Pollinator Habitat Conservation Along Roadways, Volume 6: Hawaii (2023)

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3-1 Chapter 3 Imperiled Pollinator Profiles Hawaii is home to a unique assemblage of pollinators. While there are almost 1,000 species of moths in Hawaii, there are only two species of butterflies. There are also 63 species of yellow-faced bees in Hawaii, as well as several species of bees that have been introduced from other places. Hawaii is home to eight species of pollinators listed under the federal Endangered Species Act (ESA) as of March 2022. This includes seven species of yellow- faced bees and one moth. However, there are other declining pollinators in this state that are not protected by the ESA. This section provides profiles of 33 species of imperiled pollinators found in Hawaii (Table 3-1). In addition to ESA-listed species, pollinators that have the potential to be listed in the future based on NatureServe rankings and expert opinion are included. These include 22 imperiled yellow-faced bees, Hawaii’s two native butterfly species, and one additional moth. This is not an exhaustive list of all declining pollinators in the region; the focus is on species in need of conservation that have a broad distribution and those that are more likely to affect departments of transportation (DOTs). An overview of the basic biology of bees, butterflies, and other insect pollinators can be found in Chapter 2. This section presents information on life history, distribution, threats, and habitat requirements for imperiled pollinators in this region. Information on known adult flight times (i.e., the breeding period) and larval active times (for butterflies and moths; larval bees live within nests) are included. The profiles also include a list of important plants that are used by each species as host plants or for pollen and/or nectar. Some of these plants are nonnative species or noxious weeds. These species are included in the profiles, as the information may be useful, but using them in revegetation efforts is not recommended (see Chapter 7 for more). Some basic conservation recommendations and information on the effects of roadside management on each species are also provided, when such information is available. Insects generally tend to be less well studied than vertebrates; therefore, very little is known about the biology and habitat requirements of some of these species. Other pollinator species may be better studied, but rigorous studies of the effects of different management practices on the species or their habitat may still be lacking. The best possible recommendations are made based on the available information. The profiles provide information on life cycles, host plants, habitat needs, and adult flight times that can be used to tailor maintenance and revegetation decisions (Chapters 6 and 7). General active times for adults and larvae are also provided. It should be noted that active times may shift regionally, such as with elevation or latitude, or from year to year with changes in climate. However, consultation with local experts, as well as biologists from state and federal agencies, is recommended to help DOTs develop meaningful management plans for species of interest in their areas, because the profiles cannot capture the site- specific nuances that should be considered. As the profiles indicate, habitat loss is a primary driver of species’ declines. Roads can be a source of habitat fragmentation (Box 5-3). However, with investments in high-quality revegetation (revegetation that prioritizes usage of native plants and high plant diversity, including an abundance of flowering plants that provide pollen and nectar or act as host plants for butterflies) where appropriate, roadsides can also provide an ideal opportunity

Chapter 3. Imperiled Pollinator Profiles  3-2 to increase habitat connectivity for many species, including pollinators. It is important to note that although some of the species profiled in this section may be unlikely to use roadside habitat for breeding, individuals may still use roadside habitat for nectar or pollen or as movement corridors. Therefore, it is worthwhile to invest in high-quality habitat restoration projects along roadsides near natural areas, preserves, and populations of imperiled species. Another common threat to many imperiled butterflies in the United States is noxious and invasive plants that displace required host plants. Here again, investing in high-quality habitat restoration using native plants can benefit imperiled pollinators, even if those pollinators are not using roadside habitat, by helping to slow the spread of invasive plants into key habitat areas. This section includes profiles of pollinator species that may not occur in roadside habitat, but may be found in other DOT land holdings, such as mitigation areas. Table 3-1. List of profiled imperiled pollinator species in the Hawaii region. Scientific Name  Common Name  Status#  ESA Listed Pollinator Species  Hylaeus anthracinus  Anthricinan yellow‐faced bee  Endangered   Hylaeus assimulans  Assimulans yellow‐faced bee  Endangered  Hylaeus facilis  Easy yellow‐faced bee  Endangered  Hylaeus hilaris  Hilaris yellow‐faced bee  Endangered  Hylaeus kuakea  Hawaiian yellow‐faced bee  Endangered  Hylaeus longiceps  Longhead Hawaiian yellow‐faced bee  Endangered  Hylaeus mana  Hawaiian yellow‐faced bee  Endangered  Manduca blackburni  Blackburn’s sphinx moth  Endangered  Declining Pollinator Species  Hylaeus akoko  Hawaiian yellow‐faced bee  GNR  Hylaeus anomalus  Anomalous yellow‐faced bee  G2  Hylaeus dimidiatus  Dimidiatan yellow‐faced bee  GNR  Hylaeus finitimus  Finitiman yellow‐faced bee  GH  Hylaeus flavifrons  Hulan yellow‐faced bee  G1G3  Hylaeus fuscipennis  Darkwing Yellow‐faced Bee  G1G3  Hylaeus gliddenae  Hawaiian yellow‐faced bee  GNR  Hylaeus hula  Hulan yellow‐faced bee  GNR  Hylaeus kona  A yellow‐faced bee  GNR  Hylaeus mauiensis  Maui yellow‐faced bee  GH  Hylaeus melanothrix  Melanothrix yellow‐faced bee  GH  Hylaeus nalo  A yellow‐faced bee  GNR 

Chapter 3. Imperiled Pollinator Profiles 3-3 Scientific Name  Common Name  Status#  Hylaeus niloticus  Obscuratan yellow‐faced bee  GH  Hylaeus ombrias  Ombrias yellow‐faced bee  GNR  Hylaeus paradoxicus  Hawaiian yellow‐faced bee  GNR  Hylaeus perkinsianus  Perkin’s yellow‐faced bee  G1G3  Hylaeus perspicuus  Perspicuan yellow‐faced bee  GH  Hylaeus psammobius  Psammobian yellow‐faced bee  GNR  Hylaeus satelles  Satellus yellow‐faced bee  GH  Hylaeus simplex  Simple yellow‐faced bee  GH  Hylaeus solaris  A yellow‐faced bee  GNR  Hylaeus volatilis  Volatile yellow‐faced bee  G1G3  Tinostoma smaragditis  Fabulous Green Sphinx of Kauai  G1G2  Udara blackburni  Blackburn’s bluet  G4  Vanessa tameamea  Kamehameha butterfly  G3G4  # Status of pollinators is either the ESA status for listed species or is taken from NatureServe (accessed  March 2022) for species not listed under the ESA.    G1: Critically Imperiled. At very high risk of extinction due to extreme rarity (often five or fewer  populations), very steep declines, or other factors.   G2: Imperiled. At high risk of extinction due to very restricted range, very few populations (often 20 or  fewer), steep declines, or other factors.   G3: Vulnerable. At moderate risk of extinction due to a restricted range, relatively few populations  (often 80 or fewer), recent and widespread declines, or other factors.   G4: Apparently Secure. Uncommon but not rare; some cause for long‐term concern due to declines or  other factors.   G5: Secure. Common; widespread and abundant.   G#G#: Range Rank. A numeric range rank (e.g., G2G3) is used to indicate the range of uncertainty in  the status of a species or community.   GNR: No status rank.   GH: Possibly extinct.   T#: Infraspecific Taxon (for subspecies or varieties). The status of infraspecific taxa is indicated by a  “T‐rank” following the species’ global rank. For example, the global rank of a critically imperiled  subspecies of an otherwise widespread and common species would be G5T1.  

Chapter 3. Imperiled Pollinator Profiles  3-4 3.1 Federally Listed as Endangered Hawaiian Yellow‐ Faced Bees: Anthricinan yellow‐faced bee (Hylaeus  anthracinus), Assimulans yellow‐faced bee (H.  assimulans), Easy yellow‐faced bee (H. facilis), Hilaris  yellow‐faced bee (H. hilaris), Hawaiian yellow‐faced  bee (H. kuakea), Long‐head Hawaiian yellow‐faced bee  (H. longiceps), Hawaiian yellow‐faced bee (H. mana)      Jan  Feb  Mar  April  May  June  July  Aug  Sept  Oct  Nov  Dec  Adult                                                                                                Adult yellow‐faced bee (top left and right). Adult flight times (i.e., breeding period) are shown in the chart  (bottom). Photo credit: John@lahainaphotogrpahy.      Order: Hymenoptera Family: Colletidae Distribution: Hawaii Where Hawaiian yellow-faced bees occur: Hawaii is home to 63 endemic yellow-faced bee species—the only bees native to the islands. Hawaiian yellow-faced bees are known to exploit nearly the entire range of rainfall and elevation on the main islands and occur across all habitat types; this makes them unique compared to nearly all other native insect groups found throughout the archipelago. See the following sections for specific habitat associations for listed and unlisted species. How to recognize: Yellow-faced bees in the genus Hylaeus are a group of small to moderately sized, generally slender bees with sparse, short hairs on the body. Bees in the genus are commonly known as yellow-faced bees for their facial markings, which are often yellow to white. Typically, it is the males of the group who possess distinguishing facial markings, which can often aid in species identification; most females are largely all black with fewer structural and color characters, making them more difficult to identify. Many females require examination in the hand and associated males can help with identification. In shape they resemble wasps, and their lack of pollen-carrying structures gives them a hairless appearance. However, both males and females do possess hairs. On closer inspection, they can be distinguished from wasps by their plumose (branched) hairs, which are unbranched in wasps.

