Patterns of extinction across Hawaiian Lepidoptera offer lessons from a diverse, neglected, and vulnerable endemic fauna

doi:10.21203/rs.3.rs-4993822/v1

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Patterns of extinction across Hawaiian Lepidoptera offer lessons from a diverse, neglected, and vulnerable endemic fauna

https://doi.org/10.21203/rs.3.rs-4993822/v1

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Although insects make up the overwhelming majority of the described life on Earth, virtually nothing is known about the conservation status for all but the most charismatic taxa (i.e., butterflies, dragonflies) and many are at risk of imminent extinction. Given that island faunas are particularly vulnerable to extinction, we examined the conservation status of all 935 described species of native Lepidoptera in Hawaiʻi as a model to better understand levels of threat in poorly-known groups and how their rates of extinction might warrant actions to conserve them and inform future management. Using 100- and 50-year cut-offs for the time since each species was last recorded, we consider 269 species (28.8%) to be “presumed extinct,” 85 species (9.1%) to be “possibly extinct,” and 581 species (62.1%) to be “extant” or “likely extant,” giving an extinction rate of 37.9%. Baseline data such as these are crucial in establishing which taxa persist and which lineages may be most vulnerable to extinction in the immediate future. Continued field surveys and identification of museum specimens are critical to more precisely quantify extinction rates, focusing on taxa requiring the most immediate conservation action, and may result in the rediscovery of some “extinct” species. We discuss discrepancies in extinction rates across taxa and between islands as well as summarizing major threats and obstacles to protecting the Hawaiian Lepidoptera fauna going forward. Conservation specifically geared towards Hawaiian Lepidoptera is almost non-existent, which is inconsistent with the extreme rarity and high rates of extinction across many lineages.

insects

extirpation

moths

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The native flora and fauna of the Hawaiian Islands suffer the highest rates of extinctions globally, so much so that Hawaiʻi is often dubbed “the extinction capital of the world” (Eyre 2000). Extinction rates for its birds, land snails, and plants are well-documented (Régnier et al. 2015, Paxton et al. 2018, Rønsted et al. 2022) and the causes for their declines are generally well-understood. However, little is known about the extinction rates for native Hawaiian insects, although their decline has long been noticed and forewarned (Perkins 1913, Zimmerman 1948, Dunn 2005). Research, funding, and public support for insect conservation have been marginal to non-existent over the past ~130 years (Howarth and Gagné 2012), resulting in limited data being available for virtually all species, even some that are federally listed (e.g., Elliott et al. 2021). Given very limited taxonomic, biological, and distributional data, compounded with significant diversity (>6,000 described native species), it is difficult to fully assess the conservation status of all Hawaiian insect species. Similar rapid, broad evaluations of entire insect orders have been undertaken in New Zealand (e.g., Andrew et al. 2012; Leschen et al. 2012; Hoare et al. 2015), demonstrating that it is possible, despite incomplete biological and distributional information. Appraising the Hawaiian insect fauna with similar criteria (see Hitchmough et al. 2007) would serve as a badly needed global model for insect conservation assessments going forward since the Hawaiian fauna is nearly entirely endemic and relatively restricted geographically, often to just small fractions of individual islands, thus simplifying range wide evaluations. Relative assessments in Hawaiʻi like this can offer results that might be relevant to other regions which are either more diverse, difficult to survey, or otherwise hard to assess.

