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Hawaiian hoary bat

The Hawaiian hoary bat (Lasiurus cinereus semotus), known as ʻōpeʻapeʻa in Hawaiian, is an endangered subspecies of the hoary bat (Lasiurus cinereus) and the sole extant native terrestrial mammal in the Hawaiian archipelago, having colonized the islands via a single long-distance dispersal event from North America.^[1]^[2] It inhabits forested habitats across islands including Hawaiʻi, Maui, and Kauaʻi, where it roosts solitarily in trees and forages nocturnally on insects using echolocation.^[3]^[4] Listed as endangered under the U.S. Endangered Species Act since 1970, the species faces threats from habitat degradation, collisions with wind turbines, and low reproductive rates, with no reliable population estimates available despite acoustic and genetic monitoring efforts.^[5]^[6] Recent genomic analyses reveal moderate genetic diversity but signs of historical bottlenecks and ongoing decline, underscoring its isolation and vulnerability as the most geographically remote bat population on Earth.^[7]^[8]

Taxonomy and evolutionary history

Classification and nomenclature

The Hawaiian hoary bat belongs to the order Chiroptera, suborder Yangochiroptera, superfamily Vespertilionoidea, family Vespertilionidae, subfamily Vespertilioninae, and tribe Lasiurini.^[9] It was first described scientifically by Harrison Allen in 1890 as Atalapha semota based on specimens from the Hawaiian Islands, published in the Proceedings of the United States National Museum. This name was later revised to Lasiurus cinereus semotus, recognizing it as a subspecies of the mainland hoary bat (Lasiurus cinereus), which ranges across the Americas.^[10] Molecular analyses, particularly Baird et al.'s 2015 phylogenetic revision of the Lasiurini tribe using mitochondrial and nuclear DNA, revealed deep genetic divergence between Hawaiian populations and continental L. cinereus, with the Hawaiian lineage forming a distinct clade separated by millions of years of isolation.^[11] This led to elevation of the Hawaiian form to full species status as Lasiurus semotus in subsequent taxonomic updates, including those by NatureServe and the Mammal Diversity Database, reflecting its endemic status and lack of gene flow with North American populations.^[12]^[9] However, the U.S. Fish and Wildlife Service retains the trinomial Lasiurus cinereus semotus in Endangered Species Act listings, prioritizing historical nomenclature for regulatory continuity despite the genetic evidence.^[1] The genus name Lasiurus derives from Greek roots lasios (hairy) and oura (tail), alluding to the furred tail membrane typical of the genus.^[13] The species epithet semotus is Latin for "remote" or "set apart," denoting the population's extreme geographic isolation in Hawaii.^[2] In the Hawaiian language, it is called ʻōpeʻapeʻa, a name evoking its folded wings resembling frayed tapa cloth, and it was designated Hawaii's official state mammal on April 21, 2015.^[14] Common English names include Hawaiian hoary bat, reflecting the hoary (frosted) appearance of its fur from white-tipped hairs.^[15]

Phylogenetic origins and colonization of Hawaii

The Hawaiian hoary bat (Lasiurus cinereus semotus, potentially elevated to full species status L. semotus based on genetic divergence) belongs to the vespertilionid tribe Lasiurini, with its closest continental relatives in the North American hoary bat (L. cinereus cinereus).^[16] Phylogenetic analyses using mitochondrial DNA (mtDNA) and nuclear markers reveal that Hawaiian populations form distinct clades, indicating a basal divergence from North American lineages rather than simple subspeciation.^[7] Single nucleotide polymorphism (SNP) data from whole-genome sequencing further confirm shared ancestry with continental L. cinereus, but with significant genetic differentiation, including reduced heterozygosity and unique alleles consistent with isolation and genetic drift post-colonization.^[17] Colonization of the Hawaiian archipelago by hoary bats occurred via overwater dispersal from North America, as the islands lack other native terrestrial mammals and bats are capable of long-distance flight aided by fat reserves.^[18] Molecular evidence from cytochrome b mtDNA and microsatellite loci supports at least two independent dispersal events: an ancient founding event establishing a divergent "ancient clade" across islands, followed by a more recent colonization introducing lineages closely allied with modern continental L. c. cinereus.^[19] These events likely involved trans-Pacific flights of approximately 4,000 km, feasible under tailwind conditions as modeled by flight simulation tools like FLIGHT, which demonstrate sufficient energy stores in pregnant or fat-laden individuals for nonstop travel from the California coast.^[18] No evidence supports human-mediated introduction, as bats predate Polynesian arrival around 1,000 years ago, and fossil records or indigenous oral histories align with pre-human presence.^[19] Post-colonization, genetic structure shows island-specific variation, with lower effective population sizes (e.g., Ne estimates of 1,000–10,000 across Hawaii) reflecting bottlenecks and limited gene flow, though admixture from multiple founders mitigated inbreeding.^[20] This history underscores the bat's remarkable vagility within Vespertilionidae, contrasting with flightless mammals' absence in Hawaii, and informs conservation by highlighting vulnerability to further isolation-induced drift.^[7]^[19]

Physical characteristics

Morphology and adaptations

The Hawaiian hoary bat (Lasiurus cinereus semotus) is a medium-sized insectivorous bat with a head-body length of approximately 60-70 mm, forearm length of 40-48 mm, and body mass ranging from 14-25 g, notably smaller than its mainland conspecific L. c. cinereus which averages 20-35 g.^[21]^[22] Its fur is dark brown with white or silvery tips, creating a frosted or "hoary" appearance that provides camouflage against tree bark and foliage during daytime roosting.^[23] The ears are short, rounded, and hairy-fringed, with a dark facial mask, while the tail membrane is moderately developed and furred. Wings are long and narrow, with a wingspan of 26-30 cm, featuring high aspect ratio suited for efficient gliding and agile maneuvers.^[21]^[24] Morphological adaptations reflect insular evolution, including reduced body size enabling consumption of proportionally larger, hard-bodied insects like beetles despite smaller dentition compared to mainland populations.^[22] Wing parameters, such as increased tip area and length ratios, enhance maneuverability in cluttered Hawaiian forest habitats while preserving long-distance flight capability necessary for oceanic colonization over 4,000 km from North America.^[22]^[2] The bat employs frequency-modulated echolocation calls varying by habitat—longer, multi-harmonic pulses in open areas and shorter, steeper sweeps in cluttered vegetation—to detect prey during aerial hawking.^[25] Fur coloration and solitary foliage-roosting behavior further adapt it for crypsis, minimizing predation risk in predator-scarce but structurally complex island ecosystems.^[23]

