side of the Kalaupapa peninsula are largely rocky and devoid of
vegetation, but contain scattered patches of native coastal vegetation,
similar to Ka Lae on the island of Hawaii (Magnacca 2007a, p. 181).
Oahu
Hylaeus anthracinus was historically known from seven sites on the
island of Oahu, although two of the coastal sites were not conclusively
identified by Perkins and the exact locations cannot now be determined
(Perkins 1899, p. 100). This species appears to have declined
precipitously since Perkins' collecting period on Oahu (1892-1906) and
is currently only known from two sites, Kaena Point NAR and Mokuauia
(Goat Island). Between 1997 and 2008, H. anthracinus was not found
during surveys of five of its historical Perkins-era collection sites.
Several of these sites no longer provide suitable habitat for Hylaeus
species because native vegetation has been removed during urbanization,
or the sites are dominated by invasive, nonnative vegetation. These
sites include Honolulu, Waikiki, ``the Honolulu mountains,'' Waialua,
and the Waianae coast (Liebherr and Polhemus 1997, pp. 345-347; Daly
and Magnacca 2003, p. 55). Between 1999 and 2002, researchers searched
coastal habitat at Makapuu and Kalaeloa (Barber's Point), but did not
find any species of Hylaeus (Magnacca, pers. comm. 2008a). The coastal
habitat at both sites is degraded and dominated by nonnative
vegetation. Descriptions of the two known sites follow:
(A) Kaena Point NAR: Between 1998 and 2008, Hylaeus anthracinus was
collected at Kaena Point, which is located on Oahu's northwest-most
point (Daly and Magnacca 2003, p. 55; Sahli, University of Hawaii at
Manoa, pers. comm. 2008). Kaena Point contains the best intact native
coastal habitat on Oahu, and is an excellent example of that type of
ecosystem in the Northwestern Hawaiian Islands. It provides habitat for
nesting seabirds, monk seals, native plants, and other native species
(Magnacca 2007a, p. 181). The primary activities within this NAR
include recreation, hiking, nature study, education, and the
observation of wildlife (DLNR 2007, p. 20). While illegal off-road
driving was once a concern, a physical barrier is now in place that
prevents vehicular access, and native vegetation is regenerating and
being restored by the Kaena Point Ecosystem Restoration Project (DLNR
2007, p. 20; Magnacca 2007a, p. 181). In partnership with several
agencies including the Service, the DLNR is building a predator-proof
fence to prevent nonnative species, such as cats and dogs that threaten
nesting seabirds, from entering 59 ac (24 ha) of coastal habitat within
Kaena Point NAR (https://www.state.hi.us/dlnr/dofaw/kaena/index.htm).
(B) Mokuauia (Goat Island): From the lack of records, it appears
Perkins and other early naturalists did not search Mokuauia or Oahu's
other offshore islets for yellow-faced bees. Recently, Hylaeus
anthracinus was found on this islet by Service biologists during
general surveys of the islet (S. Plentovich, Service, pers. comm.
2008). Mokuauia, an offshore islet in Laie Bay located on Oahu's
northeast coast, encompasses 13 ac (5.3 ha) and reaches a maximum
elevation of 15 ft (4.5 m). The entire islet is a State Seabird
Sanctuary and is managed by the State's Department of Forestry and
Wildlife (DOFAW). The entire islet was designated as critical habitat
for the endangered plant Sesbania tomentosa in 2003, and the DOFAW is
actively restoring native vegetation and controlling nonnative species.
Mokuauia is easily accessed by the public and is a popular destination
for small boats, kayaks, and swimmers on weekends.
Lanai
Hylaeus anthracinus has not been observed on Lanai for over 100
years and is likely extirpated from this privately owned island. This
species was not observed at any of the recently surveyed sites,
including Manele Bay, where it was collected by Perkins in 1899
(Magnacca 2007a, p. 182; Magnacca, pers. comm. 2008a). However, other
Hylaeus species were collected at seven of the eight locations surveyed
(Daly and Magnacca 2003, pp. 217-229).