Chapter 3. Imperiled Pollinator Profiles 3-5 Life cycle: Adult Hawaiian yellow-faced bees can be observed year-round. They are solitary nesters and, depending on the species, may nest in cavities either above or below ground; however, specific nesting preferences are unknown for most species. As solitary bees, reproductive females will provision their own nests and construct brood cells (chambers where the young bees develop) in a nesting cavity or tunnel. One of their common names (cellophane bees) refers to the cellophane-like material the bees use to line their nests and make them waterproof. An egg is laid within each cellophane-lined chamber and supplied with pollen and nectar. Yellow-faced bees lack external pollen-carrying structures and instead carry a pollen-nectar mixture to the nest internally within their crop. Female bees establish nests in pre-existing cavities in plant material such as hollow stems and branches, in holes created by wood-boring insects, under bark or rocks, or in burrows in the soil. Some species of Hawaiian yellow-faced bees may also utilize human-made structures that provide suitable nest cavities, such as coral retaining walls in coastal habitats, and artificial nests with pre-drilled holes. Cuckoo bee life cycle: The Hawaiian Islands are home to the only cuckoo bees in the family Colletidae. Cuckoo yellow-faced bees are parasites of other yellow-faced bee species. Five species of Hawaiian yellow-faced bees are cuckoo bees or cleptoparasites: rather than build their own nest or provision for their young, a cuckoo bee will lay its eggs in the nests of other yellow-faced bee species. Cuckoo bee larvae hatch and feed on the provisioned food within the host nest and may directly kill the host egg or larva or cause it to starve by consuming the food stores. Habitat needs: Availability of food and nesting resources is a key feature in determining the success of yellow-faced bees. All Hawaiian Hylaeus species are highly dependent on intact native plant communities; they are rarely observed utilizing nonnative plants and are tied to habitats with a diversity of native forage plants. As the only native Hawaiian bees, they are likely important pollinators of native plants from the coastal strand to high- elevation forests. Hawaiian yellow-faced bees may nest in vegetation or within the ground. Stem- or cavity-nesting species need vegetation (woody, hollow, or pithy-stemmed plants) or other materials with pre-existing cavities (e.g., coral rubble) to provide nest sites. Ground-nesting species need undisturbed substrate and generally require relatively dry conditions. Adults are active year-round; therefore, it is important to maintain a bloom of floral resources throughout the year to supply yellow-faced bees with a diversity and abundance of food. Top reasons for decline, if known: The main threats known to yellow-faced bees are habitat loss and destruction, and the introduction of invasive plants and animals. Many formerly abundant and widespread species are now extremely rare and restricted to small populations due to many other factors including pesticide exposure, climate change (for coastal species: sea level rise; increased storm frequency), pathogens, and parasites, as well as the invasion of nonnative species. Nonnative predators such as ants prey directly on the immobile immature life stages. Feral ungulates and other introduced mammals destroy, degrade, and alter habitat, interfering with the reproduction of native plants. Additionally, introduced ants and bees are thought to be significant competitors for floral resources. Conservation recommendations: Conservation efforts for yellow-faced bees will help address known threats, including nonnative predators, invasive plants, and habitat

Chapter 3. Imperiled Pollinator Profiles  3-6 destruction from ungulates in lowland and higher-elevation habitats. The main conservation and recovery efforts for Hawaiian yellow-faced bees include planting native floral species and nest plants utilized by stem-nesting species, creating new or managing existing habitat for all life history needs, protecting habitat from destruction by nonnative ungulates, and providing connectivity among suitable habitat to promote the long-term persistence of current populations. Woody, hollow, or pithy-stemmed vegetation and ground substrate can be left undisturbed when possible to provide nesting opportunities. Artificial nests (e.g., wood blocks with pre-drilled holes) that offer protection from invasive crawling predators can be provided to augment nesting resources for cavity nesters. Additionally, for rare and lesser-known species, there is a continued need to survey for extant populations and document the existence of each species. Potential impacts of roadside management: Intensive mowing and invasive species management may deplete floral resources and potentially disturb roadside nests. Above- ground cavity- and tunnel-nesting bees and ground-nesting species may be sensitive year- round to management that could disturb nests (e.g., ground excavation, mowing) or harm bees in nests (e.g., chemical applications). Indiscriminate spraying of herbicides on roadsides can also affect the health of yellow-faced bees by removing floral resources. Native seed mixes in planned revegetation along roadsides will be more effective in supporting yellow-faced bees than nonnative seed mixes. Because these bees are active year-round, revegetation projects intended to support these pollinators should ensure that at least three species will be in bloom at all times during the year. 3.1.1 Anthricinan Yellow-Faced Bee (Hylaeus anthracinus) Status: Endangered How to recognize: H. anthracinus is a medium-sized bee with clear to smoky wings, a black body, and dark brown to black legs that are unmarked except for a pale area on the tibia of the first pair of legs. Males of this species have yellow on the face that is shaped like an oval, which encompasses most of the lower part of the face; yellow facial markings do not extend above the base of the antennae. Females of the species are entirely black and lack facial markings. Where it occurs: Usually inhabits dry coastal strand habitat; may rarely occur in dry lowland forest. Populations on rocky shorelines appear to be restricted to an extremely narrow corridor of vegetation composed predominantly of native and naturalized trees and shrubs between the sandy beaches and inland vegetation. This species has been recorded on Molokai, Oahu, Kahoolawe, Maui, and Hawaii, but may exist on other islands as well. Habitat needs: Adults of this species have been observed year-round and require a diversity of plants that flower throughout the year (Table 3-2). Of the Hawaiian yellow-faced bees, nest preferences are best known for female H. anthracinus, which are found to nest in hollow stems of coastal plants. Nest plants include native naupaka kahakai (S. taccada) and naturalized tree Island‐level distribution for H. anthracinus 

Chapter 3. Imperiled Pollinator Profiles 3-7 heliotrope (T. argentea); this species is also known to use pre-existing cavities in coral rubble and artificial nest blocks for nesting. H. anthracinus is thought to be a host of the cleptoparasitic yellow-faced bee H. hilaris. This species visits several native coastal shrubs and herbaceous food plants for nectar and pollen (Table 3-2). Table 3-2. Plants used by Hylaeus anthracinus. Species Name  Common Name  Notes  Nest Plants  Scaevola taccada  Beach naupaka  Native  Tournefortia argentea  Tree heliotrope  Nonnative; naturalized  Nectar Plants  Argemone glauca  Kala/smooth pricklypoppy  Native  Capparis sandwichiana  Maiapilo/native caper  Native  Chamaesyce celastroides  Akoko/’ekoko  Native  Chamaesyce degeneri  Akoko/beach sandmat  Native  Scaevola coriacea  Naupaka/dwarf naupaka  Native; endangered  Scaevola taccada  Naupaka kahakai/beach naupaka  Native  Myoporum sandwicense  Naio  Native  Sesbania tomentosa  Ohai/Oahu riverhemp  Native; endangered  Sida fallax  Ilima/yellow ‘ilima  Native  3.1.2 Assimulans Yellow-Faced Bee (Hylaeus assimulans) Status: Endangered How to recognize: This is a medium-large bee with slightly smoky to smoky wings. It is distinguished from other coastal yellow-faced bees by its relatively large size. The male is black with yellow facial markings. The male’s face can be entirely yellow below the antennae, or the yellow markings may be interrupted by a narrow to broad band of black on the clypeus, forming three separate facial spots; facial markings extend above the base of the antennae. Females are dark in color and are largely unmarked, except for the fore tibia, which is marked by a pale area.

Chapter 3. Imperiled Pollinator Profiles  3-8 Where it occurs: Dry coastal strand and lowland dry shrublands and forest habitat. This species has been recorded in Kahoolawe, Lanai, Maui, and Oahu. Habitat needs: Nesting preferences are unknown. H. assimulans is thought to be a host of the cleptoparasitic yellow-faced bees H. hilaris and N. volatilis. Known food plants are native dryland shrubs and herbaceous plants, including ilima (Sida fallax) and shrubland nehe (Lipochaeta lobata) (Table 3-3). Table 3-3. Plants used by Hylaeus assimulans. Species Name  Common Name  Nectar Plants  Lipochaeta lobata  Nehe/shrubland nehe1  Lycium sandwicense  Ohelo kai/Hawai’i desert‐thorn1  Osteomeles sp.  Ulei/osteomeles1  Scaevola sp.  Naupaka1  Sida fallax  Ilima/yellow ‘ilima1  1 Native  3.1.3 Easy Yellow-Faced Bee (Hylaeus facilis) Status: Endangered How to recognize: H. facilis is a medium-sized bee with smoky wings. Males have an oval- shaped, yellow facial marking that does not extend above the base of the antennae. Males may be distinguished from H. simplex, its sister species, by the large gonoforceps (part of the external genitalia) that are visible externally. Females are very similar to and difficult to reliably distinguish in the field from the common and widespread H. difficilis and the rare H. simplex unless there is an accompanying male. Where it occurs: Dry coastal, and dry to mesic shrubland and forest. Historically one of the most common dryland species, found in Oahu, Molokai, Lanai, and Maui. However, it is now extremely rare, and only three individuals have been recorded from Oahu and Maui over the last 50 years. Habitat needs: Historic records document this species nesting within dead twigs and plant stems hollowed by burrowing beetles; however, more recent Island‐level distribution for H. assimulans  Island‐level distribution for H. facilis 

Chapter 3. Imperiled Pollinator Profiles 3-9 monitoring surveys have not been able to confirm these observations. This species is also thought to be a host of the cleptoparasite H. volatilis. Native host plant visitation has not been documented for this species; however, one floral record exists for this species on the naturalized tree heliotrope (Tournefortia argentea). Other yellow-faced bees in similar habitat often utilize community-dominant trees, shrubs, and sub-shrubs, including koa (Acacia koa), lehua (Metrosideros polymorpha), naupaka (Scaevola spp.), and akoko (Chamaesyce spp.), the latter of which have representatives that are themselves listed as endangered under the ESA.   Table 3-4. Plants used by Hylaeus facilis. Species Name  Common Name  Nest Plants  Broussaisia arguta  Kanawao/Hawaiian hydrangea1  Clermontia grandiflora  Oha/bog clermontia1  Smilax sandwicensis  Akaawa/Hawai’i greenbrier1  Nectar Plants  Tournefortia argentea  Tree heliotrope2  1 Native  2 Nonnative; naturalized  3.1.4 Hilaris Yellow-Faced Bee (Hylaeus hilaris) Status: Endangered How to recognize: H. hilaris is one of the most colorful of all Hawaiian yellow-faced bees. Males are distinguished by the yellow facial marks that cover most of the face and the reddish-brown abdomen with unusual bands of white hairs. While most female Hawaiian yellow-faced bees lack distinct facial marks, females of this species possess some coloration on the face that is yellowish to reddish brown. The female abdomen is dark with light hair bands on the segments and the forelegs are yellowish brown. The reddish abdomen, light- colored abdominal hair bands, and large facial marking help distinguish this bee from other yellow-faced bees. Where it occurs: Restricted to coastal strand habitat. It was previously known to occur in Lanai and at the Wailuku Sand Dunes in Maui. The Maui site was lost to habitat loss and degradation. In recent years, this species has only been recorded on Molokai Island.