Sexual dimorphism and variation

The Hawaiian hoary bat (Lasiurus cinereus semotus) displays sexual dimorphism characterized by greater body size in females compared to males, consistent with patterns observed in many vespertilionid species where larger female mass supports reproductive demands. Adult females average 17.9 g in mass, while males average 14.2 g.^[3] This dimorphism extends to overall body dimensions, with Hawaiian individuals approximately 30-45% smaller than mainland conspecifics (L. c. cinereus), though proportional sex differences in wingspan and forearm length remain similar, typically spanning 25-30 cm across both sexes.^[2]^[26] Morphological variation within sexes is limited, reflecting the subspecies' insular isolation and reduced genetic diversity, with minimal geographic differences across Hawaiian islands in pelage (frosted gray-brown fur with white-tipped hairs) or cranial features.^[2] Females may exhibit slightly greater variability in mass during reproductive seasons due to pregnancy and lactation, but no pronounced intrasexual polymorphisms in dentition, skull shape, or echolocation call structure have been documented.^[27] These traits underscore adaptations for solitary, foliage-roosting arboreal life rather than sexual selection-driven ornamentation.

Distribution and habitat

Historical and current range across islands

The Hawaiian hoary bat (Lasiurus cinereus semotus) was first collected in 1861 and has been documented historically across five of the main Hawaiian Islands: Hawaiʻi, Maui, Oʻahu, Kauaʻi, and Molokaʻi.^[28]^[3] No historical records exist for Lānaʻi, Kahoʻolawe, or Niʻihau, likely due to limited suitable forested habitats on these smaller or more arid islands.^[3] Early accounts, including specimens from the 19th century, confirm presence on these islands, with observations tied to native forests and koa-ohia woodlands.^[4] Current distribution mirrors the historical range, with confirmed detections on Hawaiʻi, Maui, Oʻahu, Kauaʻi, and Molokaʻi, but excludes Niʻihau, Lānaʻi, and Kahoʻolawe.^[14]^[1] Acoustic surveys and visual sightings indicate persistent but low-density populations, particularly on the larger islands of Hawaiʻi and Kauaʻi, where bats are more commonly detected.^[12] On Oʻahu, recent observations include Waimea Valley, the Koʻolau and Waiʻanae ranges, Waikīkī, Ford Island, North Shore, and Pearl Harbor National Wildlife Refuge.^[1] Molokaʻi records are sporadic, reflecting overall rarity outside core areas. Population declines since the mid-20th century have reduced encounter rates across islands, though the species remains extant without evidence of island-specific extirpations.^[4]^[3]

Habitat preferences and elevational distribution

The Hawaiian hoary bat (Lasiurus cinereus semotus) utilizes diverse habitats including native and non-native forests, forest edges, and open areas across the main Hawaiian Islands. It forages primarily in open fields, along cluttered forest margins, over lava flows, streams, ponds, and the ocean, typically 1 to over 150 meters above ground or water, preying on nocturnal insects such as moths and beetles.^[3] Roosting is solitary in tree foliage, favoring trees taller than 4.6 meters (15 feet), with documented use of species like Metrosideros polymorpha (ʻōhiʻa), Acacia koa, and non-natives such as eucalyptus and Java plum, though no definitive preference for native vegetation over non-native has been established.^[3]^[29] Occupancy models from acoustic surveys on Hawaiʻi Island reveal a significant positive association with mature forest cover but no strong correlation with dominance by koa (Acacia koa) or net primary productivity.^[30] Elevational distribution spans from sea level to over 4,000 meters, with the majority of observations occurring below 2,286 meters, though records extend to 4,023 meters on Hawaiʻi Island.^[3] The species undertakes seasonal altitudinal migration, concentrating in coastal lowlands during the breeding season (May–October) for pupping and dispersing to interior highlands up to 3,600 meters during the non-breeding winter period (November–March).^[30]^[29] This pattern aligns with higher arrival rates and lower departure rates at high elevations, potentially driven by prey abundance or cooler conditions prompting torpor-like states, and individuals can traverse elevations from sea level to above the cloud line within a single night.^[30]^[3] Multi-year occupancy data indicate overall positive associations with elevation (odds ratio 1.533–1.874), with mean occupancy rates of 0.63 across surveyed sites.^[30] Island-specific variations exist, with prominent winter use of high-elevation caves on Mauna Loa for foraging, but such behaviors require further confirmation across islands.^[3]

Ecology and behavior

Foraging and diet

The Hawaiian hoary bat (Lasiurus cinereus semotus) is strictly insectivorous, with moths (Lepidoptera) comprising the dominant component of its diet, including taxa from at least 18 families such as Noctuidae and Geometridae.^[31] Other consumed orders include Coleoptera (beetles), Diptera (flies), Hemiptera, and Blattodea (cockroaches), encompassing both native Hawaiian insects and non-native species, some of which are agricultural pests.^[31] ^[15] Genetic analysis of fecal samples confirms this composition, with no evidence of significant predation on caterpillars or vegetation despite occasional observations in mainland congeners.^[31] Foraging occurs primarily through aerial hawking, facilitated by echolocation calls with a primary frequency of approximately 30 kHz, which are higher-pitched and more variable than those of mainland hoary bats.^[15] Bats exhibit selectivity in prey choice and target insects in open habitats such as forest edges, gaps, clearings, above the canopy, along roads, and over water bodies, spending more time in woodlands than grasslands.^[3] ^[32] On the island of Hawai'i, radiotracking of 28 individuals revealed moderately large foraging ranges compared to other insectivorous bats, with documented nightly movements supporting year-round activity, though seasonal shifts to higher elevations occur during winter.^[33] The Hawaiian hoary bat (Lasiurus cinereus semotus) is a foliage-roosting species that typically roosts solitarily in tree canopies during the day, selecting perches in the upper third of trees for thermoregulation and protection from predators.^[3] Roost trees include both native species like Metrosideros polymorpha (‘ōhi‘a) and non-native ones such as eucalyptus (Eucalyptus spp.) and paperbark (Melaleuca quinquenervia), often with diameters at breast height exceeding 225 cm, canopy heights over 20 m, and located at forest edges or in areas with slopes greater than 15°.^[34] Bats roost at heights ranging from 1 to 26 m above ground, rarely using lava tubes, rock crevices, or artificial structures.^[35] Some individuals exhibit roost fidelity by returning to the same tree or area nightly over periods of at least two weeks, while others switch roosts frequently, potentially varying seasonally.^[3] Females form temporary maternity roosts with their pups until the young reach 6–7 weeks of age, after which pups become independent and roost solitarily.^[35] The cryptic nature of these foliage roosts, combined with the bat's low population density, complicates detection and monitoring efforts.^[3] Socially, the Hawaiian hoary bat is predominantly solitary, with no documented large aggregations or complex social structures beyond mother-pup pairs.^[35] Observations indicate dispersed distributions and minimal inter-individual interactions during non-reproductive periods, reflecting adaptations to its island environment and foliage-roosting lifestyle.^[3] Limited data suggest potential social calls or associations during migration or foraging, but these remain unconfirmed for the subspecies and are inferred cautiously from mainland hoary bat behaviors.^[34]

