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Community Composition of Bats in Cusuco National Park, Honduras, a Mesoamerican Cloud Forest, Including New Regional and Altitudinal Records

Pamela Medina-Van Berkum1*, Kevina Vulinec1,2, Declan Crace1, Zeltia López Gallego11, and Thomas Edward Martin1

1Operation Wallacea Ltd., Wallace House, Old Bolingbroke, Spilsby, Lincolnshire, PE23 4EX, UK. 2Delaware State University,Department of Agriculture and Natural Resources, Dover, DE 19901, USA.*Corresponding author.

Neotropical Naturalist, No. 3 (2020)

Mesoamerican cloud forests support diverse bat communities, but remain poorly studied. Here, we provide an overview of the bat community of Cusuco National Park (CNP), Honduras, an area of tropical cloud forest and adjacent lowland habitats. Our results are based on one of the longest-running bat surveys completed regionally to date. Mist nets were deployed over 14 eight-week research seasons running June through August 2006 to 2019. A total of 6,854 bats were caught, with 59 species identified, including two major range extensions (Natalus lanatus and Eptesicus brasiliensis) and three species at higher elevations than previously reported. Species richness estimators predicted 63 bat species in the park. We highlight CNP as an important center of bat diversity and an important conservation priority.

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No. 3 Neotropical Naturalist 2020 Community Composition of Bats in Cusuco National Park, Honduras, a Mesoamerican Cloud Park, Including New Regional and Altitudinal Records Pamela Medina-Van Berkum, Kevina Vulinec, Declan Crace, Zeltia López Gallego, and Thomas Edward Martin NEOTROPICAL NATURALIST The Neotropical Naturalist (ISSN # 2327-5472) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680-0009. Phone 207-546-2821 Ext. 4, FAX 207-546-3042. E-mail: Webpage: http://www.eaglehill. us/neon. Copyright © 2020, all rights reserved. Published on an article by article basis. Special issue proposals are welcome. The Neotropical Naturalist is an open access journal. Authors: Submission guidelines are available at neon. Co-published journals: The Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, Urban Naturalist, and Eastern Paleontologist, each with a separate Board of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Federal ID # 010379899). Board of Editors David Barrington, Department of Plant Biology, University of Vermont, Burlington, VT, USA William G. R. Crampton, University of Central Florida, Orlando, FL, USA Paulo Estefano Dineli Bobrowiec, Instituto Nacional de Pesquisas da Amazônia, Brazil Valentina Ferretti, Universidad de Buenos Aires, Argentina Danny Haelewaters, Ghent University, Belgium Matthew Halley, Drexel University, Philadelphia, PA, USA Christopher M. Heckscher, Department of Agriculture and Natural Resources, Delaware State University, Dover, DE, USA Ian MacGregor-Fors, Instituto de Ecología Mexico, Veracruz, Mexico Klaus Mehltreter, Institute of Ecology, A.C., Xalapa, Veracruz, Mexico Jorge Ari Noriega A., Universidad de los Andes, Colombia Jason M. Townsend, Biology Department, Hamilton College, Clinton, NY, USA Judit Ungvari, Florida Museum of Natural History, Gainesville, FL, USA Fredric V. Vencl, Stony Brook University, Stony Brook, NY. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA Kevina Vulinec, Department of Agriculture and Natural Resources, Delaware State University, Dover, DE, USA ♦ The Neotropical Naturalist (ISSN 2327-5472) is a peer-reviewed journal that publishes articles on all aspects of the natural history sciences of terrestrial, freshwater, and marine organisms and the environments of the neotropics from Mexico through the southern tip of South America. 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Special issue editors can rely on the publisher ’s years of experiences in efficiently handling most details relating to the publication of special issues. ♦ Indexing - As is the case with Eagle Hill's other journals, the Neotropical Naturalist is expected to be fully indexed in Elsevier , Thomson Reuters, Proquest, EBSCO, Google Scholar, and other databases. ♦ The journal's staff is pleased to discuss ideas for manuscripts and to assist during all stages of manuscript preparation. The journal has a page charge to help defray a portion of the costs of publishing manuscripts. Instructions for Authors are available ( ♦ It is co-published with the Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, Urban Naturalist, and Eastern Paleontologist. ♦ It is available online in full-text version on the journal's website ( Arrangements for inclusion in other databases are pending. Cover Photograph: Centurio senex in the cloud forests of Cusuco National Park, north-west Honduras. Photograph © Anikó Kurali. Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 1 2020 NEOTROPICAL NATURALIST 3:1–24 Community Composition of Bats in Cusuco National Park, Hounduras, a Mesoamerican Cloud Forest, Including New Regional and Altitudinal Records Pamela Medina-Van Berkum1*, Kevina Vulinec1,2, Declan Crace1, Zeltia López Gallego1, and Thomas Edward Martin1 Abstract - Mesoamerican cloud forests support diverse bat communities, but remain poorly studied. Here, we provide an overview of the bat community of Cusuco National Park (CNP), Honduras, an area of tropical cloud forest and adjacent lowland habitats. Our results are based on one of the longest-running bat surveys completed regionally to date. Mist nets were deployed over 14 eight-week research seasons running June through August 2006 to 2019. A total of 6,854 bats were caught, with 59 species identified, including two major range extensions (Natalus lanatus and Eptesicus brasiliensis) and three species at higher elevations than previously reported. Species richness estimators predicted 63 bat species in the park. We highlight CNP as an important center of bat diversity and an important conservation priority. Introduction Cloud forests are complex ecosystems which typically span a range of environmental gradients related to climate, soil, and geomorphological conditions. Consequently, cloud forests are one of the more diverse ecosystems worldwide, harbouring high numbers of endemic and threatened species, as well as providing a range of important ecosystem services (Bruijnzeel et al. 2010, Bubb et al. 2004). In these forests, more than the 40% of mammal species are bats (Bruijnzeel et al. 2010), which play a pivotal role in ecological processes, acting as seed dispersers, pollinators, and prey population regulators (Fleming 1993, Kalko et al. 1996, Medellín et al. 2000). The cloud forests of Central America, which form part of the Mesoamerican biodiversity ‘hotspot’ (Myers et al. 2000, Myers 2003), occur disjunctively from southern Mexico to Panama and support diverse and highly endemic species assemblages for most faunal groups (CEPF 2019, Powell and Palminteri 2001). However, despite their biological importance, comparatively little research has examined the composition of ecological communities in these cloud forests (Bubb et al. 2004); a trend that includes bat communities (Reid 2009). Most community-level bat research in Mesoamerican cloud forests to date has focussed on Mexico (e.g., Briones-Salas et al. 2005, Calderón-Patrón et al. 2013, Pineda et al. 2005) and Costa Rica (LaVal 2004, LaVal and Fitch 1977), with Chiroptera research in the contiguous countries of Belize, Guatemala, El Salvador, Honduras and Nicaragua largely confined to species-specific records from lowland ecosystems (e.g., Divoll and Buck 2013, Fenton 2001, Herrera et al. 2018, Loza et al. 2018, Medina-Fitoria et al. 2015, Mora et al. 2014). There are many aspects of the distribution and ecology of Honduran bat populations that remain unknown. Significant knowledge gaps remain regarding distributions of species within different habitats, as well as data on species ecology, community composition, and spatial patterns of diversity within different cloud forest ecosystems. Faunal communities in cloud forests are facing severe threats from deforestation (Bubb et al. 2004, Mulligan 2010) and climate change (Neate-Clegg et al. 2019, Pounds et al. 1999). Without an understanding of regional species distributions, it is difficult 1Operation Wallacea Ltd., Wallace House, Old Bolingbroke, Spilsby, Lincolnshire, PE23 4EX, UK. 2Delaware State University, Department of Agriculture and Natural Resources, Dover, DE 19901, USA. Manuscript Editor: Fredric Vencl Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 2 to identify conservation priority areas or assess the conservation value of specific Mesoamerican cloud forest ecosystems. This study attempts to partially address this research gap by summarizing data from one of the longest-running bat surveys in northern Central America in order to characterize the cloud forest bat community of Cusuco National Park (CNP), north-west Honduras. We report all species recorded from these surveys, as well as provide altitudinal delimitations for all species in our inventory and identify notable range extensions and altitudinal records. We construct species accumulation curves and non-parametric species richness estimators to extrapolate overall diversity within CNP and examine species turnover and feeding guild dominance across the Park’s altitudinal gradient. Thus, we provide the most detailed overview of bat community structure in a northern Central American cloud forest published to date. Methods Study site CNP (15°32′31″ N, 088° 15′49″ W) is a 23,440 ha protected area spanning the departments of Cortés and Santa Bárbara in north-west Honduras (Fig. 1). It is located in the Sierra de Omoa, part of the Merendón mountain range, and spans an altitudinal range of 500–2,242 m (Lenkh 2005). The area of CNP is divided into a 7,690 ha core zone where socio-economic activity is nominally prohibited, and an encompassing 15,750 ha buffer zone where some limited agriculture and other land use is permitted (Martin and Blackburn 2009). Average annual precipitation is 2,788 mm, with 45% of precipitation falling in the wettest months between October–December (Fundación Ecologista 1994). Mean day time temperatures in summer (June–July) range from 21 °C at 1,150 m (range: 18 °C–23.5 °C) to 15 °C at 2,200 m (13 °C–16.5 °C) (Jones 2020). Broadly, all ecosystems within the Park fall into the “moist tropical forest” Holdridge life zone (FAO 2002), although as with many “cloud forest” National Parks, CNP represents a mosaic landscape with different vegetation types across its various elevational belts. These include cleared land, coffee plantations, and remnant patches of lowland forest at lower altitudes (500–1,200 m asl), pine-oak forest and tropical montane cloud forest in the middle-upper altitudinal zones (1,200–2000 m asl), and “Bosque enano” elfin forest on the highest ridges of the park (>2,000 m asl) (Lenkh 2005). A long-term biodiversity monitoring program run by