Chapter 3. Imperiled Pollinator Profiles  3-10 Habitat needs: H. hilaris is one of five species of cleptoparasitic Hawaiian yellow-faced bees, which lay their eggs in the nests of other yellow-faced bee species. Host bees that co-occur in coastal strand habitat are thought to include the endangered H. anthracinus, H. assimulans, and H. longiceps. Nectar plant visitation is unknown for this species; however, other yellow-faced bees in similar habitat often utilize native coastal vegetation, such as akoko (Chamaesyce spp.), naupaka (Scaevola spp.), naio (Myoporum sandwicense), and ilima (Sida spp.). 3.1.5 Hawaiian Yellow-Faced Bee (Hylaeus kuakea) Status: Endangered How to recognize: H. kuakea is a small bee with slightly smoky wings. Males are unusual in their ivory facial marking on the clypeus, which extends narrowly to the adjacent paraocular area next to the eyes. It is similar in appearance to H. anthracinus but differentiated by the facial mark and longer hairs on the head. The female has not been described. Where it occurs: H. kuakea occurs in lowland mesic forests that occur on the leeward slopes. This species likely has narrow habitat requirements and may strictly depend on these rare habitats, which serve as transitional zones between dry and wet environments. It is currently known only from the Waianae Mountains in Oahu. Habitat needs: This species is known only from sporadic records but was more recently documented utilizing the food plant Koolau Range sandmat (Chamaesyce rockii). It likely visits additional native plants in mesic forests, which other yellow-faced bees in similar habitat are known to frequently visit, including akoko (Chamaesyce olowaluana) and naupaka (Scaevola spp.). Nest substrate is unknown for this species, but it may nest in wood similar to other related yellow-faced bees. This species is thought to be a host of the cleptoparasitic yellow-faced bee H. hilaris. Island‐level distribution for H. hilaris  Island‐level distribution for H. kuakea 

Chapter 3. Imperiled Pollinator Profiles 3-11 3.1.6 Longhead Hawaiian Yellow-Faced Bee (Hylaeus longiceps) Status: Endangered How to recognize: H. longiceps is a medium-sized dark bee with clear to slightly smoky wings. They are distinguished by their long head and the facial markings on the male. The male’s face is entirely yellow below the antennal sockets. The markings extend along the sides of the face to form broad stripes above the scape (the last antennal segment at the base), which is twice as long as it is wide. The female is entirely black and usually unmarked (some may have a small yellow spot on the legs, near the front of the tibia), but has distinct punctuation (round pits) on the front of the head. The female may be distinguished from co-occurring H. anthracinus and H. flavipes by its erect dark hairs on abdominal segment 6. Where it occurs: This species was once common in dry coastal and dry lowland habitat. Remaining populations are restricted to a few populations in small remnant patches of coastal strand and lowland habitat. It is known from Oahu, Molokai, Lanai, and Maui islands. Habitat needs: Nesting requirements are unknown for this species, but it may nest in the ground similar to other related ground-nesting species. H. longiceps visits a variety of native shrubs and trees for nectar and pollen (Table 3-5). Table 3-5. Plants used by Hylaeus longiceps. Species Name  Common Name  Notes  Nectar Plants  Chamaesyce degeneri  akoko/beach sandmat  Native  Lipochaeta lobata  nehe/shrubland nehe  Native  Myoporum sandwicense  naio  Native  Santalum ellipticum  iliahialoe  Native  Scaevola coriacea  naupaka/dwarf naupaka  Native; endangered  Scaevola sericea  naupaka kahakai/beach naupaka  Native  Sesbania tomentosa  ohai/Oahu riverhemp  Native; endangered  Sida fallax  ilima/yellow ‘ilima  Native  Tournefortia argentea  tree heliotrope  Nonnative; naturalized  Vitex rotundifolia  pohinahina/roundleaf chastetree  Native  Island‐level distribution for H. longiceps 

Chapter 3. Imperiled Pollinator Profiles  3-12 3.1.7 Hawaiian Yellow-Faced Bee (Hylaeus mana) Status: Endangered How to recognize: H. mana is the smallest of all Hawaiian yellow-faced bees. The male’s face is mostly yellow below the base of the antennae, and the yellow extends dorsally in a narrowing stripe. There is a shallow groove underneath the scape. The legs and pronotum are extensively marked with yellow. The female has three yellow lines on the face, one near each eye as well as a transverse stripe at the apex of the clypeus; the other marks are similar to those of the male. It can be distinguished from related species by its extremely small size; the male’s narrow, strongly arched process of the eighth sternum; and the female’s extensive facial marks and transverse rather than longitudinal mark o the clypeus. Where it occurs: H. mana occurs in lowland mesic forests that serve as transitional zones between dry and wet environments. Records suggest that this species likely has limited habitat requirements and is restricted to a narrow zone on several ridges across Ko’olau range on Oʻahu. Habitat requirements: H. mana is known only from sporadic records but has primarily been documented utilizing the food plant iliahi or sandalwood (Santalum freycinetianum). This bee may be a specialist on Santalum; however, additional potential mesic forest food plants, which are visited by other yellow-faced bees, include akoko (Chamaesyce olowaluana), lehua (Metrosideros polymorpha), Styphelia tameiameiae, and naupaka (Scaevola spp.). Nest substrate is unknown for this species, but it may nest in wood similar to other related yellow-faced bees. 3.2 Other Yellow‐Faced Bees: Hawaiian yellow‐faced  bee (Hylaeus akoko), Anomalous yellow‐faced bee (H.  anomalus), Dimidiatan yellow‐faced bee (H.  dimidiatus), Finitiman yellow‐faced bee (H. finitimus),  Very yellow‐faced bee (H. flavifrons), Darkwing yellow‐ faced Bee (H. fuscipennis), Hawaiian yellow‐faced bee  (H. gliddenae), Hulan yellow‐faced bee (H. hula), Kona  yellow‐faced bee (H. kona), Maui yellow‐faced bee (H.  mauiensis), Melanothrix yellow‐faced bee (H.  melanothrix), a yellow‐faced bee (H. nalo), Obscuratan  yellow‐faced bee (H. niloticus), Ombrias yellow‐faced  bee (H. ombrias), Hawaiian yellow‐faced bee (H.  paradoxicus), Perkin’s yellow‐faced bee (H.  perkinsianus), Perspicuan yellow‐faced bee (H.  perspicuus), Psammobian yellow‐faced bee (H.  psammobius), Satellus yellow‐faced bee (H. satelles),  Island‐level distribution for H. mana 

Chapter 3. Imperiled Pollinator Profiles 3-13 Simple yellow‐faced bee (H. simplex), a yellow‐faced  bee (H. solaris), Volatile yellow‐faced bee (H. volatilis)  While seven species of Hawaiian yellow-faced bees profiled above are listed as endangered under the ESA, many others receive no federal protections despite their apparent rarity and imperilment. Twenty-two additional species of imperiled yellow-faced bees are highlighted separately here. Far less is known about their status and life history; however, they generally have small, isolated populations that are vulnerable to many of the same threats facing endangered yellow-faced bees and would benefit from conservation actions. Several species treated here were listed as a group in 1984 and designated as “candidate endangered” species (Category 2; now considered “species of concern”) by the U.S. Fish and Wildlife Service. This group of yellow-faced bees was thought to be endangered, but not enough information existed to justify a formal endangered listing. Several additional species included here were listed as “probably extinct” (Category 3A). In 2007, conservation categories were assigned to all yellow-faced bees described at the time by Hawaiian entomologist K. Magnacca to understand their current status based on more recent collections (see Table 3-6). While some of these species once thought to be extinct are now known to be extant, others have not been recovered; however, more surveys are needed to locate extant populations in suitable habitat and establish the extent of their distribution and abundance. When known, floral resources and habitat information are included for each species, but nesting preferences are largely unknown. Conservation recommendations and impacts from roadside management will be similar to those for listed species.

3-14 Table 3-6. Other Hawaiian Hylaeus spp. without formal federal conservation status. Scientific Name  Common Name  Habitat  Floral Resources  Conservation Category*  Global  Conservation  Status**  Hylaeus akoko  Hawaiian yellow‐ faced bee  Mesic forest  Chamaesyce  olowaluana  5  GNR  Hylaeus  anomalus  Anomalous  yellow‐faced bee  Mesic and wet  forest  Acacia koa, Psychotria  sp.  6‡  G2  Hylaeus  dimidiatus#  Dimidiatan  yellow‐faced bee  Dry forest  Chamaesyce  olowaluana  4  GNR  Hylaeus  finitimus#  Finitiman yellow‐ faced bee  Coastal  unknown  6  GH  Hylaeus  flavifrons#  Very yellow‐faced  bee  Coastal  Portulaca oleracea,  Scaevola sericea, Sida  fallax  4  G2  Hylaeus  fuscipennis#  Darkwing yellow‐ faced bee  Wet and mesic  forest  Acacia confusa  3  G2  Hylaeus  gliddenae  Hawaiian yellow‐ faced bee  Mesic forest  unknown  6  GNR  Hylaeus hula#  Hulan yellow‐ faced bee  Dry and mesic  forest  Chamaesyce  olowaluana, Santalum  paniculatum  5  GNR  Hylaeus kona#  Kona yellow‐ faced bee  Dry forest  Bidens menziesii,  Chamaesyce  olowaluana, Myoporum  sandwicense  4  GNR  Hylaeus  mauiensis  Maui yellow‐ faced bee  Wet forest  unknown  6  GH  Hylaeus  melanothrix  Melanothrix  yellow‐faced bee  Wet forest  unknown  6  GH  Hylaeus nalo  A yellow‐faced  bee  Unknown; only  known from a  single 1914  specimen  unknown  6  GNR  Hylaeus niloticus  Obscuratan  yellow‐faced bee  Coastal; dry forest  and shrubland  unknown  6  GH  Hylaeus ombrias#  Ombrias yellow‐ faced bee  Coastal; dry forest  Bidens menziesii,  Chamaesyce  olowaluana, Scaevola  4  GNR 

Chapter 3. Imperiled Pollinator Profiles 3-15 Scientific Name  Common Name  Habitat  Floral Resources  Conservation Category*  Global  Conservation  Status**  sericea, Sida fallax,  Tetramolopium sp.,  Tribulus cistoides  Hylaeus  paradoxicus  Hawaiian yellow‐ faced bee  Dry and mesic  forest  Chamaesyce  olowaluana, Myoporum  sandwicense  5  GNR  Hylaeus  perkinsianus#  Perkin’s Yellow‐ faced Bee  Coastal; dry forest  and shrubland  unknown  1  G2  Hylaeus  perspicuus#  Perspicuan  yellow‐faced bee  Mesic and wet  forest  unknown  6  GH  Hylaeus  psammobius#  Psammobian  yellow‐faced bee  Coastal  Bacopa monnieri,  Sesuvium  portulacastrum  4  GNR  Hylaeus satelles#  Satellus yellow‐ faced bee  Wet forest  unknown  6  GH  Hylaeus simplex#  simple yellow‐ faced bee  Dry forest and  shrubland; mesic  forest  unknown  6  GH  Hylaeus solaris  A yellow‐faced  bee  Coastal  Scaevola sericea, Sida  fallax  4  GNR  Hylaeus volatilis#  Volatile Yellow‐ faced Bee  Coastal; dry forest  and shrubland  Styphelia tameiameiae  3  G2   # This species was petitioned for listing under the ESA but was not listed.  * Conservation category: 1: Uncommon to abundant, habitat safe; 2: Rare but widespread, habitat safe; 3: Missing from one or  more islands; 4: Restricted to endangered habitat; 5: Very rare, probably endangered; 6: No recent collections, endangered.  ‡ More recent collections exist (2013 surveys on Oahu).  ** Status of pollinators is taken from NatureServe (accessed March 2022).    G1: Critically Imperiled. At very high risk of extinction due to extreme rarity (often five or fewer populations), very steep  declines, or other factors.   G2: Imperiled. At high risk of extinction due to very restricted range, very few populations (often 20 or fewer), steep  declines, or other factors.   G3: Vulnerable. At moderate risk of extinction due to a restricted range, relatively few populations (often 80 or fewer),  recent and widespread declines, or other factors.   G4: Apparently Secure. Uncommon but not rare; some cause for long‐term concern due to declines or other factors.   G5: Secure. Common; widespread and abundant.   GNR: No status rank.   GH: Possibly extinct.   