Reproduction and life history

The Hawaiian hoary bat follows a defined seasonal reproductive cycle, with distinct phases including pre-pregnancy from mid-December to March, pregnancy from April to mid-June, lactation from mid-June to mid-August, and post-lactation from mid-August to mid-December.^[36] Breeding activity, including mating, primarily occurs from September to December.^[3] Due to the bat's solitary and cryptic nature, detailed observations of mating behaviors remain limited, though acoustic monitoring indicates elevated activity during pregnancy and lactation phases, particularly at lower elevations.^[36] Females typically give birth to one or two pups in mid-June, consistent with patterns observed in the mainland hoary bat subspecies.^[5]^[3] Pups are born non-volant and remain dependent on their mothers, who form temporary maternity roosts in foliage, often in urban or forested trees where females have been observed with one or two young.^[5] Young achieve flight capability by late August or early September, marking the transition to independence amid peak fledging activity that coincides with population occupancy maxima.^[5]^[36] Life history traits reflect adaptation to an insular environment, with bats concentrating in coastal lowlands during the breeding season (April to August) before shifting to higher elevations in non-breeding periods.^[36] Longevity data are sparse, but mitigation models estimate an average lifespan of around 10 years, though wild survival may be shorter due to threats like habitat loss and predation.^[5] Overall, reproductive success appears tied to mature forest availability, with lactating females showing higher site persistence in suitable habitats.^[36]

Population genetics and dynamics

Genetic diversity and structure

The Hawaiian hoary bat (Lasiurus semotus) exhibits low overall genetic diversity, consistent with its status as a small, isolated island population derived from multiple trans-Pacific colonization events from North American hoary bats (L. cinereus). Analysis of mitochondrial cytochrome oxidase subunit I (COI) sequences from 321 individuals across four main Hawaiian Islands (Hawaiʻi, Maui, Oʻahu, and Kauaʻi) revealed haplotype diversity ranging from 0.000 to 0.567 per island, with nucleotide diversity similarly low at 0.000–0.002, indicating limited variation and potential historical bottlenecks.^[20] ^[8] Nuclear microsatellite loci (18 markers genotyped in 298 individuals) showed higher but still modest heterozygosity (mean observed 0.58–0.64 across islands), with no significant deviation from Hardy-Weinberg equilibrium, suggesting recent population stability rather than inbreeding depression.^[20] Population genetic structure is weak but detectable among islands, with pairwise F_ST values for microsatellites ranging from 0.01 to 0.06, implying ongoing but restricted gene flow, likely via rare long-distance dispersal.^[20] Bayesian clustering analyses (e.g., STRUCTURE) identified two main genetic clusters primarily differentiating Kauaʻi from the other islands, though admixture is common, supporting a model of multiple founding events rather than a single bottlenecked lineage.^[7] Mitochondrial data corroborate this, with shared haplotypes across islands but island-specific variants, and phylogenetic reconstructions placing L. semotus as a distinct clade diverging from continental L. cinereus approximately 0.7–1.3 million years ago, followed by repeated colonizations estimated at 800–2,000 years ago based on genomic sequence data from whole-genome and reduced-representation sequencing.^[7] ^[20] Effective population sizes (N_e) estimated via linkage disequilibrium methods yield contemporary values of 1,227–1,532 for nuclear markers, exceeding census estimates and indicating demographic resilience despite low diversity, while coalescent-based historical N_e from microsatellites suggests long-term stability around 1,000–2,000 individuals pre-human impacts.^[20] These patterns reflect causal dynamics of island biogeography: initial founder effects reduced diversity, but recurrent immigration mitigated severe inbreeding, contrasting with expectations of panmixia in a highly mobile species. Genetic differentiation from mainland populations supports taxonomic elevation to full species status, emphasizing adaptive divergence in isolation.^[7]

Population estimates and trends

No reliable census estimates of the Hawaiian hoary bat (Lasiurus cinereus semotus) population size exist, owing to the species' cryptic nocturnal habits, solitary roosting in tree foliage, and the limitations of detection methods such as acoustic monitoring, which primarily measure activity rather than absolute abundance.^[27]^[4] Genetic analyses provide effective population size (N_e) estimates as proxies for breeding population viability, revealing contemporary island-specific values of approximately 1,265 on Hawaiʻi Island (95% CI: 314–∞), 106 on Maui (55–344), 21 on Oʻahu (12–39), and 91 on Kauaʻi (17–∞), based on microsatellite loci from 298 individuals sampled between 1988 and 2020.^[20] These N_e figures indicate small, fragmented breeding populations, with historical N_e inferred to be substantially larger (e.g., 248–62,162 from microsatellites), suggesting past bottlenecks and reduced genetic diversity (haplotype diversity ranging from 0.125 on Kauaʻi to 0.669 on Maui).^[20] Population viability analyses using matrix models parameterized with expert-elicited survival and reproduction data (adult mortality 35–48%, juvenile 54–66%) project slightly positive intrinsic growth rates (r ≈ 0.003–0.009, λ ≈ 1.003–1.019) under baseline conditions without additional anthropogenic mortality, assuming a hypothetical census size of around 1,000 individuals for sensitivity testing.^[37] However, these models highlight vulnerability, as annual take exceeding 0.5–1% of the population could shift trajectories toward decline or extinction risk over decades.^[37] Acoustic monitoring efforts, including a five-year study (2007–2011) on Hawaiʻi Island with mean occupancy of 0.63 across sites, indicate stable to slightly increasing detection probabilities during the breeding season (June–October), with a Bayesian posterior probability of increase at 0.567.^[30] Subsequent surveys on multiple islands (e.g., 2019–2021 on Marine Corps Base Hawaiʻi, 2022 on Oʻahu) confirm persistent activity tied to forest cover and elevation but yield no island-wide abundance trends due to inconsistent sampling and methodological challenges in converting call detections to density.^[38] Overall, empirical data do not demonstrate ongoing population decline, though low N_e, island isolation (significant F_ST differentiation, e.g., 0.780 between Maui and Hawaiʻi), and unquantified historical reductions underpin the endangered status, with monitoring gaps limiting causal inferences on current dynamics.^[20]^[4]