Summary of Hylaeus anthracinus Range and Distribution
Hylaeus anthracinus was historically known from numerous coastal
and lowland dry forest habitats up to 2,000 ft (600 m) in elevation, on
the islands of Hawaii, Lanai, Maui, Molokai, and Oahu. Currently, this
species is known from a total of 13 sites in a few small patches of
coastal and lowland dry forest habitat: one location on Kahoolawe, five
locations on the island of Hawaii, two locations on Maui, three
locations on Molokai, and two locations on Oahu. In addition, in 2004 a
single individual of H. anthracinus was collected in montane dry forest
habitat on the island of Hawaii. It is unknown if this collection was a
single vagrant individual or from an established population. The lands
on which H. anthracinus occurs are under a variety of jurisdictions,
including private (e.g., TNC), State (e.g., DHHL, DOFAW, NARs, State
Park, Seabird Sanctuary), and Federal (U.S. Army, NPS).
Specific Information on Hylaeus assimulans
Taxonomy and Description
Hylaeus assimulans was first described as Nesoprosopis assimulans
(Perkins 1899, pp. 75, 101-102); 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). 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 (Magnacca
2005b, p. 2). Likewise, the nesting habits of H. assimulans are not
known, but because the species is genetically related to other ground
nesting Hylaeus spp., it is thought to be a ground nester (Magnacca
2005b, p. 2).
Hylaeus assimulans adults have been observed visiting the flowers
of Lipochaeta lobata (nehe) and its likely primary host plant, Sida
fallax (Daly and Magnacca 2003, p. 58). Hylaeus assimulans appears to
be closely associated with plants in the genus Sida, and studies thus
far suggest this yellow-faced bee species may be more common where this
plant is abundant (Daly and Magnacca 2003, pp. 58, 217; Magnacca 2007a,
p. 183). In recent survey efforts, H. assimulans seems to be more
common in dry forest at relatively higher elevations, which may be
related to the abundance of Sida in the understory (Magnacca 2005b, p.
2). Sida spp. were less often found in coastal habitat. It is likely H.
assimulans visits several other native plants, including Acacia koa,
Metrosideros polymorpha, Styphelia tameiameiae (pukiawe), and species
of Scaevola (naupaka) and Chamaesyce (akoko),
[[Page 55176]]
which are frequented by other Hylaeus species as well (Magnacca, pers.
comm. 2008b).
Range and Distribution
Historically, Hylaeus assimulans was known from numerous coastal
and lowland dry forest habitats up to 2,000 ft (610 m) in elevation on
the islands of Lanai, Maui, and Oahu. There are no collections from
Molokai, although it is likely H. assimulans also occurred there
because all other species of Hylaeus known from Maui, Lanai, and Oahu
also occurred on Molokai (Daly and Magnacca 2003, pp. 217-229). Between
1997 and 2008, surveys for Hawaiian Hylaeus were conducted in 25 sites
on Kahoolawe, Lanai, Maui, Molokai, and Oahu. Hylaeus assimulans was
absent from six of its historical localities on Lanai, Maui, and Oahu
(Xerces Society 2009b, p. 4). Hylaeus assimulans was not observed at 19
other sites with potentially suitable habitat on Lanai, Maui, Molokai,
and Oahu, including several sites from which other native Hylaeus
species have been recently collected (Daly and Magnacca 2003, pp. 56,
217; Magnacca 2005b, p. 2; Magnacca 2007a, pp. 177, 181, 183).
Currently, Hylaeus assimulans is known from a few small patches of
coastal and lowland dry forest habitat at one location on Kahoolawe,
two locations on Lanai, and two locations on Maui (Daly and Magnacca
2003, p. 58; Magnacca 2005, p. 2). This species has likely been
extirpated from Oahu because it has not been observed since Perkins'
1899 surveys and was not found during recent surveys of potentially
suitable coastal habitat at Kaena Point, Makapuu, and Kalaeloa (Daly
and Magnacca 2003, p. 217; Magnacca 2005, p. 2; H. Sahli, unpublished
data).