3-16 3.2.1 References Daly, H. V., and K. N. Magnacca. 2003. Insects of Hawaii. Vol. 17. Hawaiian Hylaeus (Nesoprosopis) Bees (Hymenoptera: Apoidea). University of Hawai’i Press, Honolulu. 234 pp. Graham, J., C. King, and W. Haines. 2015. “The nest architecture of Hylaeus anthracinus and other coastal solitary bees.” Hawaii Conservation Conference Proceedings. August 3–6, 2015. University of Hawaii, Hilo, Hawaii. Graham, J., S. Plentovich, and C. King. 2016. Nest ecology of an endemic Hawaiian bee, Hylaeus anthracinus (Hymenoptera: Colletidae), and implications for conservation. University of Hawaii at Manoa. College of Tropical Agriculture and Human Resources. Unpublished Data. Howarth, F. G. 1985. “Impacts of alien land arthropods and mollusks on native plants and animals in Hawaii.” Hawaii’s terrestrial ecosystems: preservation and management. University of Hawaii Press, Honolulu, Hawaii. Pp. 149–179. iNaturalist. 2020. iNaturalist Network (web application): “Hylaeus.” Available: http://www.inaturalist.org/. Accessed: September 2020. Ing, K., and C. L. Mogren. 2020. “Evidence of competition between honey bees and Hylaeus anthracinus (Hymenoptera: Colletidae), an endangered Hawaiian yellow-faced bee.” Pacific Science 74(1):75–85. Koch, J. B., and H. F. Sahli. 2013. “Patterns of flower visitation across elevation and successional gradients in Hawai ‘i.” Pacific Science 67(2):253–266. Kuppler, J., M. K. Höfers, W. Trutschnig, A. C. Bathke, J. A. Eiben, C. C. Daehler, and R. R. Junker. 2017. “Exotic flower visitors exploit large floral trait spaces resulting in asymmetric resource partitioning with native visitors.” Functional Ecology 31(12):2244– 2254. Lach, L. 2005. “Interference and exploitation competition of three nectar-thieving invasive ant species.” Insectes Sociaux 52:257–262. Lach, L., and L. M. Hooper-Bui. 2010. Consequences of ant invasions. In: Lach, L., C. L. Parr, and K. L. Abbott (Eds.), Ant Ecology. Oxford University Press, Inc., Oxford. pp. 261–286. Magnacca, K. N. 2005a. Species Profile: Hylaeus anthracinus. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD- ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR. Magnacca, K. N. 2005b. Species Profile: Hylaeus assimulans. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD- ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR.

Chapter 3. Imperiled Pollinator Profiles 3-17 Magnacca, K. N. 2005c. Species Profile: Hylaeus facilis. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD-ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR. Magnacca, K. N. 2005d. Species Profile: Hylaeus hilaris. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD- ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR. Magnacca, K. N. 2005e. Species Profile: Hylaeus kuakea In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD- ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR. Magnacca, K. N. 2005f. Species Profile: Hylaeus longiceps. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD- ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR. Magnacca, K. N. 2005g. Species Profile: Hylaeus mana. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds.), Red List of Pollinator Insects of North America. CD-ROM Version 1 (May 2005). The Xerces Society for Invertebrate Conservation, Portland, OR. Magnacca, K. N. 2007a. “Conservation status of the native bees of Hawaii, Hylaeus (Nesoprosopis) (Hymenoptera: Apoidea).” Pacific Science 61(2):173–190. Magnacca, K. N. 2007b. “New records of Hylaeus (Nesoprosopis) and Ceratina bees in Hawai’i.” Bishop Museum Occasional Papers 96:44–45. Honolulu, Hawaii. Magnacca, K. N., and B. N. Danforth. 2006. “Evolution and biogeography of native Hawaiian Hylaeus bees (Hymenoptera: Colletidae).” Cladistics 22(5):393–411. Magnacca, K. N. 2018. Personal database of Hawaiian Hylaeus records. Entomological Specialist, Oahu Army Natural Resources Program. Magnacca, K. N., and C. King. 2013. Assessing the presence and distribution of 23 Hawaiian yellow-faced bee species on lands adjacent to military installations on O’ahu and Hawai’i Island. Technical Report No. 185. Pacific Cooperative Studies Unit, University of Hawaii, Honolulu, Hawai’i. 39 pp. Medeiros, M. J., J. A. Eiben, W. P. Haines, R. L. Kaholoaa, C. B. A. King, P. D. Krushelnycky, K. N. Magnacca, D. Rubinoff, F. Starr, and K. Starr. 2013. “The importance of insect monitoring to conservation actions in Hawaii.” Proceedings of the Hawaiian Entomological Society 45:149–166. Michener, C. D. 2007. The Bees of the World. 2nd Edition. Johns Hopkins University Press, Baltimore, Maryland. 972 pp. Miller, A. E., B. J. Brosi, K. N. Magnacca, G. C. Daily, and L. Pejchar. 2015. “Pollen carried by native and non-native bees in the large-scale reforestation of pastureland in Hawaii: implications for pollination.” Pacific Science 69(5):67–79. NatureServe. 2020. NatureServe Explorer (web application). NatureServe, Arlington, Virginia. Available: https://explorer.natureserve.org/. Accessed: September 2020.

Chapter 3. Imperiled Pollinator Profiles  3-18 Pan, A. D., and J.S. Wilson. 2020. “A scientific note on the behavior of the endangered Anthricinan yellow-faced bee (Hylaeus anthracinus) from South Kohala District, Hawai’i, Hawai’i.” Apidologie:1–4. Perkins, R. C. L. 1899. Hymenoptera Aculeata. In David Sharp (Ed.), Fauna Hawaiiensis. Vol. 1, Part 1. Cambridge University Press, Cambridge, United Kingdom. pp. 1–115. Perkins, R. C. L. 1913. Introduction. Pages i–ccxxvii In D. Sharp (Ed.), Fauna Hawaiiensis. Vol. 1. Cambridge University Press, London. Sakai, A. K., W. L. Wagner, and L. A. Mehrhoff. 2002. “Patterns of endangerment in the Hawaiian flora.” Systematic Biology 51(2):276–302. Schonberg, L., S. Jepsen, and S. H. Black. 2009a. Petition to list two species of Hawaiian yellow-faced bees Hylaeus anthracinus and Hylaeus longiceps as endangered under the U.S. Endangered Species Act. Submitted by the Xerces Society for Invertebrate Conservation to the U.S. Fish and Wildlife Service. Portland, Oregon. 49 pp. Schonberg, L., S. Jepsen, and S. H. Black. 2009b. Petition to list one species of Hawaiian yellow-faced bee Hylaeus assimulans as an endangered species under the U.S. Endangered Species Act. Submitted by the Xerces Society for Invertebrate Conservation to the U.S. Fish and Wildlife Service. Portland, Oregon. 34 pp. Schonberg, L., S. Jepsen, and S. H. Black. 2009c. Petition to list one species of Hawaiian yellow-faced bee Hylaeus facilis as an endangered species under the U.S. Endangered Species Act. Submitted by the Xerces Society for Invertebrate Conservation to the U.S. Fish and Wildlife Service. Portland, Oregon. 38 pp. Schonberg, L., S. Jepsen, and S. H. Black. 2009d. Petition to list one species of Hawaiian yellow-faced bee Hylaeus hilaris as an endangered species under the U.S. Endangered Species Act. Submitted by the Xerces Society for Invertebrate Conservation to the U.S. Fish and Wildlife Service. Portland, Oregon. 23 pp. Schonberg, L., S. Jepsen, and S. H. Black. 2009e. Petition to list two species of Hawaiian yellow-faced bees Hylaeus mana and Hylaeus kuakea as endangered under the U.S. Endangered Species Act. Submitted by the Xerces Society for Invertebrate Conservation to the U.S. Fish and Wildlife Service. Portland, Oregon. 24 pp. Swezey, O. H. 1954. Forest entomology in Hawaii. Bernice P. Bishop Museum Special Publication 44. 266 pp. Swezey, O. H. and E. H. Bryan. 1929. “Further notes on the forest insects of Molokai.” Proceedings of the Hawaiian Entomological Society 7(2):293–314. U.S. Fish and Wildlife Service. 1984. Endangered and threatened wildlife and plants: Review of invertebrate wildlife for listing as endangered or threatened species. Federal Register 49:21664–21673. U.S. Fish and Wildlife Service. 2008. Hawaiian Islands Plants: Listed species, as designated under the U.S. Endangered. Endangered and Threatened Wildlife and Plants; Review of Native Species That Are Candidates for Listing as Endangered or Threatened; Annual Notice of Findings on Resubmitted Petitions; Annual Description of Progress on Listing Actions; Proposed Rule Species Act. 50 CFR Part 17. 70 pp.