Threats and anthropogenic impacts

Habitat alteration and invasive species

Habitat alteration has historically and continues to pose a significant threat to the Hawaiian hoary bat (Lasiurus cinereus semotus), primarily through the reduction of native forest cover essential for roosting and foraging. Deforestation in the 19th century, particularly on Oʻahu, drastically decreased tree availability, which serves as a limiting factor for solitary foliage-roosting bats that prefer species like Metrosideros polymorpha (ʻōhiʻa) and other native trees exceeding 5 meters in height.^[3] Contemporary alterations include low-elevation conversion to agriculture and urban development, which fragment woodlands where bat activity is 2.4 times higher than in grasslands, alongside ungulate-induced degradation via rooting that disturbs up to 200 m² of soil per day, promotes erosion, and damages understory vegetation critical for insect prey.^[27]^[6]^[39] Timber harvesting and tree trimming above 4.6 meters during the June–September pupping season further exacerbate risks by destroying non-volant young and disrupting territorial behaviors.^[27] Invasive species compound habitat degradation by altering forest structure, reducing native plant diversity, and indirectly affecting prey availability. Feral pigs (Sus scrofa), an introduced ungulate, not only directly damage roost trees but also facilitate the spread of invasive plants, leading to monotypic stands that diminish biodiversity and foraging efficiency through increased vegetation clutter.^[39] At least 30 invasive plant species, including priority targets like strawberry guava (Psidium cattleianum) and mule's foot fern (Angiopteris evecta), degrade native habitats by outcompeting roost trees and reducing insect habitats, with ongoing ecosystem alteration noted across islands.^[39]^[27] The introduced coqui frog (Eleutherodactylus coqui) competes intensely for aerial insects at elevations below 670 meters, consuming 4,500–56,000 prey items per hectare nightly—primarily moth and beetle larvae that overlap with bat diets—potentially altering foraging patterns where frog densities reach high levels on Hawaiʻi, Maui, Oʻahu, and Kauaʻi.^[27] Additionally, the fungal pathogen causing Rapid ʻŌhiʻa Death (ROD), introduced via Ceratocystis species, threatens key roost trees, further eroding habitat quality amid limited evidence of direct prey competition from other invasive insectivores like ants or yellowjackets.^[39]^[6]

Direct mortality factors

The Hawaiian hoary bat experiences direct mortality primarily from collisions with human-made structures, including wind turbines and barbed wire fences. Collisions with wind turbines have been documented at facilities across the Hawaiian Islands, such as the Kaheawa Wind Farm on Maui, where post-construction monitoring from 2007 onward revealed bat carcasses beneath turbines, indicating barotrauma and direct impact as causes of death.^[5] These incidents are exacerbated by the bat's foraging behavior at heights overlapping turbine rotor-swept zones, with fatality rates varying by site-specific factors like turbine height and cut-in speeds; for instance, studies suggest that raising low wind speed curtailment from 5.5 m/s to 6.5 m/s may not significantly reduce mortality without further empirical validation.^[40] Entanglement in barbed wire fencing represents another significant factor, as bats foraging as low as 1 meter above ground can become snagged on top strands, leading to injury or death; monitoring at sites like Kaheawa Pastures estimated bat mortality rates of up to several individuals per kilometer of fence annually.^[41]^[42] Additional direct mortalities occur from strikes against communication towers and other vertical structures, though less quantified than turbine or fence impacts; historical records note such collisions as a persistent threat since at least the 1990s, potentially linked to the bat's nocturnal flight paths through altered landscapes.^[43] Vehicle collisions have been anecdotally reported but lack systematic data confirming prevalence, unlike the well-monitored structural hazards. Overall, these anthropogenic collisions contribute to elevated adult mortality, estimated at 20-50% annually in some models, though precise attribution to direct factors requires ongoing carcass surveys to distinguish from predation or disease.^[44] Mitigation efforts, such as ultrasonic deterrents, have shown promise in reducing fatalities by up to 78% for hoary bats in mainland trials, but efficacy for the Hawaiian subspecies remains under evaluation in island contexts.^[45]

Pesticides and environmental contaminants

The Hawaiian hoary bat, an insectivorous species that forages primarily on moths, beetles, and other arthropods, faces potential threats from pesticides through both direct toxicity and indirect reduction of prey abundance. Pesticide applications in Hawaiian agriculture and invasive species control may contaminate prey insects, leading to bioaccumulation in bats via the food chain, while herbicides can alter vegetation structure and reduce suitable foraging habitat. Although no studies have documented quantifiable insecticide residues in Hawaiian hoary bat tissues, general bat susceptibility to such chemicals—evidenced by mortality in other insectivorous species from ingestion—suggests plausible risk, particularly in areas with intensive pesticide use. The 1998 U.S. Fish and Wildlife Service Recovery Plan identifies pesticides as a factor potentially exacerbating population declines by limiting food resources or causing direct contamination, though it notes the absence of subspecies-specific data and recommends coordinated assessments with agricultural agencies.^[3]^[46] Specific evaluations of certain pesticides indicate limited acute risk. Citric acid, registered as a minimum-risk pesticide for controlling invasive coqui frogs (Eleutherodactylus spp.), was tested for exposure pathways in a 2014 study using big brown bats (Eptesicus fuscus) as surrogates. Oral doses of 542–759 mg/kg induced emesis, with lethality at 2,811 mg/kg, but field simulations showed bats retain minimal amounts during aerial spraying (0.02 g dry acid on effigies) and avoid voluntary consumption, leading to the conclusion of very low toxic exposure risk for Hawaiian hoary bats, albeit with possible sublethal effects like fur matting from high-volume ground applications. Rodenticide screening of 29 Hawaiian hoary bat samples (9 livers, 20 carcasses) in 2018 detected no confirmed quantifiable residues of common anticoagulants (e.g., difethialone, diphacinone) or bromethalin, contrasting with higher exposure rates in native birds of prey; unconfirmed traces in one sample were deemed potential false positives, underscoring minimal evidence of secondary poisoning despite widespread rodenticide use in Hawaii.^[47]^[48] Broader environmental contaminants, such as heavy metals, remain underexplored for this subspecies, with no targeted residue analyses identified; continental hoary bats (Lasiurus cinereus) show mercury accumulation in some regions, but Hawaiian populations' isolation and diet suggest differing exposure profiles. Overall, while ecological reasoning supports pesticide-related vulnerabilities—given the bat's dependence on uncontaminated insect prey—empirical confirmation of significant impacts is lacking, highlighting data gaps in residue monitoring and prey contamination studies.^[3]