Kahoolawe
Although not historically known from Kahaoolawe (Daly and Magnacca
2003, Magnacca, pers. comm. 2008a), Hylaeus assimulans was discovered
in 1997 near the high cliffs of Kamohio Bay in the center of the
southern coast of the island (Daly and Magnacca 2003, p 217). The
species was absent from one other site on the island in lowland habitat
on the east coast at Pali o Kalapakea where other Hylaeus species were
collected (Daly and Magnacca 2003, pp. 217-229).
Lanai
On Lanai, Perkins found Hylaeus assimulans in low numbers within
uninhabited coastal habitat at Awalua in northwest Lanai, and in the
Koele mountains at an elevation of 2,000 ft (610 m) (Perkins 1899, p.
102). Between 1998 and 2006, seven sites with potentially suitable
habitat on private lands, including Mt. Koele and Awalua, were
surveyed, and H. assimulans was found only near Manele Road and Polihua
Road in small pockets of native vegetation (Magnacca, pers. comm.
2008b). Descriptions of these sites follow:
(A) Manele Road: In 1999, Hylaeus assimulans was collected in
lowland dry forest along Manele Road at 600 ft (180 m) in elevation,
north of Manele Beach in southern Lanai (Daly and Magnacca 2003, p.
217). Researchers observed the canopy was dominated by invasive
Prosopis pallida trees and the understory had a dense stand of Sida
fallax, the likely primary host plant of H. assimulans (Magnacca, pers.
comm. 2008b). However, with the exception of a few stunted plants at
the roadside where moisture had accumulated, the rest of the stand of
Sida fallax had senesced (reached maturity) or possibly died. Native
plants at this site appeared to be drought-intolerant and probably did
not provide consistent habitat for Hylaeus throughout the year
(Magnacca 2007a, p. 183; Magnacca, pers. comm. 2008a).
(B) Polihua Road: In 1999, two specimens of H. assimulans were
collected in lowland dry forest along Polihua Road at 1,000 ft in
elevation (300 m) in central Lanai (Daly and Magnacca 2003, p. 58).
Both sites are on private land, and we are unaware of any recent or
current land management in these areas.
Maui
Perkins collected Hylaeus assimulans from coastal habitat at the
Wailuku sand hills, and from an unknown site labeled ``Maui'' (Daly and
Magnacca 2003, p. 58). Although other rare Hylaeus spp. were collected
from the Waiehu dunes area during surveys conducted in 1999 and 2001,
H. assimulans, as well as several other species once collected there by
Perkins, were not found (Daly and Magnacca 2003, pp. 217-229; Magnacca,
pers. comm. 2008a). Between 1998 and 2006, researchers surveyed six
potentially suitable habitat locations island-wide, and H. assimulans
was found within small pockets of native plants in only two of these
sites (Daly and Magnacca 2003, p. 217; Magnacca, pers. comm. 2008a).
However, researchers believe H. assimulans may exist in potentially
suitable habitat in rugged and inaccessible portions of west Maui
(Magnacca, in litt., 2010, p. 1). Descriptions of these two sites
follow:
(A) Lahainaluna: In 1999, Hylaeus assimulans was collected in dry
lowland forest at 1,800 ft (550 m) in elevation on the west side of
Maui. The site is with the State's West Maui NAR. Established in 1986,
the NAR's management plan calls for the control and removal of feral
ungulates, and the control of selected priority invasive plant species
(https://hawaii.gov/dlnr/dofaw/nars/reserves/maui/west-maui).
(B) Waikapu: In 2000, researchers collected Hylaeus assimulans in
lowland dry shrubland dominated by the native shrub, Dodonaea viscosa
(aalii) at 400 ft (120 m) elevation in Waikapu Valley, which is south
of Iao Valley on the east side of west Maui (Daly and Magnacca 2003, p.
217). The 10,000-square ft (.09-square-ha) site is privately owned and
surrounded by a fence to exclude nonnative axis deer (Axis axis). The
fence was built in the mid-1980s by the Native Hawaiian Plant Society,
and is currently managed by inspecting the fence for breaks; removing
nonnative, invasive weeds; and collecting seeds of native plants for
propagation. There have been two major fires in the past 5 years in the
vicinity of the fenced area, although neither fire has burned within
the enclosed area (H. Oppenheimer, Plant Extinction Prevention Program,
pers. comm. 2008).