The only attempt to quantify the conservation status of the native Hawaiian insect fauna occurred in the 1980’s (Gagné 1982, Gagné and Christensen 1985), when just 27 species of Hawaiian Lepidoptera were evaluated for potential inclusion under the Endangered Species Act (ESA). The USFWS (1984) considered an additional 13 Hawaiian Lepidoptera species for listing, but in 1996 removed 19 of the 40 due to “persuasive evidence of extinction,” another 19 due to lack of biological or distributional information (“data deficient”), and one because it was found to be more abundant and widespread than previously believed (USFWS 1996), finally leaving only Manduca blackburni to be listed as endangered in 2000. The data used for these determinations was so sparse that of the 19 species considered to be extinct, Haines et al. (2004) reported the rediscovery of five species of Omiodes and Heddle et al. (2000) reported the rediscovery of Tinostoma smaragditis, although the latter has not been seen since 1998 and some of the former are likely still critically endangered. This suggests that the USFWS declaration of extinction was hasty for the rare moths that were rediscovered, though USFWS never reevaluated its mistakes. Even worse, no subsequent effort was made by USFWS to re-evaluate or conserve any of the non-listed species. Austin and Rubinoff (2023b) assessed the conservation status of native Tortricidae, reporting the rediscovery of three species not seen in >90 years, but considered 21 species as “presumed extinct” (32% of the described Hawaiian tortricid fauna). Presently, the USWFS lists 28 species of Hawaiian insect as “endangered” (35% of the United States’ total endangered insect fauna), including 14 species of fruit flies (Drosophila), 7 species of yellow-faced bees (Hylaeus), and 6 species of damselflies (Megalagrion), but only one moth, the aforementioned Manduca blackburni. The true number of Hawaiian insects needing conservation attention and protection is certainly much higher though there is no ongoing plan to collect the data needed to assess unlisted species.

These trends also hold true on a global scale. As of 2023, The International Union for Conservation of Nature (IUCN) Red List, the leading authority on global biodiversity conservation and extinction, evaluated 60 species of insects as extinct (IUCN 2023). Of these, 26 (43%) are from Hawaiʻi. For Lepidoptera, the numbers are even more alarming: 11 of the 21 (52%) IUCN-listed extinct species and 22 of the 57 (39%) critically endangered species are from Hawaiʻi; an additional 3 species are considered “endangered” and 9 are “vulnerable.” Still, this represents only a fraction of the total endemic Lepidoptera diversity of Hawaiʻi. The remaining 894 species (95%) would be considered “data deficient” under IUCN criteria and are of unknown conservation status. At this rate, and given that the Hawaiian insect fauna is only ~50% described (Hembry et al. 2021), it would take 1,600 years to assess just the endemic Lepidoptera species. There is evidence that poorly known insect species are in fact the most vulnerable to extinction (Boyle et al. 2024), making current inaction even harder to justify. As global extinctions continue to accelerate at an ever-alarming rate (Ceballos et al. 2015, Urban 2015), most of these species will disappear before they are ever discovered, and many of the described species will be gone without any conservation action being taken. It is more critical than ever to evaluate the conservation status and extinction risk for as much of the planet’s threatened insect fauna as possible. We assess the Hawaiian insect fauna as a model for future work and to demonstrate both the utility and urgency of such evaluations in precipitating conservation actions.

We visited the following museum collections to record the most recently collected specimen of each species: the Bernice Pauahi Bishop Museum (BPBM), the Hawaiʻi Department of Agriculture (HDOA) collection, the Natural History Museum, London (NHMUK), the University of Hawaiʻi Insect Museum (UHIM), and the U.S. National Museum of Natural History (USNM). These museums constitute the repositories for all major historical and modern surveys of the Hawaiian Lepidoptera fauna, including all holotypes. Because of a lack of taxonomic expertise, many specimens were unidentified so, with the exception of the mega-diverse genus Hyposmocoma, we also attempted to identify all the unidentified material in these collections to species whenever possible. In some instances, genitalia dissections were made in order to obtain a positive identification. All identifications were based on comparisons with original descriptions, figures, and illustrations; photographs or personal examination of type material (primarily in NHMUK, BPBM, and UHIM); and figures in Zimmerman’s (1958a, b, 1978) monumental Insects of Hawaii series. In all but a few instances in which taxonomic monographs have been published or convincing molecular evidence is available, the range of species was considered to be limited to the island from which they were described (“type locality” in Table S1), as most studies to date have revealed few multi-island species distributions in insects (Hembry et al. 2021). Identifications were sometimes hampered by the poor condition of type material or vague labels on type material (i.e., “Sandwich Islands”). In these cases, we used our best judgment in identifying specimens and the type locality is listed as “‘Hawaiʻi’” in Table S1. Species described from the Hawaiian Islands and only known from there, but likely to be adventive (see Zimmerman 1978, Nishida 2002, Austin and Rubinoff 2022), are excluded from our list. A few distinctive species without more recent voucher specimens were identified based on photos shared online at iNaturalist and cited as such.