Conservation and management

Legal status and recovery efforts

The Hawaiian hoary bat (Lasiurus semotus, formerly Lasiurus cinereus semotus) has been listed as endangered under the U.S. Endangered Species Act (ESA) since October 13, 1970, due to population declines and insufficient data on its status at the time of listing.^[3] It is also listed as endangered under Hawaii state law and designated as the state land mammal in 2015.^[29] In November 2021, the U.S. Fish and Wildlife Service (USFWS) proposed downlisting the subspecies to threatened status, citing improved knowledge of its distribution and resilience to some threats, but this proposal was not finalized, and the bat remains federally endangered as of 2025.^[49] ^[14] The 1998 USFWS Recovery Plan for the Hawaiian hoary bat outlines specific recovery criteria, including maintaining self-sustaining populations on at least three islands, protecting essential habitat, and reducing threats through research and management actions such as invasive species control and habitat restoration.^[3] Recovery efforts emphasize monitoring via acoustic surveys and camera traps to estimate population trends, with ongoing USGS-led studies documenting its island-wide distribution and habitat use to inform conservation priorities.^[4] Key initiatives include habitat conservation plans for wind energy developments, which require mitigation such as restoring 40 acres of native forest per estimated bat pair impacted by turbine collisions, as recommended by the Hawaii Endangered Species Recovery Committee (ESRC).^[50] Recent guidance from 2024 updates these measures, prioritizing data collection on population status and habitat needs to support recovery.^[51] Federal and state agencies collaborate on threat mitigation, including guidelines to minimize bat mortality at wind facilities through curtailment during high-risk periods and enhanced roost protection in native woodlands.^[52] Despite these efforts, challenges persist due to limited funding and the species' solitary nature, which complicates population assessments, though acoustic monitoring has revealed stable detections on multiple islands since the 2010s.^[1] No captive breeding programs exist, with recovery focusing instead on in-situ habitat enhancement and empirical research to verify demographic viability.^[3]

Mitigation strategies and research

Mitigation strategies for the Hawaiian hoary bat (Lasiurus cinereus semotus) emphasize reducing collision mortality at wind turbines and preserving habitat integrity. At wind energy facilities, operational curtailment—feathering blades or halting rotation below wind speed thresholds of 4-6 m/s during nighttime hours—has been implemented to decrease bat fatalities, as documented in projects like Kaheawa Wind Power, where such measures align with state guidance requiring net environmental benefits.^[53]^[40] Habitat management plans, such as the Tier 1 plan for Poamoho, involve fencing to exclude feral ungulates, invasive plant control, and enhancement of native vegetation to support foraging and roosting, with goals spanning eight years to yield measurable bat occupancy increases.^[39] During the pupping season (June 1 to September 15), guidelines prohibit trimming trees taller than 15 feet to safeguard non-volant pups, as enforced by entities like the U.S. Navy on military installations.^[54] Experimental approaches include dim ultraviolet (UV) light deterrence, which reduced bat activity near trees by altering echolocation cues, as tested by USGS researchers, warranting scaled-up trials for artificial structures.^[55] Control of invasive insectivores—such as rodents, ants, and yellowjacket wasps—via small-scale suppression has shown potential to boost bat presence by increasing prey availability, per field experiments on Hawai'i Island.^[32] Pesticide application is minimized, with studies indicating secondary poisoning risks from contaminated prey, alongside ongoing mammalian predator control benefiting bats incidentally.^[56] Research efforts prioritize filling ecological knowledge gaps to inform mitigation efficacy. Radiotelemetry tracking of 28 individuals on Hawai'i Island revealed mean nightly movements of up to several kilometers, informing foraging range protections.^[4] Mist-netting and acoustic surveys from 2018-2023 across 23 sites (33-2,341 m elevation) documented capture rates and activity patterns, aiding population trend assessments.^[57] Multi-scale roost selection studies using GPS tags identified preferences for native trees like Metrosideros polymorpha at intermediate canopy heights, guiding restoration priorities.^[58] Genetic research, initiated in 2004, analyzed mitochondrial DNA to confirm North American origins and low diversity, supporting subspecies validity and highlighting inbreeding risks.^[2] Diet analyses via fecal samples linked bat foraging to moth abundance, with prey availability influencing seasonal movements.^[59] The U.S. Fish and Wildlife Service's 1998 recovery plan calls for standardized monitoring protocols, though persistent data conflicts on status underscore needs for longitudinal studies; recent DOFAW updates integrate these into guidance.^[3]^[60] Wind farm-specific research evaluates curtailment impacts, revealing unresolved offsets for residual fatalities despite reductions.^[61]