Between 1997 and 2007, Hylaeus assimulans was not collected during
surveys of potentially suitable habitat at other locations on Maui
where other rare Hylaeus species were collected, including lowland dry
forest habitat in Kanaio NAR and coastal habitat at Manawainui Gulch
(Daly and Magnacca 2003, pp. 217-229; Magnacca, pers. comm. 2008a).
Oahu
Perkins found Hylaeus assimulans to be widespread but not
relatively abundant on Oahu (Magnacca 2005b, p. 2). His Oahu collection
sites included Honolulu (Magnacca, pers. comm. 2008a), the Kaala
mountains, the Waianae Mountains, and the Waianae coast (Perkins 1899,
p. 102; Daly and Magnacca 2003, p. 58). There are also specimens
collected by Perkins from unknown locations labeled ``Oahu'' and ``w.
coast, near sea level'' (Daly and Magnacca 2003, p. 58).
Hylaeus assimulans was not found during surveys conducted between
1998 and 2008, including surveys at one historical location (Daly and
Magnacca 2003, pp. 58, 217). Although H. anthracinus was recently found
on Mokuania (see Hylaeus anthracinus Range and Distribution), H.
assimulans was not found during surveys of potentially suitable habitat
on this off-shore islet (S. Plentovich, Service, pers. comm. 2008). The
absence of H. assimulans from potentially suitable
[[Page 55177]]
coastal habitat on Oahu suggests it has likely been extirpated from
this island (Daly and Magnacca 2003, p. 58; H. Sahli, unpublished
data).
Summary of Hylaeus assimulans Range and Distribution
Hylaeus assimulans was historically known from numerous coastal and
lowland dry habitats up to 2,000 ft (610 m) in elevation, on the
islands of Lanai, Maui, and Oahu. Currently, this species is found in a
few small patches of coastal and lowland dry forest habitat in five
locations on Kahoolawe, Lanai, and Maui. The lands on which H.
assimulans occurs are under private and State (DLNR and KIRC)
ownership.
Specific Information on Hylaeus facilis
Taxonomy and Description
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 2003, 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 H. facilis. 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).
The native host plants of adult Hylaeus facilis are unknown, but it
is likely this species visits several plants other Hylaeus species are
known to frequent, including Acacia koa, Metrosideros polymorpha,
Styphelia tameiameiae, Scaevola spp., and Chamaesyce spp. (Daly and
Magnacca 2003, p. 11). Hylaeus facilis has also been observed visiting
the nonnative Tourneforia argentea for nectar and pollen (Magnacca
2007a, p. 181).
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 H. facilis was among the most common and widespread Hylaeus
species on Oahu and all of Maui Nui (Lanai, Maui, and Molokai)
(Magnacca 2007a, p. 183). 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
2007a, p. 183). Although the species was widely collected within a
diverse range of habitats historically, 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).
Hylaeus facilis has almost entirely disappeared from most of its
historical range (Daly and Magnacca 2003, p. 7; Magnacca 2007a, p.
183). Between 1998 and 2006, 39 sites on Lanai, Maui, Molokai, and Oahu
were surveyed, including 13 historical sites. Hylaeus facilis was
absent from each of the 13 historical localities (Magnacca 2007a, p.
183) and was also 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 2007a, p. 183). Likely extirpated from Lanai, H.
facilis is currently only known from two locations, one each on the
islands of Molokai and Oahu (Daly and Magnacca 2003, pp. 81-82;
Magnacca 2005c, p. 2). In addition, in 1990, a single individual was
collected on Maui in a residential area near Makawao at 1,500 ft (457
m) in elevation. However, this site is an urbanized area devoid of
native plants, and it is likely this collection was a single vagrant
individual and not from an established population on Maui.