We also reviewed all relevant literature (e.g., Giffin and Rowe 2007; Medeiros 2009, Kobayashi et al. 2021), including unpublished Ph.D. dissertations (e.g., Heddle 2003, Oboyski 2011). When voucher material from these studies was available, we confirmed and/or updated previous authors’ identifications and cite these under the museum acronyms in which vouchers are deposited rather than the publication itself. Supplementary data was also sought from taxonomic experts (see Acknowledgments).

We then reviewed data from our own field work (2002–2024), which targeted all species of native Lepidoptera on Midway Atoll, Laysan, Nihoa, Kauaʻi, Oʻahu, Molokaʻi, Lānaʻi, Kahoʻolawe, Maui, and Hawaiʻi island (Fig. 1). These represent over 900 unique collecting events, including from nearly all of the type localities of the species collected by R.C.L. Perkins (see Evenhuis 2007) and O.H. Swezey, although many now are severely degraded by invasive species. Ultraviolet light traps and light sheets were the primary modes of collection, following standard Lepidoptera collection methods (Montgomery et al. 2021), supplemented by rearing of larvae and opportunistic netting of adult moths. All specimens from our field work are retained in the University of Hawaiʻi Insect Museum (UHIM) for future study.

We modified the criteria used by Austin and Rubinoff (2023b) to classify extinctions into one of two categories: a 100-year cut-off (pre-1924, “presumed extinct”) and a 50-year (1924–1973, “possibly extinct”). We use a relatively conservative (optimistic) approach in that species for which identified specimens are available since 1974 are considered “extant or likely extant.” Throughout the text, we cite these numbers as “[presumed extinct / possibly extinct / extant or likely extant].” This final category should not be conflated with “secure” or “stable,” as many of these species have only been seen once in decades and may be just as threatened as those in the former two categories. For the sake of simplicity, subspecies were excluded, as there are exceedingly few described subspecies of Hawaiian Lepidoptera and many of them are of uncertain taxonomic standing (i.e., some are likely deserving of full species status and others are likely synonyms). Known, but undescribed species (e.g., those listed in Zimmerman 1978) were also excluded.

Figures were created with cartopy v0.23.0, matplotlib v3.9.1, pandas v2.2.2, and seaborn v0.13.2, and edited further with Affinity Designer v1.10.8.

As of 2024, there are 935 species of native Lepidoptera described from the Hawaiian Islands (Fig. 2B; Table S1). 269 (28.8%) have not been collected in over a century (primarily only known from the type specimen/series), while an additional 85 (9.1%) have not been collected since 1973, a 50-year cutoff (Table S1). Excluding Kahoʻolawe and the Northwestern Hawaiian Islands (discussed below), rates of presumed extinctions (≥ 100 years since last record) are roughly similar between islands, varying from 23.8% (Oʻahu) to 41.7% (Lānaʻi) (Fig. 2C, Table S3). Rates of possible extinctions (50–99 years since last record) are also roughly similar, with Molokaʻi (5.3%) and Oʻahu (13.3%) having the lowest and highest respective percentages of their fauna fitting into this category. Altogether, the percentage of the native fauna that we classify as either “presumed extinct” or “possibly extinct” is 37.9%.

When the megadiverse endemic genus Hyposmocoma (360 species) is excluded (Table S2), rates of extinction fall dramatically for total species presumed extinct: from 28.8–16.0%. The percentage of species believed to be extant or likely extant rises from 62.1–74.4% without Hyposmocoma. This is true for every island except some of the Northwestern Hawaiian Islands (e.g. Laysan), where Hyposmocoma are some of the only extant native Lepidoptera, the rest having been driven to extinction by human-mediated disturbances more than a century ago. Extinction rates within Hyposmocoma (177 [49.1%] / 30 [8.3%] / 153 [42.5%]) vary dramatically between subgenera: the percentage of species extant or likely to be extant for subgenus Hyposmocoma (83 [40.5%] / 8 [3.9%] / 114 [55.6%]) is more than twice as high as that of subgenus Euperissus (94 [60.6%] / 22 [14.2%] / 39 [25.1%]).

Apart from Hyposmocoma, 31 other genera have at least one Hawaiian species which has not been seen in > 50 years (Fig. 2A, D), ranging from just 9.1% of Eupithecia to 100% of the of the species in the genera Acrapex (Noctuidae), Hypena (Erebidae), and the monotypic Paraphasis (Tortricidae) and Tritocleis (Geometridae). 34 genera appear to be entirely extant, with all described Hawaiian species having been collected within the last 50 years (Fig. 2A, D). All of these are relatively small genera, with up to only 7 species.