Debates on status and policy implications

The U.S. Fish and Wildlife Service (USFWS) proposed downlisting the Hawaiian hoary bat from endangered to threatened status in 2021, citing improved knowledge of its distribution across all main Hawaiian islands via acoustic surveys, lack of evidence for population decline, and recognition that the 1970 endangered listing was based on limited data regarding habitat loss and life history.^[61]^[49] This proposal reflected field observations indicating wider occupancy than previously documented, with stable to slightly increasing trends in breeding-season detection rates on islands like Hawaii over five-year surveys from 2010–2014.^[30] However, conservation organizations expressed concerns that downlisting could weaken protections amid ongoing threats like wind turbine collisions, arguing unresolved questions about population size and viability warranted maintaining endangered status.^[61] Genetic analyses have fueled further debate on the bat's taxonomic and conservation status, revealing two independent colonizations from North American hoary bat populations within the last 10,000–15,000 years rather than ancient isolation as a distinct endemic subspecies.^[7]^[62] This recency implies higher gene flow potential and less genetic bottleneck than assumed for long-isolated endemics, potentially justifying revised Endangered Species Act (ESA) protections, though USFWS has not finalized taxonomic changes and the subspecies remains listed as endangered pending broader acceptance of reclassification to genus Aeorestes.^[63] Critics of heightened endangerment claims note that acoustic data overestimate abundance due to migratory behavior and detectability biases, complicating delisting resistance.^[26] Policy implications center on balancing bat conservation with Hawaii's renewable energy mandates, particularly wind development, which has documented higher-than-predicted fatalities—up to three times initial estimates at operational facilities like Auwahi Wind.^[64]^[65] Habitat Conservation Plans (HCPs) under ESA Section 10 allow incidental take permits for wind projects, incorporating mitigations like nighttime turbine curtailment (reducing operations by 50–75% during peak bat activity), but developers have sought expanded take limits to accommodate actual mortality rates exceeding 1–5 bats per turbine annually at some sites.^[44]^[66] These policies trade short-term bat losses for long-term habitat offsets and renewable energy gains, as Hawaii aims for 100% clean energy by 2045, yet debates persist over whether uncertainty in population estimates (ranging from hundreds to low thousands) justifies stricter curtailment or alternative technologies like radar-based shutdowns, with some arguing overregulation hampers decarbonization without proportional recovery benefits.^[52]^[64]