the UK-based company Operation Wallacea (, has been ongoing in CNP since 2006. This program has resulted in detailed species inventories for several taxa (Hoskins et al. 2018, Martin et al. 2016), and the Park’s biodiversity is recognized to be of global importance (Le Saout et al. 2013). However, previous published bat research from CNP is restricted to a genetic analysis of populations of two fruit bat species (Asher 2009) and a methodological study examining novel means of detecting altitudinal movements in bats (Erzerberg et al. 2011). Sampling design To evaluate the bat community in CNP, we conducted mist-netting surveys for an eight-week period between June and August each year between 2006 and 2019. Initially, we conducted mistnetting at 36 sites at seven camps located between 605 and 1925 m asl (Table 1) to cover all the major altitudinal bands and associated habitat classifications present in the Park. However, the lowest altitude camp (Santo Tomas, 605 m asl) was not surveyed between 2015 and 2019, thus only 31 mist-netting sites were surveyed in these years. Within each camp, we established three to five fixed-location mist-netting sites between 300 and 600 m apart. We typically conducted mist-netting in three camps each night, for six nights a week. For each survey session at each site, we used five nets (a combination of 6 m x 2.6 m and 12 Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 3 m x 2.6 m nets, all with 36-mm mesh and five shelves). The mist nets were opened before sunset and checked every 15 to 30 minutes for six hours. Mist-netting sites were surveyed in rotation. Once a mist-netting site was sampled, it was not surveyed again for at least three nights. Mist nets were always placed in the same positions on each survey night. We did not conduct mist-netting surveys in heavy rain. We followed Sikes and Gannon (2011) with regards to protocols for capturing and handling bats. We extracted bats captured and placed them within cotton holding bags. We processed the bats by weighing the individuals (g) using pre-calibrated Pesola spring balances and took forearm measurements (mm) using vernier callipers. Sex, reproductive stage (female: non-reproductive, pregnant, lactating, post-lactating; male: scrotal, or non-scrotal), and age (juvenile or adult) were also recorded, with age being determined by the degree of ossification of the metacarpal-phalangeal joints (Baagøe 1977). We identified individuals to species level using authoritative regional field guides and taxonomic keys (Medellín et al. 2007, Mora 2016, Reid 2009, Timm and LaVal 1998). We also recorded weather conditions on each survey night (Rain = 0 or 1, Wind = 0 or 1, Moon = 0 or 1). Data analysis We used two broad approaches to examine the data: the construction of an annotated species inventory for the study site and a quantitative analysis of the diversity and structure of the bat community. We compiled a list of all species detected during surveys in CNP following the taxonomy provided by IUCN (2019), with five exceptions: Sturnira parvidens, Uroderma convexum, Lasiurus frantzii, Myotis pilosatibialis, and Natalus lanatus—all recently-split species, which are broadly recognized in the literature (e.g., Castaño et al. 2018, Medina-Fitoria et al. 2015, Rodríguez-Herrera and Sánchez 2015, Timm and LaVal 2018, York et al. 2019) but remain unrecognized by the IUCN (2019). We followed the taxonomy provided by Velazco and Patterson (2013), Mantilla-Meluk (2014), Baird et al. (2015), Mantilla-Meluk and Muñoz-Garay (2014), Figure 1. Elevation map of Cusuco National Park, showing camps and boundary of the buffer zone and core zone. Insert shows location of Cusuco National Park within Central America. Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 4 Table 1. Summary of camp locations and survey effort in study camps, and non-parametric species richness estimators of bat species in Cusuco National Park (CNP), north-west Honduras, as well as species richness estimators for each camp. Chao, Jackknife 1, and Bootstrap are non-parametric species richness estimators (Colwell and Coddington 1994, Magurran 2004). CNP Santo Tomas Buenos Aires Guanales Cortecito Base Camp El Danto Cantiles Elevation (mean m asl) 605 850 1287 1400 1572 1591 1825 Sampling effort (m2h) 398,383 24,408 48,865 56,262 44,824 129,113 44,831 55,535 Individuals captured 6,854 1,139 2,042 646 921 1,481 316 309 Capture rate 0.0172 0.0467 0.0418 0.0115 0.0205 0.0115 0.0076 0.0058 Richness observed 59 40 43 33 35 46 34 21 Chao (± SE) 62.12 (± 3.65) 49.66 (± 7.69) 56.10 (± 12.11) 44.94 (± 12.84) 40.97 (± 5.94) 49.45 (± 3.46) 43.97 (± 7.57) 40.65 (± 19.61) Jacknife 1 (± SE) 63.99 (± 2.23) 50.54 (± 3.73) 51.74 (± 2.91) 39.83 (± 2.58) 41.83 (± 3.25) 52.90 (± 2.96) 45.64 (± 4.36) 29.74 (± 4.03) Bootstrap (± SE) 61.63 (± 1.37) 44.91 (± 2.13) 46.85 (± 1.62) 36.01 (± 1.39) 38.33 (± 2.10) 49.71 (± 1.84) 39.49 (± 2.51) 24.59 (± 1.98) Average richness estimate 63 48 52 41 40 51 43 31 Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 5 and Tejedor (2005) for these four species respectively. We noted endemism and conservation status of each species. Mesoamerican hotspot endemics were defined by cross-referencing the demarcations of the hotspot defined in Myers et al. (2000) with range maps provided in Reid (2009) and IUCN (2019). International conservation status followed IUCN (2019), while national conservation status followed Hernández (2015). We determined whether our records for each