Ongoing surveys have been productive and at least four species of Lepidoptera presumed extinct by Austin and Rubinoff (2023b) are rediscovered here: Eccoptocera osteomelesana, Nuritamburia phyllanthana, Pararrhaptica capucina, and “Spheterista” urerana (Table S1). Countless other species reported here from collections in the past 22 years likely represent additional rediscoveries of species not seen in over a century, but with no published baseline data to compare to, it is difficult to make such conclusions.

The discrepancies between extinction rates on islands may reflect recent collecting bias between islands rather than true differences in rates. It is highly unlikely that Oʻahu, with its relatively low remaining amount of intact, native forest, has suffered the lowest percentage of species extinction of Lepidoptera compared to all the other main islands. Instead, we suggest that Oʻahu has benefited from much more intensive entomological surveys over the past century and as a result its fauna is much more well-documented than the other islands. Oʻahu’s extinction rate is probably the most accurate as large numbers of surveys over decades often specifically targeting these “lost” species have so far failed to find them (Austin and Rubinoff 2023b). Additional surveys on islands with seemingly high extinction rates (e.g., Lānaʻi, Molokaʻi) may result in additional rediscoveries of “extinct” species and bring their extinction rate lower than currently recognized. However, ecological damage on Lānaʻi is more severe than Oʻahu, and it is possible that extinction rates on smaller, lower islands may be higher than that of Oʻahu. Even if lower extinction rates end up being verified for other islands, they are still among the highest recorded anywhere in the world.

Detailed understanding of the ecology of species that are apparently rare will be critical in understanding the broader patterns of extinction across Hawaiian insects. Some groups provide important examples of how or why some species are particularly vulnerable. The subgenus Euperissus, which primarily have larvae that feed naked (i.e., without cases), are generally faring much more poorly than subgenus Hyposmocoma, all of which make protective silk cases as larvae. This may be partially due to the close relationship some groups of Euperissus have with their host plants, many of which are now rare or endangered. It may also reflect the limited attention taxonomists have paid to Euperissus in the past century: no new species have been described in Euperissus since 1935, whereas 50 species in subgenus Hyposmocoma have been described since then, including 49 in just the past 20 years. Some groups of Euperissus can still be encountered fairly frequently, even abundantly. However, this may be a consequence of their protected feeding strategies: many of these more resilient Euperissus groups feed internally in bark, fruits, and seeds, and at least some are known to make cases. Many of the naked wood-feeding Euperissus species are now largely absent from otherwise suitable habitat and may have been outcompeted by termites and invasive moths (e.g., Opogona spp.) as well as preyed upon by ants, making the extinction estimates unfortunately accurate. Consideration of this ecological information is important in understanding the extinctions patterns and engaging in more effective conservation actions.

Extinction rates between groups of native moths appear to vary dramatically, with some extremely diverse genera having largely avoided extirpation while other genera are entirely extinct. The monotypic endemic genus Tritocleis (Geometridae) Meyrick is only known from the holotype collected in 1896. At least one other genus (Hypena, Erebidae; 3 presumed extinct / 2 possibly extinct / 0 extant or likely extant) appears to have become entirely extinct as well, with none of the five native species having been seen since 1941. This may be the result of rodent introductions on the Northwestern Hawaiian Islands (Gagné 1982); deliberately released, non-specific biological controls aimed at Lepidoptera or other insects (Funasaki et al. 1988); and the accidental arrivals of non-native parasitoids (Henneman and Memmott 2001). Other species may be declining because of the loss or severe decline of their host plants, such as Acrolepia (leaf-miners on Nothocestrum; 2 / 0 / 2), Mapsidius (stem- and leaf-feeders on Charpentiera and Chenopodium; 1 / 2 / 2), and Philodoria (specialist leaf-miners on various native plants; 5 / 5 / 35). Some may have been lost to hybridization with invasive congeners (Helicoverpa, 1 / 2 / 1; Mythimna, 0 / 0 / 3). Some genera that appear to be hard-hit (e.g., Batrachedrodes, Paralopostega) may actually be more common than currently known, but uncommon in collections due to their small size, specialized feeding habits, or difficult-to-rear larvae. Still others are declining despite no immediately obvious factor such as Merimnetra (leaf-miners, stem-gallers, and fruit-feeders in native Rubiaceae; 7 / 5 / 6) and Pararrhaptica (leaf-rollers on native Primulaceae; 3 / 5 / 13).