Lanai
Perkins (1899) described Hylaeus facilis as ``common'' at two Lanai
locations. He noted H. facilis was collected from the Koele Mountains
at 2,000 ft (610 m) in elevation. Researchers believe the collection
locality was northwest of Puu Alii where the ridges are at an elevation
of approximately 2,000 ft (600 m). The Puu Alii summit itself is 2,800
ft (850 m) in elevation, and less likely to be the site of Perkins'
collection (Magnacca in litt. 2011, p. 36). Today this area contains
mixed native and nonnative vegetation. Researchers collected three
other species of Hylaeus in the same general area, along the Munro
Trail and Kaiholena ridge in 1999 and 2001 (Daly and Magnacca 2003, pp.
217-229). Perkins' second collection site was in montane habitat at
3,000 ft (900 m) in elevation at Haalelepaakai in the ``summit
mountains on Lanai'' (Daly and Magnacca 2003, p. 83). Researchers
surveyed this area in 1999 and 2001, and were unable to find H.
facilis, although they collected four other Hylaeus species (Daly and
Magnacca 2003, pp. 217-229). Hylaeus facilis is likely extirpated from
Lanai because it has not been relocated in over 100 years, and its
potentially suitable habitat has been extensively surveyed (Magnacca
2007a, pp. 177, 183).
Maui
Perkins collected Hylaeus facilis from three different sites on
Maui, including coastal habitat at the Wailuku sand hills (Waiehu
dunes), montane mesic forest habitat on Haleakala, and lowland wet
habitat in Iao Valley. Although other species of Hylaeus were collected
from the Waiehu dunes in 1999 and 2001, H. facilis, as well as several
other species collected by Perkins in the late 19th century, were
absent (Daly and Magnacca 2003, pp. 217-229).
Perkins (1899) collected Hylaeus facilis in montane mesic forest
habitat on Haleakala at an elevation of 5,000 ft (1,524 m) on
Haleakala, in the Olinda area where he is known to have camped while
surveying for and collecting insects (Evenhuis 2009, pp. 199-200).
These native forests were once abundant in this area up to 6,000 ft
(1,818 m) in elevation across the west slope of Haleakala, but have now
been completely converted by agriculture and other land uses (Juvik and
Juvik 1998, pp. 123-124). Hylaeus facilis and other species with
similar habitat requirements (e.g., H. difficilis, H. volcanicus) are
absent from the native,
[[Page 55178]]
wetter forest across the eastern slope of Haleakala (Daly and Magnacca
2003, pp. 219-221, 228-229).
Perkins also collected Hylaeus facilis in lowland wet habitat at an
elevation of 2,000 ft (610 m) in Iao Valley in the west Maui Mountains
(H. V. Daly, unpublished data). The terrain in Iao Valley is especially
rugged and wet, and Perkins relied on assistants to collect specimens
from this area (Liebherr and Polhemus 1997, p. 351). Even today the
vegetation in this area is predominantly native (Liebherr and Polhemus
1997, p. 351).
Since the late 1960s, there have been only two collections of
Hylaeus facilis on Maui, but neither is from a distinct population that
can be relocated. One collection was made in 1967 (Daly and Magnacca
2003, p. 221; Magnacca 2005c, p. 2), but the location is unknown
(Xerces Society 2009c, p. 7). In 1990, a single individual was
collected at Kokomo at an elevation of 1,500 ft (457 m) near Makawao,
in a residential area devoid of native plants (Daly and Magnacca 2003,
p. 221). This individual may have been a straggler blown in from a
different site altogether (Magnacca 2005c, p. 2). Researchers question
whether any viable H. facilis populations still remain on Maui
(Magnacca 2007a, pp. 183-184).
Molokai
Perkins collected Hylaeus facilis in three locations within montane
mesic forest habitat in the east Molokai Mountains (Daly and Magnacca
2003, p. 83). These locations were probably between Makakupaia and the
rim of Pelekunu Valley, where Perkins did most of his collecting
(Liebherr and Polhemus 1997, p. 347). Makakupaia is located within
TNC's Kamakou Preserve. Researchers have surveyed extensively in
similar, high-elevation habitat near Perkins' collecting area,
including Kamakou Road (3,200 ft (975 m)), Puu Kolekole (3,400 ft
(1,040 m)), and Kawela Gulch (3,600 ft (2,000 m)), and found other
Hylaeus species, but were unable to locate H. facilis (Daly and
Magnacca 2003, pp. 217-229).