Chapter 3. Imperiled Pollinator Profiles 3-19 U.S. Fish and Wildlife Service. 2016. Endangered Status for 49 Species from the Hawaiian Islands. September 30, 2016. Federal Register 81(190):67786–67860. Wagner, W. L., D. R. Herbst, and S. H. Somer. 1999. Manual of the Flowering Plants of Hawai’i. Revised Edition. Mill, S. W. (Ed.). University of Hawai’i Press, Bishop Museum Press, Honolulu, Hawai’i. 1919 pp. Warshauer, F. R., J. D. Jacobi, and J. Price. 2009. Native coastal flora and plant communities in Hawai’i: Their composition, distribution, and status. Hawai’i Cooperative Studies Unit Technical Report HCSU-014. University of Hawai’i at Hilo. 108 pp. Wilson, E. E., C. S. Sidhu, K. E. LeVan, and D. A. Holway. 2010. “Pollen foraging behaviour of solitary Hawaiian bees revealed through molecular pollen analysis.” Molecular Ecology 19(21):4823–4829. 3.3 Blackburn’s Sphinx Moth (Manduca blackburni)      Jan  Feb  Mar  April  May  June  July  Aug  Sept  Oct  Nov  Dec  Adult                                                                                                Larvae                                                                                                Adult Blackburn’s sphinx moth (top left) and known distribution (top right). Adult flight times (i.e., breeding  period; blue) and larvae active periods (green) are shown in the chart (bottom). Darker blue shades  indicate peak adult flight time. Lighter green shades indicate uncertain but likely active larval times. See  Life Cycle for more information. Photo credit: Wikimedia. Map Source: ecos.fws.gov, accessed March 2022.    Order: Lepidoptera Family: Sphingidae Status: Endangered Distribution: Hawaii. It is currently known to occur in small, localized populations on the islands of Maui, Hawaii, and Kahoolawe; it has disappeared from the islands of Kauai, Oahu, and Molokai where it occurred historically. Where it occurs: This species was once abundant throughout the Hawaiian Islands but is now reduced to a few highly localized, disjunct, low-density populations. The moth likely inhabited much of the dry and mesic shrubland and forest, which historically covered over two million acres on all the main Hawaiian Islands prior to human settlement. This moth

Chapter 3. Imperiled Pollinator Profiles  3-20 is found in association with diverse landscapes (from near sea level along the coast to montane forests up to 1,525 meters elevation) that have a mix of both native and nonnative vegetation, and it is tied to areas where host plants occur. Both the caterpillar and adult food plants occur primarily in undeveloped habitats that have a mix of native and nonnative vegetation but can also be found in more urbanized areas. How to recognize: Blackburn’s sphinx moth is one of Hawaii’s largest native insects, having a wing span of up to 12 centimeters. The body is grayish brown in color with five orange spots along each side of the abdomen. Similar to other sphinx moths in the Sphingidae family, adults have long, narrow forewings and a thick, spindle-shaped body tapered at both ends. Although similar in appearance, Blackburn’s sphinx moth can be distinguished from the closely related tomato hornworm (M. quinquemaculata, introduced to Hawaii) by the last of the five orange spots on the top of the abdomen, which are a quarter of the size in the tomato hornworm. Additionally, these two species can be distinguished by a narrow cross-band of white triangular spots in Blackburn’s sphinx moth inside the outer edge of the forewings; its wings are also decidedly broader than that of the tomato hornworm. The grayish brown forewings have light whitish markings and two arched double bands; the inner one is black edged, with white markings, which are absent in the similar tomato hornworm and tobacco hornworm (M. sexta, also introduced to Hawaii). Life cycle: Adult moths are active at night and cryptic during the day. They lay their eggs on the leaves of native and nonnative plant species in the Solanaceae family (nightshade), host plants on which their caterpillars feed (Table 3-7). Blackburn’s sphinx moth caterpillars have five instar stages, each of which last several days. When caterpillars are ready to pupate, they are known to wander from the host plant to burrow into soil, often near the roots of their host species. Blackburn’s sphinx moth caterpillars have been documented primarily between October and May; however, they have also been documented during the summer, and may likely be active all year. From egg to pupa, development time is estimated at approximately 60 days. Adult moths may emerge within 6 weeks of pupation; however, pupae may spend up to a year underground. The adult life span is unknown. Adult moths feed on nectar and are active throughout the year, although two main flight periods have been observed between January and April and July through October. Adults typically oviposit following seasonal rains, which largely occur from October through April. Both eggs and later instar larvae have been found at the same time in the field, indicating extended adult flight periods or overlap among multiple generations. Habitat needs: Blackburn’s sphinx moth has three basic habitat needs: (1) foraging resources for caterpillars (host plants); (2) foraging resources for adults (nectar plants); and (3) habitat for egg laying, development, breeding, sheltering, and resting. The full range of caterpillar host plants and adult nectar plants used by this species is not known, but documented food plants are given in Table 3-7. Top reasons for decline, if known: Blackburn’s sphinx moth has declined throughout its native range, largely due to habitat loss and degradation as well as the introduction of nonnative plants, predators, and parasitoids. Hawaiian habitats where this species occurs have been significantly transformed by human influence as well as domestic and feral ungulates. Other important factors include pesticide use and light pollution. General conservation recommendations: Critical habitat has been designated and recovery plans developed to aid in the moth’s and their endangered host plants’ recovery.

Chapter 3. Imperiled Pollinator Profiles 3-21 Restoration that may benefit this species includes establishing habitat with its known native host and nectar plants and protecting current habitat. The dependence of Blackburn’s sphinx moth on the invasive tree tobacco discourages its removal from dry forest habitat and roadsides under restoration. However, native Nothocestrum host species are more stable and persistent components of dry to mesic forest habitats than tree tobacco; consequently, as a nonnative, short-lived species, it may be affected more during extended drought or successional changes in plant communities. These attributes may indicate that native host species plantings will fare better in roadside restoration areas and will help contribute to the moth’s recovery, as well as the recovery of dry and mesic habitats in Hawaii. Roadside management recommendations: This species may be sensitive year-round to management that could disturb caterpillars and eggs on host plants (e.g., mowing, chemical applications), pupae (e.g., ground excavation, chemical applications) or adult moths seeking shelter among roadside vegetation (e.g., mowing, chemical applications). Care must be taken during roadside maintenance and management near Blackburn’s sphinx moth host plant habitat to avoid practices that disturb soil or damage host plants. Damage or loss of host plants causes direct harm to eggs and caterpillars, and ground disturbance will harm pupae within the soil. Intensive mowing and invasive species management may deplete floral resources and potentially disturb roadside host plants. Indiscriminate spraying of herbicides on roadsides can also affect the health of moths by removing floral resources. Using seed mixes that are dominant in nonnative plants instead of native flowering species will be less effective in supporting native moths. Despite the moth’s reliance on nonnative tree tobacco, a common roadside weed as a caterpillar host plant, tree tobacco has become increasingly invasive and targeted for removal during restoration and fuel-break clearing activities in some areas where the moth occurs. Targeted removal of this invasive but critical host plant presents a significant conservation challenge for roadside managers. Targeting smaller seedlings before they grow to 1 meter has been shown to limit impacts on the moth, which is found to use larger plants for reproduction. However, the preservation of tree tobacco is currently considered a conservation expedient to support populations of the moth given the rarity of native host plants. Because this species is federally listed as endangered, consultation with the U.S. Fish and Wildlife Service and a Biological Opinion will be required if invasive host plants are targeted for removal in designated critical habitat. Table 3-7. Plants used by Blackburn’s sphinx moth. Species Name  Common Name  Notes  Larval Host Plants  Nicotiana glauca  Tree tobacco  Nonnative  Nothocestrum breviflorum  Aiea/smallflower aiea  Native; endangered  Nothocestrum latifolium  Aiea/broadleaf aiea  Native; endangered  Nothocestrum longifolium*  Aiea/longleaf aiea  Native; endangered 

Chapter 3. Imperiled Pollinator Profiles  3-22 Species Name  Common Name  Notes  Nothocestrum peltatum*  Aiea/Oahu aiea  Native; endangered  Solanum americanum  Olohua/American black nightshade  Native  Solanum sandwicense  Popolo/Hawaii horsenettle  Native; endangered  Nectar Plants  Ipomoea indica  Koali awa/oceanblue morning‐glory  Native  Pleomele auwahiensis  Hala pepe/St. John Maui hala pepe  Native  Capparis sandwichiana*  Maiapilo/native caper  Native  Plumbago zeylanica*  Hiliee/wild leadwort  Native  *Presumed  3.3.1 References Cuddihy, L. W., and C.P. Stone. 1990. Alteration of native Hawaiian vegetation-effects of humans, their activities and introductions. University of Hawaii Cooperative National Park Studies Unit, University of Hawaii Press, Honolulu, Hawaii. 138 pp. Hawaii Department of Land and Natural Resources. 2015. Hawai’i’s State Wildlife Action Plan. Prepared by H. T. Harvey and Associates, Honolulu, Hawai’i. 1055 pp. Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife. 2015. Draft Habitat Conservation Plan for Game Management at Pu’u Wa’awa’a and Pu’u Anahulu. Napu’u Conservation Project. 269 pp. Elliott, C. H. 2019. The Riddle of the Sphinx: Population ecology of the endangered Blackburn’s sphinx moth, Manduca blackburni (Lepidoptera: Sphingidae) on an invasive host plant. Doctoral dissertation, University of Hawaiʻi at Mānoa. 60 pp. Gagné, W. C., and C. C. Christensen. 1985. “Conservation status of native terrestrial invertebrates in Hawaii.” Hawaii’s terrestrial ecosystems: preservation and management. University of Hawaii Cooperative National Park Resources Studies Unit, Honolulu, pp. 105–126. Gagné, W. C., and F. G. Howarth. 1982. “Conservation Status of Endemic Hawaiian Lepidoptera.” Proceedings of the Third Congress of European Lepidopterology, Societus Europaea Lepidopterologica, Karluhe. Cambridge. pp. 74–84. Howarth, F. G., and W. P. Mull. 1992. Hawaiian Insects and their Kin. University of Hawaii Press, Honolulu. pp. 124-125. Howarth, F. G., D. J. Preston, and R. Pyle. 2012. Surveying for Terrestrial Arthropods (Insects and Relatives) Occurring within the Kahului Airport Environs, Maui, Hawai’i: Synthesis Report. Hawaii Biological Survey. Bishop Museum Technical Report 58, Honolulu, Hawaii. 225 pp.