Coastal species have suffered the highest rates of extinction, with numbers and abundance of native species increasing with elevation and percent native forest coverage. Only a handful of native species can be encountered with any regularity at the lowest elevations (e.g., Tamsica, Plutella, some Hyposmocoma). Formerly abundant coastal and lowland species such as Agrotis crinigera were apparently driven to extinction by the arrival of invasive predators such as big-headed ants and the early introductions of poorly-guided attempts at biological control (mynah birds, cattle egrets, and various polyphagous parasitoid wasps and flies; Funasaki et al. 1988). Many of these formerly common native species have been largely replaced by a small set of polyphagous exotic species.

On the other hand, the few genera of native moths that appear to be surviving, even thriving, offer important management information as well. This is likely the result of their hosts remaining common even in highly disturbed and non-native forests, the ability to switch to feeding on non-native plants, and/or the predator protection derived from their specialized cases or behaviors of larvae. For example, several species of Eudonia (4 / 2 / 56) and Mestolobes (8 / 4 / 22) can often be seeing flying at dusk and night in the hundreds to thousands in highly degraded strawberry guava (Psidium cattleyanum) or ironwood (‎Casuarina equisetifolia) thickets where the larvae likely feed on the indigenous mosses that persist at the base of these trees.

Despite these glimmers of hope, the overall outlook for Hawaiian Lepidoptera is grim: though many species can still be found, exceedingly few are common or abundant. In many areas, non-native moths outnumber native species by an enormous margin. Extinction rates, high as they are, show no sign of abating. Hawaii’s rate of loss dramatically outpaces that of similar island ecosystems: New Zealand, with over 1,800 described native moth and butterfly species (roughly twice as many as Hawaiʻi), has zero species that can be “confidently considered extinct,” only 49 that are “threatened,” and 69 “at risk” (Stringer et al. 2012). The primary major threats to native Lepidoptera in Hawaiʻi likely mirror those of native insects more broadly, namely the loss of habitat and host plants due to displacement by invasive plants (Wagner and Van Driesche 2010, Medeiros et al. 2017), predation by non-native arthropods, especially ants and wasps (Gambino 1992, Krushelnycky and Gillespie, 2010), development (Polhemus 1993, Magnacca and King 2013), and climate change (Hobbelen et al. 2013, Hembry et al. 2021). The arrival of new invasive species in Hawaiʻi, both plants and animals, continues nearly unabated year after year (Matsunaga et al. 2019, Austin and Rubinoff 2022, 2023a, 2024). Whereas Hawaiian birds, plants, and charismatic marine life receive substantial conservation funding targeted to better understand species’ biology, distribution, and threats, only a tiny fraction goes towards insects (Medeiros et al. 2013), despite Hawaiian insects being more than twice as diverse as the aforementioned groups combined (even excluding undescribed insect species). Compounding this is the difficulty in accessing type material (most of which is stored in NHMUK in London), only a fraction of which is digitized and available for study. Researchers wishing to study and protect the Hawaiian fauna must fly halfway around the world to see the only identified specimens of a species that may now be extinct. Repatriation of this material to Hawaiʻi would not only benefit taxonomists working in Hawaiʻi, but also greatly accelerate the conservation of all Hawaiian insects.

Our approach is not an overestimate of the true number of extinctions in the Hawaiian Lepidoptera fauna, because in the past 50 years commensurate extinctions have been rampant in better known groups like birds and plants. Regardless, our data provides an important baseline by which to compare future conservation assessments and hopefully to guide actions. It is not yet clear how many additional “presumed extinct” species would likely warrant removal from a “data deficient” category into the “extinct” category, as additional surveys are still badly needed on all of the Hawaiian Islands. Hawaiian insect conservation lags behind all other groups in terms of support and knowledge, implying that management agencies have determined that they are under less threat, or are less important. Here we clearly show that a lack of funding to collect data, rather than any resilience is to blame for pervasive inaction. Conserving insects for their inherent value has apparently not been a compelling reason to allocate resources or protect habitat; but if investment is not made soon, critical numbers of insect species may go extinct, leaving irreplaceable gaps in ecosystems.