species represented extensions to their known range by comparing each species in our inventory with distribution maps provided by IUCN (2019), Reid (2009), and a range of local publications concerning bat records within Honduras (Appendix S1, in Supplemental File 1, available online at We used two classifications for range extensions: “major” range extensions for species that had not previously been reported in north-west Honduras and “minor” extensions for species that had previously been reported in north-west Honduras but are not specified by existing range maps to occur in the vicinity of CNP. We also reported the highest and lowest altitudes where each species was detected in CNP and determined whether any of these values represented novel altitudinal records by comparing our data to altitudinal delimitations for each species provided by IUCN (2019), Reid (2009), and consulted literature (Almazán-Núñez et al. 2018, Sánchez-Cordero 2001, Timm and LaVal 2018). We categorized each species in our inventory into broad feeding guilds: frugivores, nectarivores, insectivores, carnivores, and sanguivores based on the relevant literature (Giannini and Kalko 2004, Kalko et al. 1996, Reid 2009, Soriano 2000). Many Neotropical bat species have varied diets and can switch dietary preferences depending on environmental conditions (Lobova et al. 2009). Our categories thus reflect primary dietary preference, as indicated in the literature, rather than an absolute categorization. We assigned relative abundance categories for each species based on capture frequency, adapted from methods used in Patterson et al. (2017). These categories were: A = Abundant, (captured more than 201 times); C = common (captured between 101 and 200 times); U = uncommon (captured between 21 and 100 times); R = rare (captured less than 20 times). Statistical analysis To generate bat community diversity estimates, we first merged records of Carollia subrufa with records of C. sowelli, given that some individuals of these morphologically very similar species may have been misidentified. Specifically, separating these two species requires examination of banding patterns on the fur as well as morphometric measurements. It is possible that this was not checked by some observers in the early years of survey work, potentially leading to some C. subrufa being incorrectly identified as C. sowelli. As this potential error may have a bearing on our quantitative analysis, we took a conservative approach of lumping these species together for all such analysis. To standardize for unequal sampling effort per night, we divided the total number of bats captured per night by the total capture effort during those nights. Because we used two different sizes of mist net, sampling effort was calculated following methods described by Straube and Bianconi (2002), by summarizing the total meters of net and multiplying this by the number of hours nets were open per night (normally 540 m2/h). Considering each night as an independent sampling occasion, we constructed species accumulation curves with 1000 randomizations for observed and estimated species richness for the park and the camps. We chose three non-parametric species richness indicators for this; Chao, Jackknife 1, and Bootstrap (Colwell and Coddington, 1994, Magurran 2004), these being regarded as particularly suitable for extrapolating richness estimates within biological communities (Walther and Moore 2005). Following Sodhi et al. (2005), we took the mean value from these three estimators (rounding to the nearest whole value) as a “true” species richness estimate for CNP, given that the effectiveness of different estimators varies depending on dataset composition. Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 6 To investigate whether elevation has an influence on bat assemblages, we aggregated the captures from all mist-netting sites into five elevational ranges, each consisting of a 300 m isocline, ranging from 600 to 2100 m. We evaluated the efficiency of the sampling effort and the representation of bat communities in these elevational bands by sampling completeness estimated through coverage percentage (i.e. proportion of the total number of individuals in a community belonging to the species represented in the sample) (Chao and Jost 2012). We estimated Hill numbers (q0, q1, and q2) measures using individual based abundance rarefaction and extrapolation (Hsieh et al. 2016). This facilitates the comparison of multiple assemblages by providing estimators for inter- and extrapolation, allowing reduction of sampling effort effect, estimation of sample completeness and uses a bootstrapping method (Chao et al. 2014, Hill 1973).The q0 (Species richness) is completely insensitive to species abundance: q1 (Shannon diversity) measures the number of common species in a community, and q2 (Inverse Simpson diversity) measures the number of very abundant or dominant species in a community (Chao et al. 2014). Species diversity profiles were compared through overlapping confidence intervals (5–95%). We used linear regressions to assess whether altitudinal gradient influences the capture rate. To evaluate a priori if linear models were appropriate for the data, we performed Shapiro-Wilk tests to determine if residuals possessed a normal distribution (Gotelli 2013). The assumption was met after the data were square root-transformed. We performed all analyses using RStudio version 3.6.1 (R Development Core team 2019), using the packages