References

[1]
Species Profile for Hawaiian hoary bat(Lasiurus cinereus semotus)
Hawaiian hoary bat is an endangered endemic mammal found in the Hawaiian archipelago. Listed as a subspecies of the hoary bat (Lasiurus cinereus).
[2]
Analysis of Genomic Sequence Data Reveals the Origin and ...
Because this endangered species is of conservation concern, a clearer understanding of the population genetic structure of this bat in the Hawaiian Islands is ...
[3]
[PDF] Recovery Plan for the Hawaiian Hoary Bat - ECOS
Current Species Status: The Hawaiian hoary bat (Lasiurus cinereus semotus) is federally listed as endangered. It is known from the islands of Hawaii, Maui,.
[4]
Ecology and Distribution of the Endangered Hawaiian Hoary Bat
The Hawaiian Hoary Bat (Lasiurus cinereus semotus) is the only extant land mammal native to the Hawaiian archipelago. It is listed as endangered.<|separator|>
[5]
[PDF] Hawaiian Hoary Bat Guidance Document
c. cinereus), and is listed as endangered under both the Federal Endangered Species Act (ESA) and Hawai'i endangered species laws (Hawai'i Administrative Rules ...
[6]
[PDF] Hawaiian hoary bat responses to habitat, season, and non‐native ...
Sep 6, 2021 · The Hawaiian hoary bat (Lasiurus semotus) is the only native terrestrial mammal in the Hawaiian Islands and is listed as federally endangered ...
[7]
Analysis of Genomic Sequence Data Reveals the Origin and ...
Aug 27, 2020 · Winter distribution and use of high elevation caves as foraging sites by the endangered Hawaiian hoary bat, Lasiurus cinereus semotus.Missing: peer- | Show results with:peer-
[8]
Hawaiian hoary bat population genetics 1988 to 2020 - USGS.gov
Jan 25, 2023 · 2023 · Hawaiian hoary bat population genetics 1988 to 2020 · 10.5066/P9COQ3ZK · Corinna Pinzari, Mona Renee Bellinger, Donald K. Price, Frank J.
[9]
Lasiurus semotus • Hawaiian Hoary Bat
Taxonomy ; Suborder, : Yangochiroptera ; Superfamily, : Vespertilionoidea ; Family, : Vespertilionidae ; Subfamily, : Vespertilioninae ; Tribe, : Lasiurini.Missing: classification nomenclature
[10]
Lasiurus cinereus (hoary bat) | INFORMATION - Animal Diversity Web
Class, Mammalia mammals ; Order, Chiroptera bats ; Family, Vespertilionidae evening bats and vesper bats ; Genus, Lasiurus hairy-tailed bats.
[11]
Molecular systematic revision of tree bats (Lasiurini): doubling the ...
Sep 24, 2015 · The phylogeography of the endangered Hawaiian hoary bat is studied in detail, and hoary bats are found to have undergone multiple invasions of ...Abstract · Materials and Methods · Results · Discussion
[12]
Lasiurus semotus - NatureServe Explorer
Hawaiian Hoary Bat - Natureserve Global Rank: G2: Small range limited to the Hawaiian Islands; common on at least two islands (Kaui and Hawaii); ...
[13]
The genus and species names “Lasiurus” (meaning hairy tailed) and ...
Oct 30, 2020 · The Hoary Bat is listed as Least concern on the IUCN Red List. One subspecies, the Hawaiian hoary bat (Lasiurus cinereus semotus), is listed as ...
[14]
Wildlife Program | ʻŌpeʻapeʻa, the Hawaiian Hoary Bat
The ʻōpeʻapeʻa, or Hawaiian hoary bat, is Hawai'i's only native terrestrial mammal, with white-tipped brown and gray fur, and is the state mammal.Missing: classification nomenclature
[15]
Hawaiian Hoary Bat - Bat Conservation International
Hawaiian hoary bats are the only bat species found on the Hawaiian Islands and the only native land mammal to the islands.Missing: classification taxonomy nomenclature
[16]
Nuclear and mtDNA phylogenetic analyses clarify the evolutionary ...
Currently, the conservation status of Hawaiian hoary bats reflects the previous taxonomy: Lasiurus cinereus semotus is considered endangered; L. c. cinereus ...
[17]
Analysis of Genomic Sequence Data Reveals the Origin ... - PubMed
Sep 1, 2020 · We examine the genetic history and population status of Hawaiian hoary bats (Lasiurus semotus), the most isolated bats on Earth, and their relationship to ...
[18]
Modeling the colonization of Hawaii by hoary bats (Lasiurus cinereus)
We conducted a three-stage analysis to identify conditions under which hoary bats originally colonized Hawaii. We used FLIGHT to determine if stores of fat ...Missing: phylogenetic origins
[19]
Multiple Dispersal Events in the Founding of Hoary Bat Populations ...
Here, we use molecular data to test the hypotheses that (1) Hawaiian L. c. semotus originated via dispersal from North American populations of L. c. cinereus ...
[20]
Genetic diversity, structure, and effective population size of an ... - NIH
This manuscript was greatly improved with peer review from R. ... ʻōpeʻapeʻa or Hawaiian hoary bat (Lasiurus cinereus semotus): 5-year review summary and ...
[21]
[PDF] Biological - ECOS
Hawaiian Hoary Bat. Species Description. The Hawaiian hoary bat is medium-sized (0.5 to 0.8 ounces) nocturnal, insectivorous bat, with a wingspan of 10.5 to ...
[22]
Morphological divergence in an insular bat, Lasiurus - jstor
This has enabled the Hawaiian bat to include large, and hard-bodied insects (not taken by L. c. cinereus) in its diet despite a marked reduction in body size.
[23]
Hoary Bat - Lasiurus cinereus - A-Z Animals
The hoary bat is preyed upon by hawks, owls, kestrels, and snakes. The bat's fur color provides a degree of camouflage while it's roosting in trees. What does ...Missing: adaptations colonization
[24]
[PDF] U·M·I - ScholarSpace
This dissertation concerns the allopatric divergence of an island bat, the. Hawai \ian hoary bat, Lasiurus cinereus semotus, from its continental parent.
[25]
Variation in the echolocation calls of the hoary bat (Lasiurus cinereus)
Aug 5, 2025 · The Hawaiian hoary bat, Lasiurus cinereus semotus, foraged in both an open and a cluttered habitat. In the cluttered habitat it used slow, ...Missing: fur | Show results with:fur
[26]
Native Beauty: The Hawaiian Hoary Bat
Nov 14, 2024 · But scientists performing genetic research in 2020 found the Hawaiian bat to be distinct genetically from the northern hoary bat and found ...
[27]
[PDF] 5-YEAR STATUS REVIEW - ECOS
Sep 30, 2011 · The ʻōpeʻapeʻa or Hawaiian hoary bat is an endangered endemic mammal found in the. Hawaiian archipelago. Listed as a subspecies of the hoary bat ...
[28]
Distribution of Hawaiian hoary bat sightings and echolocation...
Although the Hawaiian hoary bat (Lasiurus cinereus semotus) was first collected in 1861 (Gray 1862) and occurs on all major volcanic islands in Hawai'i (Tomich ...<|separator|>
[29]
[PDF] Endangered Species Recovery Committee Hawaiian Hoary Bat ...
cinereus), and is listed as endangered under both the Federal. Endangered Species Act (ESA) and Hawai'i endangered species laws (Hawai'i Administrative. Rules ...
[30]
[PDF] A Five-yeAR STUdy oF HAwAiiAn HoARy BAT (LASiURUS ...
Despite the listing of “endangered” for this Hawaiian endemic subspecies by the. U.S. Fish and Wildlife Service (USFWS) since 1970 and publication of a recovery ...
[31]
Hawaiian hoary bat (Lasiurus cinereus semotus) activity, diet and ...
Hawaiian hoary bats consumed both native and non-native insect species, including several invasive species damaging to crop agriculture. Moths were the primary ...
[32]
Hawaiian hoary bat responses to habitat, season, and non‐native ...
Jan 7, 2022 · Hawaiian hoary bat foraging behavior may shift within the landscape in response to significant competition where non-native insectivore ...Missing: elevational | Show results with:elevational
[33]
Foraging range movements of the endangered Hawaiian hoary bat ...
Mar 27, 2015 · We documented nightly movements of Hawaiian hoary bats (Lasiurus cinereus semotus) on the island of Hawai'i. Based on data from 28 radiotagged ...