Hawaiʻi as a case study is by no means unusual. The Hawaiian Lepidoptera fauna exemplify unfortunate global trends, but on a much more acute and easily measurable scale. The most pressing issue for most threatened insects worldwide is the lack of basic taxonomic and biological information. Insect species are going extinct faster than they can be described (Medeiros et al. 2013, Rubinoff 2017) and only a small percentage of taxa have received any taxonomic attention since their original descriptions decades or centuries ago (Leandro et al. 2017, Wheeler and Miller 2017, Löbl et al. 2023). Host plant records and other biological information are among the most important data for conserving insects, but are unfortunately lacking for the vast majority of species (Bossart and Carlton 2002, Cardoso et al. 2011). Having a baseline estimate for the total numbers of extinct species is a crucial first step in establishing urgency and protecting what species remain. Similar published “extinction databases” would be greatly beneficial for the conservation of insect taxa elsewhere in the world, especially on islands, where insects are particularly vulnerable to extinction (Howarth and Ramsay 1991, New 2008).

Author Contribution

K.A. and D.R. conceived and wrote the manuscript. K.A. examined specimens, made identifications, and prepared figures. All authors reviewed the manuscript.

Acknowledgement

We thank Keith Arakaki (BPBM), Jim Boone (BPBM), John Brown (USNM), Neal Evenhuis (BPBM), Jeremy Frank (BPBM), David Lees (NHMUK), and Janis Matsunaga (HDOA) for collection access. We thank Ane Bakutis (PEPP, Moloka‘i), Christopher Balzotti (TNC, Hawai‘i), Keahi Bustamente (Maui DOFAW), Lance DeSilva (Maui DOFAW), Kerri Fay (TNC, Maui), Deena Gary (ANRPO), Russell Kallstrom (TNC, Moloka‘i), Cynthia King (Hawai‘i State DLNR), Peter Landon (Maui NARS), Sheri Mann (Kaua‘i DOFAW), Joe and Kristen Souza (Nani ‘Ekolu), Mapuana O’Sullivan (Kaua‘i DOFAW), Ryan Peralta (O‘ahu DOFAW), Adam Radford (TNC, Maui), David Tsuchiya (Kaua‘i DOSP), Haleakalā National Park, and Hawai‘i Volcanoes National Park for providing collection permits, site access, and logistics. We thank Marci Brimhall, Emily Brines, Monroe Bryce, Keahi Bustamente, Ryan Caesar, Ryan Chang, Saxony Charlot, Kristen Coelho, Dan Cuthrell, Camiel Doorenweerd, Jesse Eiben, Kevin Faccenda, Conrad Gillett, Will Haines, Kalli Harshmann, Alyssa Kaneshiro, Caroline Kealoha, Cynthia King, Luc Leblanc, Karl Magnacca, Jenna Masters, Steve Montgomery, Scott Nelson, Dan Nitta, Kurt Ongman, Zach Pezzillo, Spencer Pote, Kawaila Purdy, Patrick Schmitz, Melissa Simon, Malia Staab, Hubert Szczygieł, Andersonn Prestes, Adam Radford, Brad Reil, Mike San Jose, Sandra Schachat, Forest and Kim Starr, Graham Talaber, Julien Ueda, Brendan Wang, and Jon Winchester for help with field work. We thank Francis “Jay” Joy, Janis Matsunaga, and Matt Medeiros for providing data on species. We thank Camiel Doorenweerd for his assistance in creating figures. This study was supported, in part, by USDA Farm Bill project #3.0227.05: “Diagnostic Tools to Identify Exotic Tortricidae that Threaten U.S. Agriculture.” Additional funding was provided by The College of Tropical Agriculture and Human Resources (CTAHR), University of Hawaiʻi at Mānoa, USDA Cooperative State Research, Education and Extension (CSREES), Grant/Award Number: HAW00942-H; the D. Elmo Hardy Student Assistance Endowment; the Edward M. Ehrhorn Entomology Scholarship.

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Patterns of extinction across Hawaiian Lepidoptera offer lessons from a diverse, neglected, and vulnerable endemic fauna

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