Chapter 3. Imperiled Pollinator Profiles 3-23 Kitching, I. J., and J. M. Cadiou. 2000. Hawkmoths of the World. Comstock Publishing Associates, Cornell University Press. Ithaca, New York. 227 pp. Liebherr, J. K., and D. A. Polhemus. 1997. “R. C. L. Perkins: 100 years of Hawaiian entomology.” Pacific Science 51(4):343–355. Macgregor, C. J., M. J. Pocock, R. Fox, and D. M. Evans. 2015. “Pollination by nocturnal Lepidoptera, and the effects of light pollution: a review.” Ecological Entomology 40(3):187–198. Medeiros, A. C., L. L. Loope, and F. R. Cole. 1986. “Distribution of ants and their effects on endemic biota of Haleakala and Hawaii Volcanoes National Park: a preliminary assessment.” Proceedings of the 6th Conference of Natural Sciences, Hawaii Volcanoes National Park. Pp. 39–52. Medeiros, A., and E. vonAllmen. 2006. Restoration of a native Hawaiian dryland forest at Auwahi, Maui. U.S. Geological Survey Fact Sheet 2006–3035. Medeiros, M. J., J. A. Eiben, W. P. Haines, R. L. Kaholoaa, C. B. A. King, P. D. Krushelnycky, K. N. Magnacca, D. Rubinoff, F. Starr, and K. Starr. 2013. “The importance of insect monitoring to conservation actions in Hawaii.” Proceedings of the Hawaiian Entomological Society 45:149–166. NatureServe. 2020. NatureServe Explorer (web application). NatureServe, Arlington, Virginia. Available: https://explorer.natureserve.org/. Accessed: September 2020. Riotte, J. C. E. 1986. “Re-evaluation of Manduca blackburni (Lepidoptera: Sphingidae).” Proceedings of the Hawaiian Entomological Society 27:79–90. Rubinoff, D. and M. San Jose. 2010. “Life history and host range of Hawaii’s Endangered Blackburn’s Sphinx Moth (Manduca blackburni Butler).” Proceedings of the Hawaiian Entomological Society 42:53–59. Rubinoff, D., M. San Jose, and A. Y. Kawahara. 2012. “Phylogenetics and species status of Hawai’i’s endangered Blackburn’s Sphinx moth, Manduca blackburni (Lepidoptera: Sphingidae).” Pacific Science 66(1):31–41. Sakai, A. K., W. L. Wagner, and L. A. Mehrhoff. 2002. “Patterns of endangerment in the Hawaiian flora.” Systematic Biology 51(2):276–302. Shay, K. R. 2014. Pollination ecology of Hawaiian coastal plants. Ph. D. Dissertation. University of Hawaii at Manoa, Honolulu, Hawaii. 76 pp. Symon, D. E. 1999. Solanaceae in: Manual of the Flowering Plants of Hawaii. W. L. Wagner, D. R. Herbst, and S. H. Sohmer, (eds.). University of Hawaii Press and Bishop Museum Press, Honolulu. Bishop Museum Special Publications. pp. 1251–1278. Tuttle, J. P. 2007. The Hawk Moths of North America. Wedge Entomological Research Foundation. Allen Press. Lawrence, Kansas. 253 pp. US Fish and Wildlife Service. 1994a. Endangered and threatened wildlife and plants: determination of endangered or threatened status for 24 plants from the Island of Kauai, Hawaii. Federal Register 59(38):9304–9329.

Chapter 3. Imperiled Pollinator Profiles  3-24 US Fish and Wildlife Service. 1994b. Endangered and threatened wildlife and plants; determination of endangered or threatened status for 21 plants for the Island of Hawaii, State of Hawaii. Federal Register 59(43):10305–10325. U.S. Fish and Wildlife Service. 1995. Recovery Plan for the Kauai Plant Cluster. Portland, Oregon. 270 pp. U.S. Fish and Wildlife Service. 1996. Recovery Plan for the Big Island Plant Cluster. Portland, Oregon. 202+ pp. U.S. Fish and Wildlife Service. 2003. Endangered and threatened wildlife and plants; designation of critical habitat for Blackburn’s sphinx moth, final rule. Federal Register 68(111):34710–34766. U.S. Fish and Wildlife Service. 2005. Recovery Plan for the Blackburn’s Sphinx Moth (Manduca blackburni). Portland, Oregon. 125 pp. U.S. Fish and Wildlife Service. 2012a. Endangered and Threatened Wildlife and Plants; Endangered Status for 23 Species on Oahu and Designation of Critical Habitat for 124 Species; Final Rule. Federal Register 77(180):57648–57862. U.S. Fish and Wildlife Service. 2012b. Endangered and Threatened Wildlife and Plants; Listing 15 Species on Hawaii Island as Endangered and Designating Critical Habitat for 3 Species; Proposed Rule. Federal Register 77(201):63928–64018. U.S. Fish and Wildlife Service. 2016. Endangered and threatened wildlife and plants: Endangered Status for 49 Species from the Hawaiian Islands. Federal Register 81(190):67786–67857. Wagner, W. L., D. R. Herbst, and S. H. Somer. 1999. Manual of the Flowering Plants of Hawaii. Revised Edition. Mill, S. W. (ed.). University of Hawaii Press, Bishop Museum Press, Honolulu, Hawai’i. 1919 pp. Williams, F. X. 1947. “Notes and Exhibitions: Protoparce quinquemaculata blackburni (Butler).” Proceedings of the Hawaiian Entomological Society 13:10.

Chapter 3. Imperiled Pollinator Profiles 3-25 3.4 Fabulous Green Sphinx of Kauai (Tinostoma  smaragditis)    Jan  Feb  Mar  April  May  June  July  Aug  Sept  Oct  Nov  Dec  Adult                                                                                                Larvae                                                                                                Adult fabulous green sphinx of Kauai (top left) and known island‐level distribution (top right). Adult flight  times (i.e., breeding period; blue) and larvae active periods (green) are shown in the chart (bottom). Very  little information exists on this species’ phenology. Individuals have been collected in February and  October. It is probably best to assume it is active year‐round, like other pollinators in Hawaii (see Life Cycle  for more). Photo credit: Robert Young ‐ http://sphingidae.myspecies.info/file‐colorboxed/13520, CC BY  3.0, https://commons.wikimedia.org/w/index.php?curid=46785818. Map Source: ecos.fws.gov, accessed  March 2022.  Order: Lepidoptera Family: Sphingidae Status: Critically imperiled to imperiled (G1G2) Distribution: Hawaii Where it occurs: Fewer than 20 observations have been made of the Kauai-endemic fabulous green sphinx in the western part of the island. This moth likely inhabits lowland mesic forests on Kauai; a few observations are known directly from the forests themselves, while other incidental observations were made in adjacent urbanized areas as moths were likely attracted by artificial lights. How to recognize: This species is a vibrantly colored moth with brilliant green forewings and thorax. Its hindwings are brown and it has orange antennae. The moth is sexually dimorphic; males have a black spot on the forewing and a black band on the thorax, while females lack these markings. The underside of the male’s forewing and hindwing are pinkish brown, whereas those of the female are a pale green. Life cycle: Very little is known about this species’ life cycle. Most observations of the moth have been made of larval specimens that hatched from eggs in captivity; however, development time and life span were not detailed except for the egg stage, which lasted about a week before the caterpillars hatched. Habitat needs: Little information exists on this species’ habitat needs. Despite studies to determine caterpillar host plants, these and nectar plants still remain unknown. The moth

Chapter 3. Imperiled Pollinator Profiles  3-26 likely relies on native and endemic mesic forest plant species. Because this moth has only rarely been documented and its host and nectar plants still remain unknown, it is possible the plants tied to their existence are either limited in distribution or are themselves imperiled. In fact, the moth is a suspected pollinator of alula or cabbage on a stick (Brighamia insignis), which is listed as endangered and threatened with extinction in the wild. Top reasons for decline, if known: This species remains more elusive than other endemic sphinx moths, including the endangered Blackburn’s sphinx moth, which is far better documented. It remains so rare that it was thought extinct several times, only to reappear; however, this species has not been seen in over 20 years. The rarity of the moth’s habitat is likely linked to its own rarity; like Blackburn’s sphinx moth, this species occurs in globally imperiled mesic forest habitat. Hawaiian mesic forests have a great number of at- risk endemic plants, and these habitats have largely become degraded or lost due to land management practices and impacts from feral ungulates. Light pollution may be problematic near the perimeters of mesic forests, disrupting the behavior of nocturnal moths. In fact, this moth has been documented in urbanized areas near mesic forest, presumably attracted to artificial lighting, and its response to light is known from several collection occasions. Introduced wasps, which are known to attack the eggs of sphinx moths, may contribute to the decline of rare endemic species, further endangering remaining populations. General conservation recommendations: This species remains too poorly known to develop specific conservation recommendations. An attempt to list the species under the ESA failed due to insufficient information on its life history and specific threats. Protecting native mesic forest habitat, where it is thought to occur, will likely benefit this species. Studies to understand its population status, as well as caterpillar host and adult food plants, are needed in order to better protect and conserve the fabulous green sphinx of Kauai. Roadside management recommendations: Damage or loss of host plants will cause direct harm to eggs and caterpillars; however, with host plants unknown, impacts of roadside management specific to this species are difficult to define. While the full life cycle is unknown, if this species pupates in the soil similar to other known sphinx moths, it may be sensitive year-round to management that could disturb the soil (e.g., ground excavation). Adult moths seeking shelter among roadside vegetation may also be affected by mowing and chemical applications. Indiscriminate spraying of herbicides on roadsides may affect the health of moths by removing host plants and floral resources. Because this species likely utilizes native and endemic plants, roadside restoration with nonnative plantings instead of native flowering species will be less effective in supporting the fabulous green sphinx of Kauai. 3.4.1 References Cambell, C. L., and L. M. Ishii. 1993. “Larval host plant testing of Tinostoma smaragditis (Lepidoptera: Sphingidae), the Fabulous Green Sphinx of Kauai.” Proceedings of the Hawaiian Entomological Society 32:83–90.