[34]
Multi-scale assessment of roost selection by 'ōpe'ape'a, the ... - NIH
Aug 24, 2023 · A five-year study of Hawaiian hoary bat (Lasiurus cinereus semotus) occupancy on the Island of Hawai'i. University of Hawai'i at Hilo, Hawai ...
[35]
[PDF] 'Ōpe'ape'a or Hawaiian hoary bat
Oct 1, 2015 · SPECIES INFORMATION: The 'ōpe'ape'a, or Hawaiian hoary bat (Family: Vespertilionidae), is Hawai'i's only native terrestrial mammal, although ...
[36]
[PDF] A Five-yeAR STUdy oF HAwAiiAn HoARy BAT (LASiURUS ...
Conversely, the lowest period of occupancy occurred between December and March each year as bats disperse to high elevations in the annual migration.Missing: elevational | Show results with:elevational
[37]
[PDF] Exploratory population modeling exercise of Hawaiian Hoary Bat
Population trends for an array of Hawaiian Hoary Bat PVA models without take. Expert Elicitation for mainland hoary. Lowest Mortality for Lcc: 35% ad, 55% juv.Missing: "peer | Show results with:"peer
[38]
Hawaiian hoary bat (Lasiurus cinereus semotus) acoustic ...
Acoustic sampling for occurrence of the endangered Hawaiian hoary bat (Lasiurus cinereus semotus) was conducted at 18 “long-term” acoustic monitoring ...Missing: trends | Show results with:trends
[39]
[PDF] Hawaiian Hoary Bat Tier 1 Mitigation Habitat Management Plan for ...
Estimates of contemporary genetic effective population sizes1 indicate population sizes are ... Hawaiian hoary bat and currently no estimates of population ...
[40]
The Kaheawa II Wind Farm and the Fate of the Hawaiian Hoary Bat
Aug 18, 2019 · It claims that “There are no studies to date that test whether mortality rates decrease significantly when LWSC is raised from 5.5 m/s to 6.5 m/ ...
[41]
[PDF] United States Department of the Interior
Hawaiian hoary bat mortality per mile of fence. Kaheawa Wind Power. 2008. Kaheawa Pastures Wind Energy Generation Facility Habitat. Conservation Plan, Year 2 ...
[42]
[PDF] ESA Listed Species - U.S. Fish and Wildlife Service
Hawaiian hoary bats forage for insects from as low as 3 feet to higher than 500 feet above the ground and can become entangled in barbed wire used for fencing.
[43]
[PDF] Hawaiian Hoary Bat or Ōpea'ape'a. (Lasiurus cinereus semotus)
Focal Species: Hawaiian Hoary Bat or Ōpea'ape'a. (Lasiurus cinereus semotus). Synopsis: The Hawaiian Hoary Bat is the only land mammal native to Hawai'i.Missing: classification taxonomy nomenclature
[44]
[PDF] Hawaiian Hoary Bat Guidance for Renewable Wind Energy ...
Hawaiian Hoary Bat because threats and factors that limit the bat population are unknown. ... Adult mortality rates: ranging from 20-50 percent annual mortality.
[45]
Ultrasonic acoustic deterrents significantly reduce bat fatalities at ...
Our results indicate deterrents significantly reduced bat fatalities for Lasiurus cinereus and Tadarida brasiliensis by 78% and 54%, respectively.<|separator|>
[46]
[PDF] Farmers Can Help our Hawaiian Hoary Bats
Some insecticides can also affect bats through bioaccumulation of chemicals through the food chain. Herbicides can alter vegetation structure and community.
[47]
[PDF] Potential Citric Acid Exposure and Toxicity to Hawaiian Hoary Bats ...
Feb 14, 2014 · A high dose delivered orally (2,811 mg/kg) was toxic to the big brown bats and emesis occurred in 1 bat dosed as low as the 759 mg/kg level. No ...
[48]
[PDF] Screening of Hawaiian bird and bat carcasses for rodenticide residues
Sep 14, 2018 · In this study, we sought to evaluate the evidence for exposure of a suite of Hawaiian birds and the Hawaiian hoary bat to common rodenticides by ...
[49]
FWS Proposes Downlisting Bat From Endangered to Threatened
Nov 1, 2021 · ... U.S. Fish and Wildlife Service is seeking to reclassify the 'ōpe'ape'a or Hawaiian hoary bat from endangered to threatened. The species was ...<|separator|>
[50]
[PDF] Endangered Species Recovery Committee Hawaiian Hoary Bat ...
Hoary bats have constituted the highest proportions of fatalities at most continental U.S. facilities, ranging from 9 to 88 percent of all bat. Page 6. APPROVED ...
[51]
[PDF] Hawaiian Hoary Bat Guidance for Wind Energy Project Habitat ...
Although population estimates for the Hawaiian hoary bat are not currently available, one acoustics study suggests that bat numbers on the island of. Hawai'i ...
[52]
Hawaiian Hoary Bat Guidance for Wind Energy Project Habitat ...
Feb 1, 2024 · The Hawaiian hoary bat (Lasiurus cinereus semotus) is listed as endangered under state and federal law. The operation of wind turbines in Hawai' ...
[53]
[PDF] Hawaiian Hoary Bat Guidance for Wind Energy Projects
Jan 25, 2021 · ... Hawaiian hoary bat, but insects such as flies, termites, crickets, and leaf bugs are also sometimes consumed. Insects and other arthropods ...
[54]
Protecting the Hawaiian Hoary Bat During Pupping Season
Jul 20, 2023 · To protect Hawaiian Hoary Bat pups, avoid trimming trees over 15 feet during pupping season (June 1-September 15), as pups cannot fly and hang ...
[55]
Dim ultraviolet light as a means of deterring activity by the Hawaiian ...
Dim UV light was tested to reduce bat activity near trees, and results showed a decrease in bat activity, justifying further testing.Missing: mitigation strategies
[56]
[PDF] Hawaiian Hoary Bat - Marine Corps Base Hawaii
The Hawaiian hoary bat, or 'Ōpe'ape'a, is brown to grey with white tips, has a 10.5-13.5 inch wingspan, and is endemic to the Hawaiian Islands.Missing: classification taxonomy nomenclature<|separator|>
[57]
Hawai'i Island ʻōpeʻapeʻa (Hawaiian hoary bat; Lasiurus semotus ...
Aug 24, 2023 · 'Ōpe'ape'a (Hawaiian hoary bats; Lasiurus semotus) were surveyed at 23 sites on Hawaiʻi Island from 33 to 2,341 m elevation from May 2018 to ...<|separator|>
[58]
Multi-scale assessment of roost selection by 'ōpe'ape'a, the ...
Aug 24, 2023 · Multi-scale assessment of roost selection by 'ōpe'ape'a, the Hawaiian hoary bat (Lasiurus semotus) ... differences between female and male bats.<|control11|><|separator|>
[59]
Hawaiian hoary bat (Lasiurus cinereus semotus) activity, diet and ...
Habitat use, diet, prey availability, and foraging ecology of the endangered Hawaiian hoary bat (Lasiurus cinereus semotus, Vespertilionidae), was examined ...
[60]
Wildlife Program | Hawaiian Hoary Bat Guidance Document
The Division of Forestry and Wildlife (DOFAW ) is in the process of updating the last version of the guidance document (updated January 2021).
[61]
Feds Say Hawaiian Hoary Bats Aren't Endangered Despite ...
Nov 8, 2021 · One effective technique to reduce bat deaths, Phillips said, calls for wind farm developers to lock the turbine blades to stop them from ...
[62]
Researchers Trace Hawaiian Hoary Bat Origins - The Wildlife Society
one ...
[63]
Federal Register :: Endangered and Threatened Wildlife and Plants
Apr 24, 2023 · Therefore, we are withdrawing that portion of the direct final rule concerning the listed entity Hawaiian hoary bat ( Lasiurus cinereus semotus) ...Missing: ESA | Show results with:ESA
[64]
Wind Farms Want Permission To Kill More Bats — A Lot More
Aug 14, 2018 · Hawaii wind energy projects pose a greater threat to the endangered Hawaiian hoary bat population than was originally expected. PF Bentley ...
[65]
[PDF] Bats and Wind Energy: Impacts, Mitigation, and Tradeoffs
Nov 15, 2018 · The ecological consequences of bats avoiding wind facilities would depend on whether increased mortality or habitat availability are limiting.
[66]
Federal Register, Volume 83 Issue 106 (Friday, June 1, 2018)
Jun 1, 2018 · ... endangered Hawaiian hoary bat; the endangered Hawaiian goose; and the endangered Hawaiian petrel. Three of the applicants were authorized to ...