vegan (Oksanen et al. 2019), and iNEXT (Hsieh et al. 2016), and data were plotted using the package ggplot2 (Wickham 2016). Results Field work in CNP between 2006 and 2019 comprised 398,383 m2 h of total survey effort. A total of 7,192 individual bats were captured, of which 6,854 (95.7%) could be reliably identified, representing 59 species, 34 genera, and six families (Table 2). Eight (13.6%) of these species are endemic to the Mesoamerican biodiversity hotspot. Two species, Natalus lanatus and Eptesicus brasiliensis, were only the second country records for Honduras and represented major range extensions in CNP. A further three records represent minor range extensions (Enchisthenes hartii, Myotis velifer and Dermanura azteca). We detected three species, Centurio senex, Myotis albescens and Platyrrhinus helleri, at a higher altitude than previously reported elsewhere in the literature. Centurio senex was found 300 m higher than previously reported and the other two species at 100 meters higher. No species were detected at a lower altitude than previously reported. Regarding conservation status, Bauerus dubiaquercus is classified as Near-Threatened by IUCN (2019), and Hylonycteris underwoodi is classified as Threatened in Honduras (Hernández 2015). All species detected in CNP are summarized in Table 2. Phyllostomidae represented 86% of the total captures yielded in CNP, followed by Vespertilionidae (11%), Molossidae (1.4%), Mormoopidae (1.2%), and Natalidae and Thyropidae, each representing 1% of captures. The ten most abundant species caught in CNP comprised 77% of all captures. On the other hand, 26 species (44% of all species detected) had fewer than 20 captures during the study period and accounted for 1.8% of the total captures. In most cases, these rarely-caught species were high-flying insectivorous bats, which are difficult to capture in mist nets; hence our results cannot be interpreted as indicative of abundance or habitat preferences. However, acoustic recordings yielded a considerable amount of activity by many of these high-flying insectivorous species (K. Vulinec, pers. obvs.). Captures were dominated by two species: Artibeus jamaicensis and Sturnira hondurensis, which together constituted >2,000 individuals representing 31.3% of the total captures (an average of 80 Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 7 and 70 individuals captured per year, respectively). Among the common species found in the park, B. dubiaquercus and H. underwoodi have a Threatened conservation status, and both species are present in all the elevation ranges surveyed. Few species showed restricted elevation ranges, despite being commonly captured. Species such as Carollia castanea, Platyrrhinus helleri, Molossus rufus, and Vampyressa thyone were mainly captured within villages located in the buffer zone at lower altitudes between 700 and 1,000 m asl. On the other hand, Chrotopterus auritus and Chiroderma villosum were mainly captured in the core zone of the Park at an altitude around 1,400 m asl. Moreover, most of the rarely captured aerial insectivores, such as Lasiurus ega, L. frantzii, L. blossevillii, N. mexicanus, and N. lanatus were captured between 1,500 and 1,800 m asl (Table 2). Non-parametric species estimates (Table 1) predict an overall richness of 63 species to be present in CNP, suggesting our survey effort has detected the majority of understorey bat species (93.6%) likely to be present in the study area. This is corroborated by the species accumulation curve for CNP (Fig. 2), the trajectory of which suggests overall richness to be slightly higher than that yielded by our sampling effort. Bat richness varied significantly between the survey camps (Table 1). According to non-parametric species estimates, Base Camp, Buenos Aires, and Santo Tomas were the camps with greatest bat richness (each with >45 species). Cantiles, located at the highest altitude, was the camp with the lowest recorded richness (21 species). Sample coverage curves indicated that coverages were above 95% for all elevation ranges, with elevation range V being the only range that did not yield 100% coverage (Fig. 3a), suggesting that the community of understory bats in all elevations was adequately sampled. This finding implied that it is likely unnecessary to correct for sample completeness at the lowest coverage, as it did not differ drastically from the highest coverage value. Hill numbers show that taxonomic diversity and richness of species assemblages varied substantially with elevation in CNP (Fig. 3). The comparison of species richness (q0) showed three distinctive groups: the first group which covers three elevation ranges from 700 to 1500 m asl ( I, II and III ) had a similar number of species (q0 = 40, 42 and 42, respectively); the second group, IV, which covers an elevation range from 1500 to 1800 m asl had the highest richness (q0 = 47), and V, the highest elevation range, had the lowest richness (q0 = 22) (Fig. 3b). However, if we take into account the relative abundance values of all species, species diversity showed different patterns. Shannon diversity (q1) showed a decline in diversity with increasing elevation (Fig. 3c), and taking into account the dominant species, Simpson diversity (q2), showed that as elevation increases, community assemblages become increasingly dominated by a smaller number Figure 2. Species accumulation curve of bat community of Cusuco National Park, north-west Honduras. of species (Fig. 3d). Moreover, the capture rates showed a similar pattern, yielding a significant decrease with elevational increase (R2 adj = 0.63, N = 36, F = 61.2, df = 34, P < 0.001). Of the six general trophic guilds assessed in CNP, insectivorous species were the most speciose group (27 species, n = 988), while frugivorous species were the most abundant guild (20 species, n = 5049) (Fig. 4a). On average, per year, frugivorous bats accounted for 72% of the captures, mainly because of the high presence of A. jamaicensis, S. hondurensis, Dermanura tolteca, and Carollia sowelli. Carnivorous and omnivorous species were the least abundant and least diverse guilds, each represented by just two species, with fewer than five individuals of each captured per year (Fig. 4a). Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 8 All trophic guilds were present in all the different camps in the park. However, dominance of feeding guilds differed across the Park’s altitudinal gradient. Frugivorous, nectarivorous, and insectivorous bats were present across the entire elevational gradient of CNP. Insectivorous bats, the most speciose guild within CNP, were recorded along the entire elevational gradient but were mainly captured at higher altitudes. Sanguivorous bats were mainly captured at lower elevations (buffer zone) close to the villages, whereas carnivorous species were mainly captured in the core zone at elevations higher than 1300 m asl (Fig. 4b). Species accounts Major range extensions Eptesicus brasiliensis Desmarest, 1819: Twenty-seven individuals were caught: 21 adults, five juveniles, and one bat whose age and sex were not recorded. Of those, 16 were females and 10 were males. Sixty-nine percent of the females showed signs of reproductive activity (four were pregnant, three lactating and two post-lactating). Sixty-three percent of the males caught had the testes Figure 3. Species diversity based on Hill numbers from bat assemblages of Cusuco National park, north-west Honduras. a) Sample completeness (as measured by sample coverage) with respect to sample size. The dot marks the highest coverage achieved in our samples. b) Species diversity based on species richness (q = 0), c) Shannon diversity (q = 1) and d) Simpson diversity (q = 2) across elevations ranges in Cusuco National Park. The vertical lines represent the bootstrapped 95% confidence intervals. Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 9 descendent. Eptesicus brasiliensis was trapped in all elevation ranges studied but mainly captured between 1,300 and 1,500 m asl. This represented the second official record of this bat species in Honduras. Previously, there was only one record in Choluteca, southern Honduras at 1,082 m asl (Espinal and Mora 2012). Natalus lanatus Tejedor, 2005: Only two male juveniles of this species were captured within CNP. The first capture was on July, 21, 2015 at 19:15, and the second individual was captured on June 29, 2019 at 00:30. This species has been recorded in Costa Rica (Rodríguez-Herrera et al. 2011) and Nicaragua (Medina-Fitoria et al. 2015). The presence of the species in Honduras has been recently confirmed through the re-examination of two specimens captured in 1963 in Francisco Morazán and La Paz (Turcios-Casco et al. 2020). Here, we corroborate the presence of this species in Honduras and demonstrate its range extends to the north-west of the country. Additionally, in CNP, the individuals were captured at an elevation of 1,650 m asl. So far, the species was only known from lowland to middle elevation (1,200 m asl) in México and tropical rainforest in Nicaragua (1,200 m asl) and Costa Rica (1,405 m asl). The individuals showed the characteristic bicolour ventral long hairs at base of claws of N. lanatus (Rodríguez-Herrera et al. 2011, Tejedor 2005).This insectivorous species was described in 2005 (Tejedor 2005); however, López-Wilchis et al. (2012) concluded that there is not enough evidence to separate the N. lanatus individuals from N. mexicanus. High elevational range records Centurio senex Gray, 1842: A total of 316 individuals were captured. Within the adults, 171 were females, with 76% showing signs of reproductive activity (60 pregnant, 19 lactating, and 51 post lactating). Regarding the males, 86 were adults, of which 44% had the testes descendent. Centurio senex was reported at 1650 m asl (Timm and LaVal 2018) in Costa Rica; however, we captured this species in CNP at 1950 m asl (18 individuals). This species was widespread within the park and was found in a range of habitats including closed-canopy forest, coffee plantations, and humaninduced forest clearings. Occasionally large numbers were caught in a short period of time (up to 20 individuals per night). This was probably due to our nets being in close proximity to temporal feeding resources, ssuch as fruiting palms and figs. However, this requires further micro-habitat assessments to confirm. Within Honduras, this species has been captured in La Esperanza, Chichicaste Figure 4. Trophic guilds within the bat community of Cusuco National Park, north-west Honduras. a) Number of individual bats within each feeding guild captured per year. Mean (± SD) of bats captured depending on their trophic guilds. Inset Pie chart displays the number of species belonging to a trophic guild. b) Altitudinal distribution of bats according to their trophic guild. Violin chart show the distribution shape of the data where width of columns indicates the proportion of a guild found at each altitudinal level. Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 10 (LaVal 1969), Choluteca (Valdez and LaVal 1971) and Francisco Morazán (McCarthy et al. 1993). Myotis albescens (Geoffroy, 1806): A total of 18 individuals were captured in CNP. Apart from one female juvenile, all of the individuals were male, of which 50% had the testes descendent. This species has been reported to be maximal 1500 m asl in Mexico but in CNP was been captured mainly at 1600 m asl. This species has been previously reported in Choluteca (McCarthy 1993), and Santa Bárbara (Turcios-Casco et al. 2019). Platyrrhinus helleri (Peters, 1866): A total of 23 individuals were captured in CNP. Of the adults, 14 females, 86% showed signs of reproductive activity (six pregnant, three lactating and three postlactating), while all adult males (three) had the testes descendent. This species was captured between 750 to 1600 m asl, but mainly at 1,000 m asl. In Honduras, it has been previously reported in Comayagua (LaVal 1969), Atlantida (Dick 2013), and Gracias a Dios (Medina-Fitoria and Turcios-Casco 2019). Threatened species Bauerus dubiaquercus Van Gelder, 1959 (IUCN Near Threatened): A total of 118 individuals were caught in CNP, primarily during the first two hours after sunset (83% of the captures). Of the adults, 58 were males; 76% of these had the testes descendent. Among the females, 54 were adults, and 74% showed signs of reproductive activity (nine were pregnant, nine lactating, and 22 post-lactating). Bauerus dubiaquercus was trapped between 700 and 1950 m asl, with 40% of the bats caught between 1600 and 1700 m asl. This species was primarily captured close to rivers and streams within closed-canopy forest habitats within the core zone of the park and rarely recorded in human-induced clearings or coffee plantations. This indicates a certain level of intolerance towards disturbed habitat types. The only other records of this species in Honduras are in Olancho (Graciolli and Dick 2009, Pine et al. 1971). Hylonycteris underwoodi Thomas 1903 (IUCN Least Concern, PCMH National Threatened): A total of 112 individuals were caught. Within the 37 adult females, 29 showed signs of reproductive activity (21 were pregnant, four lactating and four post-lactating). A total of 64 males were caught, of which 39 had the testes descendent. Hylonycteris underwoodi was trapped between 750 and 1900 m asl, with 44% of the bats trapped at 1400 m asl. Within Honduras, the species has been also recorded in Atlántida (Merida and Cruz, 2014), Olancho (Portillo-Reyes et al. 2016), and Gracias a Dios (Turcios-Casco and Medina-Fitoria 2018). Discussion The results of our surveys produced what we believe to be the most complete description of a bat community in a northern Central American cloud forest to date. Given that the known chiropteran richness of Honduras is 114 species (Turcios-Casco et al. 2020, Siles and Baker 2020), CNP supports at least 52% of all species found nationally in just c. 0.2% of its land area, and 34.7% of the 170 species known to occur in Central America (Rodríguez-Herrera and Sánchez 2015). The finding that the lower-lying areas of the Park support the highest diversity of bat species, with richness decreasing with elevation, conforms with general theories regarding species turnovers across altitudinal gradients (e.g. Nunes et al. 2016). Our results also contribute records regarding both the spatial range and altitudinal delimitations of bat species. We reaffirm the presence of Natalus lanatus in Honduras, and report a major extension range for Eptesicus brasilienses, both species having been previously reported from southern Honduras (Espinal and Mora 2012, Turcios- Casco et al. 2020). Additionally, our data provides high-altitude records for three species. All these findings highlight CNP as an important area for conservation priority. In Honduras, four areas for priority conservation of bats have been described (Hernández 2015). CNP is included in one of those areas (Gulf of Honduras), but specific information about Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 11 Table 2. Summary of bat species detected in Cusuco National Park, north-west Honduras, from 2006-2019. Taxonomy follows IUCN (2019), Mantilla-Meluk (2014), Mantilla-Meluk and Muñoz-Garay (2014), Baird et al. (2015), Tejedor (2005), Velazco and Patterson (2013) and York et al. (2019). Species marked * are regionally endemic to the Mesoamerican biodiversity hotspot as defined by Myers et al. (2000). Species indicated † are assessed as near-threatened by the IUCN (2019) and species indicated Ω are assessed as nationally near-threatened within Honduras by Hernandez (2015). Species marked in bold represent major range extensions and underscored species represent minor range. Letters provided in parenthesis after each species name indicates primary dietary guild as defined by Kalko et al. (1996), Reid (2009) and Soriano (2000): C = carnivorous, F = frugivorous, I = insectivorous, N = nectivorous, O = omnivorous, S = sanguivorous. Forearm and weight values provided are the mean values of all adult individuals measured, with standard deviation values to two decimal places. Sample size is given in parenthesis. Pregnant individuals were excluded. Juvenile measurements were excluded in all cases where sample sizes included less than10 adults, otherwise marked as ×. N/A indicates that no individuals of the corresponding sex were caught for that species. Relative abundance estimates are