Chapter 3. Imperiled Pollinator Profiles 3-27 Funasaki, G. Y., P. L. Lai, L. M. Nakahara, J. W. Beardsley, and A. K. Ota. 1988. “A review of biological control introductions in Hawaii: 1890 to 1985.” Proceedings of the Hawaiian Entomological Society 28:105–160. Heddle, M. L., K. R. Wood, A. Asquith, and R. G. Gillespie. 2000. “Conservation status and research on the Fabulous Green Sphinx of Kaua’i, Tinostoma smaragditis (Lepidoptera: Sphingidae), including checklists of the vascular plants of the diverse mesic forests of Kaua’i, Hawai’i.” Pacific Science 54(1):1–9. Heddle, M. L. 2004. Tinostoma smaragditis. The IUCN Red List of Threatened Species 2004: e.T21913A9339981. Available: https://dx.doi.org/10.2305/ IUCN.UK.2004.RLTS.T21913A9339981.en. Accessed: September 2020. Kitching, I. J., and J. M. Cadiou. 2000. Hawkmoths of the World. Comstock Publishing Associates, Cornell University Press. Ithaca, New York. 227 pp. Macgregor, C. J., M. J. Pocock, R. Fox, and D. M. Evans. 2015. “Pollination by nocturnal Lepidoptera, and the effects of light pollution: a review.” Ecological Entomology 40(3):187–198. NatureServe. 2020. NatureServe Explorer (web application). NatureServe, Arlington, Virginia. Available: https://explorer.natureserve.org/. Accessed September 2020. Sakai, A. K., W. L. Wagner, and L. A. Mehrhoff. 2002. “Patterns of endangerment in the Hawaiian flora.” Systematic Biology 51(2):276–302. 3.5 Hawaiian Blue Butterfly (Udara blackburni) (aka  Green Hawaiian blue, Blackburn’s blue, Blackburn’s  bluet, Koa blue butterfly, Koa Pulelehua, Hawaiian  hairstreak butterfly)     Jan  Feb  Mar  April  May  June  July  Aug  Sept  Oct  Nov  Dec  Adult                                                                                                Larvae                                                                                                Adult Hawaiian blue butterfly (top left) and known island‐level distribution (top right). Adult flight times  (i.e., breeding period; blue) and larvae active periods (green) are shown in the chart (bottom). See Life  Cycle for more. Photo credit: Kim Starr/Flickr Creative Commons. 

Chapter 3. Imperiled Pollinator Profiles  3-28 Order: Lepidoptera Family: Lycaenidae Status: Apparently secure (G4) Distribution: Hawaii. This species is described from the larger Hawaiian Islands, including Kauai, Oahu, Molokai, Maui, Lanai, and Hawaii; it is not known from Niihau and was only more recently documented on Kahoolawe. Where it occurs: The Hawaiian blue butterfly inhabits shrubland and forests from the coast to alpine woodlands up to about 2,700 meters (8,900 feet) in elevation. It is commonly associated with tropical woodlands on lower slopes, valleys, and forest edges, where its larval host plants grow. Although mainly associated with native forests, the Hawaiian blue butterfly is often seen in the lowlands, including urban gardens. Koa forests, where this species is commonly known, can be found in a diversity of Hawaiian ecosystems from dry, mesic, and wet lowland and montane areas. How to recognize: The Hawaiian blue is a small blue butterfly with a wingspan of 22 to 29 millimeters. Adult males are deep purplish-blue on the upper side of the wings, sometimes appearing a very dark olive-brown, while the upper side of the forewings of the female is a similar purplish blue but paler with a black, very broad outer border. The female hindwing is black with a blue patch similar in color to that of the forewing. The undersides of both male and female wings are turquoise-green. Additionally, the female thorax has an abundance of conspicuous green scales on its upper side (dorsally), which are absent or almost so on males. It is the only butterfly in the Lycaenidae family native to the Hawaiian Islands, although at least eight different related species in the same family have been documented in Hawaii. The green undersides of the Hawaiian blue’s wings, which lack spots, easily distinguish it from the other related introduced species. Adults usually hold their wings upright (folded up), displaying the glittery green underside of the wings. Life cycle: Despite being one of two endemic butterflies to the Hawaiian Islands, little is known about the life history of this small native butterfly. Adult butterflies lay eggs on host plants (Table 3-8) and their caterpillars feed on flowers, buds, and young leaves, where they are usually well camouflaged. The caterpillar is known to remove the anthers of the koa flower head in order to access the ovary and calyx (collectively, the sepals, which are the outermost parts of a flower bud), which it consumes. When the caterpillar is ready to pupate, it spins a thin layer of silk onto a substrate and then attaches itself to the silk layer by looping a thread over its body. The pupa may attach itself lengthwise to a leaf or seed with a few threads. The pupal stage of the Hawaiian blue may last about 8 to 10 days. All life stages may be present or active all year, but activity may vary depending on habitat and elevation. Habitat needs: The Hawaiian blue is tied to habitats with its native host plants (Table 3- 8). This species’ main host plant, koa (Acacia koa), occurs in a variety of habitats from all of the main Hawaiian Islands except Kahoolawe and Niihau. Another primary caterpillar host plant for this species is a’ali’i (Dodonaea viscosa), a native shrub in the Sapindaceae family (soapberries), which is adapted to a wide range of habitats. On the island of Kahoolawe, the Hawaiian blue caterpillar is known to be associated with a’ali’i restoration plantings, because koa is not found on the island. Other documented caterpillar host plants include both native and nonnative shrubs and trees from several different families—

Chapter 3. Imperiled Pollinator Profiles 3-29 in the Fabaceae family (legumes): manila tamarind (Pithecellobium; nonnative) and monkeypod (Samanea saman; nonnative, naturalized); in the Celastraceae family (bittersweet): olomea (Perrottetia sandwicensis); in the Utricaceae family (nettles): mamaki (Pipturus albidus); and in the Rubiaceae family (coffee, madder): Hedyotis sp. Hawaiian blue caterpillars also feed on other native and nonnative legumes, soapberries, nettles, bittersweet, and coffee plants. Top reasons for decline, if known: Although direct threats to this species are not known, major threats to Hawaiian Lepidoptera include habitat loss and degradation, loss of host plants, and nonnative animal and plant introductions. One of the main threats to the species is likely habitat loss, primarily from the loss of adult and larval foraging habitat due to damage by past and present management practices, feral ungulates, and the invasion of nonnative plants. Historical land use practices have severely degraded or destroyed native koa forests on the Hawaiian Islands, which contain this species’ primary host plant. Some of the most serious threats to native Lepidoptera likely come from parasitic wasps, which directly parasitize eggs, caterpillars, and pupae, or from introduced ants, which prey on caterpillars or exclude them from suitable habitat. General conservation recommendations: Conserving and restoring populations by reducing or eliminating the presence of nonnative weeds and planting native host plants can be an effective conservation strategy to address loss of habitat for this species. Reducing threats that diminish or degrade habitat such as feral ungulates can also help promote native habitat restoration. In addition, carefully planned management in suitable habitat areas can help ensure the butterfly is protected from any potential adverse actions. Roadside management recommendations: Intensive mowing and invasive species management may deplete floral resources and potentially disturb host plants occurring along roadsides. Because they have overlapping life stages throughout the year, the Hawaiian blue may be sensitive to year-round management that could disturb or remove host or nectar plants (e.g., indiscriminate herbicide application, mowing). Using seed mixes with predominantly nonnative plants instead of native flowering species will be less effective in supporting the Hawaiian blue butterfly. Table 3-8. Plants used by Hawaiian blue butterfly. Species Name  Common Name  Notes  Larval Host Plants  Acacia koa  koa oha/koa  Native; primary food plant  Dodonaea viscosa  a’ali’i /Florida hopbush  Native; primary food plant  Hedyotis spp.  starviolet  Native  Perrottetia sandwicensis  olomea  Native  Pithecellobium dulce  monkeypod  Nonnative  Pipturus albidus  mamaki/Waimea pipturus  Native 

Chapter 3. Imperiled Pollinator Profiles  3-30 Species Name  Common Name  Notes  Samanea saman  raintree  Nonnative; naturalized  Nectar plants  Acacia koa  koa oha/koa  Native  Bidens micrantha  ko’oko’olau/grassland beggarticks  Native  Dubautia menziesii  mountain dubautia  Native  Metrosideros polymorpha  lehua/’Ohi’a lehua  Native  Myoporum spp.  naio/Myoporum  Native  Osteomeles anthyllidifolia  ulei/Hawaii hawthorn  Native  Santalum freycinetianum  iliahi/sandalwood  Native  Senecio madagascariensis  Madagascar ragwort  Nonnative  Waltheria indica  hala uhaloa/sleepy morning  Native  3.5.1 References Blackburn, T. 1882. “Descriptions of the larvae of Hawaiian Lepidoptera.” The Entomologist’s Monthly Magazine 19:55–56. Cuddihy, L. W., and C. P. Stone. 1990. Alteration of native Hawaiian vegetation-effects of humans, their activities and introductions. University of Hawaii Cooperative National Park Studies Unit, University of Hawaii Press, Honolulu, Hawaii. 138 pp. Gagné, W. C., and C. C. Christensen. 1985. “Conservation status of native terrestrial invertebrates in Hawaii.” Hawaii’s terrestrial ecosystems: preservation and management. University of Hawaii Cooperative National Park Resources Studies Unit, Honolulu, Hawaii. pp. 105–126. Gagné, W. C., and F. G. Howarth. 1982. “Conservation Status of Endemic Hawaiian Lepidoptera.” Proceedings of the Third Congress of European Lepidopterology, Societus Europaea Lepidopterologica, Karluhe. Cambridge. pp. 74–84.  Haines, W. 2018. Personal communication with Michele Blackburn and Candace Fallon, the Xerces Society. Research Entomologist, Hawaii Invertebrate Program, Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife, University of Hawaii, Center for Conservation Research and Training, Honolulu, Hawaii. 22 October 2018. Haines, W. 2019. Personal communication with Michele Blackburn, Conservation Biologist, the Xerces Society. Research Entomologist, Hawaii Invertebrate Program, Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife

Chapter 3. Imperiled Pollinator Profiles 3-31 University of Hawaii, Center for Conservation Research and Training. January 8–10, 2019. Howarth, F. G., and W. P. Mull. 1992. Hawaiian Insects and Their Kin. University of Hawaii Press. Honolulu, Hawaii. 160 pp. Liebherr, J. K., and D. A. Polhemus. 1997. “R. C. L. Perkins: 100 years of Hawaiian entomology.” Pacific Science 51(4):343–355. NatureServe. 2020. NatureServe Explorer (web application). NatureServe, Arlington, Virginia. Available: https://explorer.natureserve.org/. Accessed: September 2020. Patrick, B., and H. Patrick. 2012. Butterflies of the South Pacific (Vol. 240). Otago University Press. Dunedin, New Zealand. 240 pp. Robinson, G. S., P. R. Ackery, I. J. Kitching, G. W. Beccaloni, and L. M. Hernández. 2010. HOSTS - A Database of the World’s Lepidopteran Hostplants. Natural History Museum, London. Available: http://www.nhm.ac.uk/hosts. Accessed: September 2020. Scott, J. A. 1992. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press. Stanford, California. 668 pp. Starr Environmental. 2020. “Documented plant and arthropod observations on Hawaii.” Available: http://www.starrenvironmental.com/resources/. Accessed: September 2020. Swezey, O. H. 1910. “The Feeding Habits of Hawaiian Lepidoptera.” Proceedings of the Hawaiian Entomological Society 2(3):131–143. Tuley, N. C. 1878. “Description of a new species of butterfly from the Sandwich Islands.” Entomologist’s Monthly Magazine 15:9–10. Wagner, W. L., D. R. Herbst, and S. H. Somer. 1999. Manual of the flowering plants of Hawai’i. Revised Edition. Mill, S. W. (Ed.). University of Hawai’i Press, Bishop Museum Press, Honolulu, Hawai’i. 1919 pp. Williams, F. X. 1947. “Notes and Exhibitions for the Year 1946.” Proceedings of the Hawaiian Entomological Society 13(1):1–31. Zimmerman, E. C. 1948. Insects of Hawaii: a manual of the insects of the Hawaiian Islands, including an enumeration of the species and notes on their origin, distribution, hosts, parasites, etc. University of Hawaii Press, Honolulu, HI. 222 pp. Zimmerman, E. C. 1958. Insects of Hawaii, Volume 7, Macrolepidoptera. Honolulu, Hawaii, University of Hawaii Press. 549 pp.