In 2005, researchers collected Hylaeus facilis in coastal habitat
at Kuololimu Point, within Kalaupapa National Historical Park (KNHP) on
the southeast coast of the Kalaupapa peninsula (Magnacca 2007b, pp. 44-
45). This area, located on the east side of the peninsula, is largely
rocky and devoid of vegetation, but contains scattered patches of
native coastal vegetation similar to habitat at Ka Lae on the island of
Hawaii (Magnacca 2007a, p. 181). The park is cooperatively managed by
the NPS, and the State of Hawaii's DHHL, DLNR, DOH, and DOT (NPS 2006
(https://www.nps.gov/kala/index.htm)).
Oahu
Perkins collected Hylaeus facilis from six sites on Oahu (Daly and
Magnacca 2003, p. 83). One site described by Perkins was coastal
habitat in Honolulu. Although the exact location is unknown, Honolulu
coastal habitat has been completely developed for urban land use and
there is no potentially suitable coastal habitat remaining in Honolulu
for Hylaeus species. Perkins also described collecting Hylaeus species
from mountains in Honolulu, and although the exact locations are
unknown, these sites are presumed to be near known sites where he
collected, including Waiolani Ridge, Lanihuli Ridge, Nuuanu Valley, and
Konahuanui (Liebherr and Polhemus 1997, p. 348). While these mountain
areas are largely undeveloped, many are dominated by nonnative
vegetation. Researchers have surveyed potentially suitable native
habitat near Perkins' collection sites and found other species of
Hylaeus, but not H. facilis (Daly and Magnacca 2003, pp. 217-229).
Descriptions of the five remaining suitable habitats follow:
(A) Makaha Valley: Perkins (1899) collected H. facilis at an
elevation of 3,000 ft (900 m) in the upper part of Makaha Valley, on
Oahu's northwest side. There have been no surveys for Hylaeus in this
area since Perkins' collections, but researchers have observed this
area now lacks suitable Hylaeus habitat due to development,
urbanization, and conversion of native habitat to nonnative vegetation
(Magnacca, pers. comm. 2008c). Some of the upper reaches of Makaha
Valley contain patches of native vegetation, but much of the native
vegetation has been destroyed by brush fires (Liebherr and Polhemus
1997, p. 347).
(B) Mount Kaala: Perkins (1899) collected Hylaeus facilis at 2,000
ft (610 m) in elevation on Mt. Kaala, possibly within what is now Mt.
Kaala NAR. This area is a mix of dry and mesic forest communities (DLNR
1990, p. 3), and is generally characterized as predominantly native
vegetation (Liebherr and Polhemus 1997, p. 348). This area has not been
extensively resurveyed for Hylaeus spp. because much of it is either
inaccessible (due to either private or U.S. Army ownership), or too
rugged in general, requiring a long and steep approach along the Dupont
Trail on the north slope of Mt. Kaala.
(C) Waianae Mountains: Perkins (1899) collected Hylaeus facilis in
the Waianae Mountains, ``upland from Waianae'', likely in dry lowland
forest, although the exact location is unknown. In 2008, researchers
surveyed potentially suitable habitat in the Waianae-Kaala Forest
Reserve (FR), but did not find H. facilis (Magnacca, pers. comm. July
2008c).
(D) Tantalus: Perkins collected Hylaeus facilis in lowland mesic
habitat on ``Tantalus'' (Liebherr and Polhemus 1997, p. 348), which
today is in close proximity to the urban core of Honolulu. This area is
a mix of residential development and undeveloped sites dominated by
nonnative plants, including various species of Phyllostachys spp.
(bamboo), Acacia confusa (Formosa koa), Eucalyptus robusta (swamp
mahogany), and Aleurites moluccana (kukui) (USDA 2001 https://soilseries.sc.egov.usda.gov/OSD_Docs/T/TANTALUS.html). Habitat
dominated by nonnative plants does not support viable populations of
Hylaeus, and no species have been reported from this area since
Perkins' collections despite more recent surveys in the few small,
widely separated areas containing native plant hab