Endangered and Threatened Wildlife and Plants; 12-Month Finding on Five Petitions To List Seven Species of Hawaiian Yellow-faced Bees as Endangered

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DEPARTMENT OF THE INTERIOR Fish and Wildlife Service 50 CFR Part 17 [Docket No. FWS-R1-ES-2010-0012; MO 92210-0-008] 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:

nual 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 http://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

her 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

une 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

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 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

ixed transect route and recording the insects observed.

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

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

ed 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

ing 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 Lowland dry Lowland mesic Lowland wet Montane mesic Montane dry H. anthracinus, 13 populations HI, MA, MO, OA HI, KAH, (*LA), MA, (*MO), (*OA) N/A N/A HI. 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), (*OA) (*MA), OA (*MO) N/A. 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. 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)

s 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

s, 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).

i, 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

006g). 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 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)

ai (Daly and Magnacca 2003, p. 53).

Life History

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)

rveys 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

nacca 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 ( http://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 ( http://www.kohanaiki.org/ ))

7b, p. 44). Kohanaiki is an area of land granted to indigenous Hawaiians in 1995 for cultural and recreational preservation and pursuits (Kohanaiki Ohana 1995 ( http://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 ( http://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 http://www.hawaiistateparks.org/hawaiistateparks/parks/hawaii/index.cfm?park_id=47 ).

/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)

ted 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

, 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).

ated 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 ( http://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:

r 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

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)

rkins-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 ( http://www.state.hi.us/dlnr/dofaw/kaena/index.htm )

he 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 ( http://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

acca, 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)

o 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), Hylaeus species as well (Magnacca, pers. comm. 2008b).

Range and Distribution

tameiameiae (pukiawe), and species of Scaevola (naupaka) and Chamaesyce (akoko), 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

at 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

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 ( http://hawaii.gov/dlnr/dofaw/nars/reserves/maui/west-maui )

hainaluna: 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 ( http://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

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

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)

us 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)

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

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)

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,

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)

e.g., H. difficilis, H. volcanicus ) are absent from the native,

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)

mus 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 ( http://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)

llected, 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)

g the Dupont Trail on the north slope of Mt. Kaala.

(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

nd 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— http://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)

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 (Meade-Waldo 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

) 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.

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 group, no cleptoparasites are known from the family Colletidae (Daly and Magnacca 2003, p. 9).

to become cleptoparasites. Among the world's bees, other than the Hawaiian Hylaeus group, no cleptoparasites are known from the family Colletidae (Daly and Magnacca 2003, p. 9).

The larvae of 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)

cted 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 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

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).

Molokai

Although Hylaeus hilaris was never collected on Molokai by Perkins, in 1918, Fullaway (1918, p. 396) collected the species at an unspecified site

ese 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).

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

Specific Information on Hylaeus kuakea

Taxonomy and Description

Hylaeus kuakea was first described by Daly and Magnacca (2003, pp

ed 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.

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 well-defined Hylaeus species groups. Its facial marks are similar to those of the H. difficilis group and to H. anthracinus, but it can be distinguished by its unusual ivory facial marking covering the clypeus (the lower face region). Hylaeus kuakea also resembles H. anthracinus, but has a denser, more distinct arrangement of setae (sensory hairs) on the head and generally narrower marks next to the compound eyes (Daly and Magnacca 2003, p. 125; Magnacca 2005e, p. 2). Only 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)

s believed to be related to other wood-nesting Hawaiian Hylaeus species (Magnacca and Danforth 2006, p. 403).

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 forest-inhabiting 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 lowland mesic forest areas due their remote and rugged locations, small size, and distant spacing among large areas of nonnative forest

agnacca, 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 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 (Meade-Waldo 1923, p. 1). Daly and Magnacca (2003, pp. 133-134) most recently described the species as H. longiceps. Hylaeus longiceps is a small to medium-sized, black bee with clear to slightly smoky-colored wings. Its distinguishing characteristics are its long head and the facial marks of the male. The lower face of the male is marked with a yellow band that extends at the sides of the face in a broad stripe above the antennal sockets. The area above the clypeus (lower face region) is very long and narrow, and the scape (the first antennal segment) is noticeably twice as long as it is wide. The female is entirely black and unmarked (Daly and Magnacca 2003, p. 133).

Life History

The diet of the larval stage of Hylaeus longiceps is unknown, although the larvae are presumed to feed on stores of pollen and nectar collected and deposited in the nest by the female adult (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)

133).

Life History

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) 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).

us 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 1999, researchers collected H

pwreck Beach. Descriptions of these three locations follow:

(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)

nted.

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 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)

ix 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).

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

, 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).

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).

t 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).

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 koa -dominated, lowland mesic forest on Oahu (Daly and Magnacca 2003, p. 138). This type of forest is increasingly rare and patchily distributed on Oahu (Smith 1985, pp. 227-233; Juvik and Juvik 1998, p. 124; Wagner et al. 1999, pp. 66-67, 75).

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— http://hawaiitrails.ehawaii.gov/trail.php?TrailID=OA+09+008).

Summary of Hylaeus mana Range and Distribution

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;

(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.

e 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. 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 yellow-faced bees for foraging of nectar and pollen, any further loss of this habitat may endanger their long-term chances for conservation and recovery

n 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 yellow-faced 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 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)

-building, and past agricultural activities (Cuddihy and Stone 1990, pp. 94-95). The native 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

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 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

i, 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 lowland mesic forest habitat on the island of Oahu. Because we lack information on the historical range of H. kuakea and H

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 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

ry 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 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)

e 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 soil-water 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 1990, p. 74; D'Antonio and Vitousek 1992, p. 73; Vitousek et al. 1997, p. 6)

ive 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 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

eclining 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 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

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

a 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 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

agnacca 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 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)

s deer are found on Lanai, Maui, Molokai, and Oahu (HCA 2007). At least 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 State-designated 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.

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 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, human-exacerbated 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

a relatively new, human-exacerbated 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 grass-fueled 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, H

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, 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)

as 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 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 yellow-faced 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

aining Area. The area that burned in this fire is near the Kaena Point NAR, where two of the yellow-faced 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 yellow-faced bees are known from military lands, at least one species, H

e, 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 yellow-faced 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 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 ( http://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

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

uman 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

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)

ne 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-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

ssion 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 yellow-faced 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)

e threats to the Hawaiian yellow-faced 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 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 Niño (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 Niño have yet to be made (Benning et al. 2002, pp. 14,248-14,249)

n precipitation caused by global climate change. However, future changes in precipitation are uncertain because they depend in part on how El Niño (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 Niño 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 Niño-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 Niño-La Niña (a different disruptive extreme weather and climate pattern that can alternate with El Niño) 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

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.

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