Chapter 3. Imperiled Pollinator Profiles  3-32 3.6 Kamehameha Butterfly (Vanessa tameamea)    Jan  Feb  Mar  April  May  June  July  Aug  Sept  Oct  Nov  Dec  Adult                                                                                                Larvae                                                                                                Adult Kamehameha butterfly (top left) and known island‐level distribution (top right). Adult flight times  (i.e., breeding period; blue) and larvae active periods (green) are shown in the chart (bottom). See Life  Cycle for more. Photo credit: Will Haynes/Hawaii DLNR‐DOFAW.  Order: Lepidoptera Family: Nymphalidae Status: Vulnerable to apparently secure (G3G4) Distribution: Hawaii. The butterfly is currently known to occur on the islands of Kauai, Oahu, Molokai, Maui, and Hawaii; it has disappeared from some sites on Oahu, including Tantalus where it occurred historically and may no longer be found on Lanai. Where it occurs: The Kamehameha butterfly is primarily a butterfly of mesic and wet forests. It is tied to areas where host plants occur, which are typically found in shady habitats or wet canyons with other native vegetation in areas of moderate to heavy rainfall. How to recognize: Adult Kamehameha butterflies are large, conspicuous orange and black butterflies with a wing span of about 3 inches or more. This butterfly is sexually dimorphic; females may be distinguished from the male by having a pair of white, rather than light orange, spots on the front margin of the forewing. In both sexes the pattern of the upper wing surface varies little between individuals, although the pattern on the underside can be highly variable (varying from grayish brown to olive green). Three other species in the Vanessa genus, which are similar in appearance, have been introduced to Hawaii. The Kamehameha butterfly can be distinguished from the other species of Vanessa by the number of white (female) or light orange (male) patches in the black area on the upper surface of the forewings. The Kamehameha butterfly has only three major white patches in this area, while the other species have additional small white spots. Life cycle: Adults may live around a month and appear to fly all months of the year. Although large and conspicuous, adults spend most of their time flying around the canopy, making them elusive and difficult to detect or document. They are fast, strong fliers that are most active on sunny days where they can be found with wings outstretched sunning

Chapter 3. Imperiled Pollinator Profiles 3-33 themselves in forest clearings. Adults may be seen resting on the trunk or limbs of their caterpillar host plants or other nearby trees. Males and females are attracted to fermenting sap from the wounds of trees, particularly on the trunks of koa. Females seek out host plants in the nettle family (Urticaceae) to lay their eggs. Eggs are laid singly on leaves or stems. Young caterpillars are known as leaf rollers; when they emerge they build a shelter by cutting a crescent-shaped flap in the surface of a host plant leaf. The cut leaf flap, which becomes the shelter, is bent over and fastened to the edge of the leaf surface with silk. The shelter is almost always folded downward and tied down to the underside of the leaf. Caterpillars tend to feed on younger leaves near the ends of branches and feed from the leaf margins toward the center (rather than cutting shelter flaps in the middle of the leaf). The caterpillar uses its shelter as a hide-out as it grows, eating portions of the leaf shelter until it can no longer support the caterpillar. It will also venture out to eat and return for protection but will eventually move on to create a new shelter when it outgrows its current one. Typically, only early caterpillar instar stages (one through four) build shelters. The fifth and final instar does not usually make a shelter but continues to feed on exposed leaves and may use silk to tie leaves together. The caterpillar stage typically lasts about a month after hatching from the egg and up to pupation, and development from egg to adult is around 40 to 45 days. Caterpillars often remain on host plants to pupate. The pupal stage may last about 10 days. Habitat needs: The Kamehameha butterfly is primarily a butterfly of wet landscapes where its caterpillars feed on the main host plant, mamaki (Pipturus albidus), a native understory shrub. The range of host plants used by the butterfly is not entirely known, but it is suspected to feed on a variety of Urticaceae (nettle) plants (Table 3-9). They are tied to both low- and high-elevation habitats that have native nettles. Unlike their caterpillars, which are host specific, adult Kamehameha butterflies are generalists that feed on nectar from a variety of flowers but also from tree sap that oozes from wounds of koa trees. Adults may be found at rest on the bark of trees or flying along the forest edge or in small clearings. While little has been documented on the forage plants of adult butterflies, they likely feed on nectar from a wide range of blooming plants, including those of the caterpillar host plant (Table 3-9) Top reasons for decline, if known: Like most declining insects in Hawaii, its populations are affected largely as a result of habitat loss and degradation as well as the introduction of nonnative predators and parasitoids. General conservation recommendations: Conserving and restoring populations by reducing or eliminating the presence of nonnative weeds and planting native plants can be an effective conservation strategy to address loss of habitat for this species. Recovery efforts for this species include planting native host plants and protecting immature stages from invasive predators. Restoration that may benefit this species includes establishing habitat with its known native host and nectar plants and protecting habitat from various threats. Reducing threats that diminish or degrade habitat such as feral ungulates can also help promote native habitat restoration. In addition, carefully planned management in suitable habitat areas can help ensure the butterfly is protected from any potential adverse actions. Roadside management recommendations: Intensive mowing and invasive species management may deplete floral resources and potentially disturb host plants occurring along roadsides. Because it has overlapping life stages throughout the year, the Kamehameha butterfly may be sensitive to year-round management that could disturb or

Chapter 3. Imperiled Pollinator Profiles  3-34 remove host or nectar plants (e.g., indiscriminate herbicide application, mowing). Using seed mixes with predominantly native flowering species will support this species better than seed mixes with nonnative plants. Table 3-9. Plants used by Kamehameha butterfly. Species Name  Common Name  Notes  Larval Host Plants  Boehmeria grandis  Akolea/Hawaii false nettle  Native  Neraudia sp.  Ma’aloa  Native; some species  endangered  Pipturus albidus  Mamaki/Waimea pipturus  Native  Touchardia latifolia  Olona  Native  Urera spp.  Opuhe/urera  Native  Nectar Plants  Hibiscadelphus giffardianus  Hau kuahiwi/Kilauea hau  kuahiwi  Native; endangered  Pipturus albidus  Mamaki/Waimea pipturus  Native  3.6.1 References Blackburn, T. 1882. “Descriptions of the larvae of Hawaiian Lepidoptera.” The Entomologist’s Monthly Magazine 19:55–56. Gorelick, G. A., and R. S. Wielgus. 1968. “Notes and observations on the biology and host preferences of Vanessa tameamea (Nymphalidae).” Journal of the Lepidopterists’ Society 22(2):111–114. Haines, W. 2019. Personal communication with Michele Blackburn, Conservation Biologist, the Xerces Society. Research Entomologist, Hawaii Invertebrate Program, Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife University of Hawaii, Center for Conservation Research and Training. January 8–10, 2019. Howarth, F. G., and W. P. Mull. 1992. Hawaiian Insects and Their Kin. University of Hawaii Press, Honolulu. pp. 124–125. NatureServe. 2020. NatureServe Explorer (web application). NatureServe, Arlington, Virginia. Available: https://explorer.natureserve.org/. Accessed: September 2020. Patrick, B., and H. Patrick. 2012. Butterflies of the South Pacific (Vol. 240). Otago University Press. Dunedin, New Zealand. 240 pp.

Chapter 3. Imperiled Pollinator Profiles 3-35 Pratt, L. W., J. R. VanDeMark, and M. Euaparadorn. 2010. Limiting factors of five rare plant species in mesic forests, Hawai’i Volcanoes National Park. 151 pp. Robinson, G. S., P. R. Ackery, I. J. Kitching, G. W. Beccaloni, and L. M. Hernández. 2010. HOSTS - A Database of the World’s Lepidopteran Hostplants. Natural History Museum, London. Available: http://www.nhm.ac.uk/hosts. Accessed: September 2020. Scott, J. A. 1992. The butterflies of North America: A Natural History and Field Guide. Stanford University Press. Stanford, California. 668 pp. Swezey, O. H. 1910. “The Feeding Habits of Hawaiian Lepidoptera.” Proceedings of the Hawaiian Entomological Society 2(3):131–143. Swezey, O. H., and F. X. Williams. 1932. “Some Observations on Forest Insects at the Nauhi Nursery and Vicinity on Hawaii.” Proceedings of the Hawaiian Entomological Society 8(1):179–190. Tabashnik, B. E., W. D. Perreira, J. S. Strazanac, and S. L. Montgomery. 1992. “Population ecology of the Kamehameha butterfly (Lepidoptera: Nymphalidae).” Annals of the Entomological Society of America 85(3):282–285. Wagner, W. L., D. R. Herbst, and S. H. Sohmer. 1999. Manual of the Flowering Plants of Hawai’i, Vols. 1 and 2. University of Hawai’i and Bishop Museum Press. Honolulu, Hawaii. xviii + 1919 pp. Weber, P. 1936. “Notes and Exhibitions.” Proceedings of the Hawaiian Entomological Society 9(2):145. Williams, F. X. 1928. “The Kamehameha Butterfly, Vanessa tammeamea Esch.” Proceedings of the Hawaiian Entomological Society 7(1):164–169.