denoted as follows: A= Abundant (captured more than 200 times) C = common (captured between 101 and 200 times); U = uncommon (captured between 21 and 100 times); R = rare (captured less than 20 times). Min and max elevation display the lowest and highest points each species was detected in the study area. Elevations marked in bold represent species captured at higher elevation than previously reported. Known max elevation shows the maximum elevation each species is indicated to occur in the literature (IUCN 2019; Timm and LaVal, 2018 b, Almazán-Núñez et al. 2018c and Sánchez-Cordero 2001d). Minimum known elevations for each species are not shown as none of our records in Cusuco National Park were lower than indicated in the literature. Family Scientific name Forearm ♀ (mm) Forearm ♂ (mm) Mass ♀ (g) Mass ♂ (g) Captures Min elevation in CNP Max elevation in CNP Known max elevation DESMODONTINAE Desmodus rotundus (S) 60.93 ± 1.43 (49) 56.87 ± 2.73 (94) 35.66 ± 5.65 (45) 31.12 ± 3.48 (89) A (201) 700 1700 2700 Diphylla ecaudata (S) 56.68 ± 1.92 (6) 54.6 ± 2.36 (7) 34.17 ± 1.94 (6) 28.83 ± 6.91 (6) U (21) 1400 1900 1900 GLOSSOPHAGINAE Anoura geoffroyi (N) 43.42 ± 1.52 (6)× 43.47 ± 0.94 (7)× 15.33 ± 1.03 (6)× 15.86 ± 1.35 (7)× R (13) 750 1900 2500 Choeroniscus godmani (N) 33.95 ± 1.21 (6) 33.79 ± 1.24 (21) 9.8 ± 3.27 (5) 10.55 ± 2.75 (21) U (55) 700 1700 1650b Glossophaga commissarisi (N) 34.81 ± 1.22 (64) 34.19 ± 1.23 (56) 10.06 ± 2.01 (39) 10.18 ± 1.95 (53) C (157) 750 1700 2000 Glossophaga leachii * (N) 36.36 ± 2.34 (19) 34.78 ± 1.01 (10) 11.19 ± 4.42 (8) 9.6 ± 1.26 (10) U (39) 750 1700 2400 Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 12 Glossophaga soricina (N) 35.91 ± 1.21 (87) 35.06 ± 1.21 (62) 11.01 ± 1.73 (37) 10.22 ± 2.19 (59) A (212) 700 1700 2600 Hylonycteris underwoodi*Ω (N) 34.67 ± 2.22 (40) 33.45 ± 1.18 (54) 10.84 ± 2.39 (19) 8.35 ± 1.12 (56) C (113) 750 1900 2600 STENODERMATINAE Artibeus jamaicensis (F) 61.61 ± 1.79 (296) 61.22 ± 1.93 (421) 47.73 ± 7.24 (196) 45.86 ± 4.42 (419) A (1130) 700 1700 1700 Artibeus lituratus (F) 67.39 ± 4.25 (35) 67.73 ± 3.92 (10) 63.83 ± 11.88 (20) 60.33 ± 10.43 (9) U (53) 700 1650 1700 Centurio senex (F) 43.03 ± 1.22(171) 42.99 ± 1.24 (86) 21.01 ± 2.78 (111) 21.49 ± 2.48 (85) A (316) 750 1950 1650b Chiroderma salvini (F) 49.15 ± 2.15 (17) 48.84 ± 1.28 (17) 30.7 ± 3.47 (10) 28.1 ± 2.04 (15) U (48) 750 1650 1700 Chiroderma villosum (F) 45.36 ± 2.27 (10) 43.93 ± 0.12 (3) 25.22 ± 4.94 (9) 17.5 ± 6.71 (2) R (13) 1000 1400 1700c Dermanura azteca (F) 42.06 ± 1.02 (30) 40.78 ± 1.07 (16) 17.41 ± 3.26 (17) 17.28 ± 3.32 (16) U (71) 800 1950 3000 Dermanura phaeotis (F) 40.15 ± 1.94 (30) 39.09 ± 2.82 (13) 16.08 ± 2.69 (26) 14.93 ± 3.27 (14) U (55) 700 1700 1650b Dermanura tolteca (F) 40.89 ± 1.1 (204) 40.43 ± 1.46(281) 17.55 ± 3.76(145) 15.95 ± 2.46 (282) A (779) 700 1950 1850b Dermanura watsoni (F) 40.29 ± 1.15 (26) 40.27 ± 1.04 (61) 18.25 ± 3.76 (18) 14.93 ± 2.69 (58) C (146) 700 1850 1850b Enchisthenes hartii (F) 39.89 ± 1.64 (50) 38.63 ± 2.75 (23) 18.24 ± 2.86 (40) 17.13 ± 3.47 (23) U (85) 750 1950 2600 Platyrrhinus helleri (F) 39.36 ± 0.86 (14) 38.93 ± 0.12 (3) 18.13 ± 1.89 (8) 15 ± 1 (3) U (23) 700 1600 1500 Sturnira hondurensis * (F) 44.62 ± 1.48 (397) 44.86 ± 1.82 (257) 23.44 ± 3.16 (347) 25.42 ± 3.49 (260) A (1018) 700 1950 2000 Table 2 continued. Family Scientific name Forearm ♀ (mm) Forearm ♂ (mm) Mass ♀ (g) Mass ♂ (g) Captures Min elevation in CNP Max elevation in CNP Known max elevation Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 13 Sturnira parvidens * (F) 39.42 ± 1.83(150) 39.73 ± 1.72 (100) 17.51 ± 2.75 (128) 18.42 ± 3.42 (100) A (406) 700 1950 2450d Uroderma convexum (F) 44 ± 2.16 (4) × 43 (1) × 20 ± 2.45 (4) × 19 ± 1.41 (2) × R (7) 750 1000 1500 Vampyressa thyone (F) 32.08 ± 0.88 (14) 33.04 ± 3.28(10) 10.44 ± 1.13 (9) 9.7 ± 2.66 (10) U (37) 800 1450 1500 Vampyrodes major (F) 55.77 ± 2.6 (27) 54.19 ± 2.27 (31) 39.53 ± 4.79 (15) 36.47 ± 5.02 (31) U (63) 700 1650 1600b CAROLLINAE Carollia castanea (F) 38.95 ± 3.46 (12) 36.24 ± 1.38 (17) 18.91 ± 7.71 (11) 14.83 ± 1.92 (18) U (51) 700 1300 1500b Carollia perspicillata (F) 44.22 ± 1.84 (64) 44.26 ± 1.54 (53) 22.05 ± 3.24 (55) 23.39 ± 3.21 (54) C (197) 700 1300 1650b Carollia sowelli * (F) 40.62 ± 1.63 (201) 40.64 ± 2.63 (189) 18.13 ± 3.28 (172) 18.91 ± 4.23 (173) C (521) 700 1700 1700 Carollia subrufa * (F) 40.67 ± 1.15 (3) 39.50 ± 0.70 (2) 15 ± 1.41 (2) 14.5 ± 4.95 (2) (5) 1200 PHYLLOSTOMINAE Chrotopterus auritus (C) 83 ± 1.8 (3) 82.63 ± 2.25 (7) > 100 (2) > 100 (7) R (11) 1000 1400 2000 Lonchorhina aurita (F) 50.5 ± 0.71 (2) N/A 15 (1) N/A R (2) 1400 1600 1600 Lophostoma brasiliense (F) 35.55 ± 0.64 (2) 36.7 ± 0.42 (2) 9 ± 1.41 (2) 12.5 ± 2.12 (2) R (5) 700 1250 1300b Micronycteris microtis (I) 34.67 ± 2.31 (3) 35.65 ± 0.83 (10) 7 ± 1.73 (3) 8.3 ± 0.92 (10) R (17) 700 1700 2600 Micronycteris minuta (I) 34.35 ± 0.49 (2) 35.5 ± 1.32 (3) 9.5 ± 4.95 (2) 9.33 ± 3.21(3) R (6) 750 800 800 Table 2 continued. Family Scientific name Forearm ♀ (mm) Forearm ♂ (mm) Mass ♀ (g) Mass ♂ (g) Captures Min elevation in CNP Max elevation in CNP Known max elevation Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 14 Micronycteris schmidtorum (I) 38 ± 1.41 (2) 34.57 ± 2.37 (7) 8 (1) 8.58 ± 2.42 (6) R (9) 1000 1650 1600b Mimon cozumelae (F) 58 (1) 56.4 ± 0.85 (2) 22 (1) 36 ± 16.97 (2) R (3) 1000 1000 1300b Phyllostomus discolor (O) 64.5(1) N/A 42.5 (1) N/A R (1) 1050 1050 1300b Phyllostomus hastatus (O) 91.56 ± 1.25 (7) 91.9 ± 0.14 (2) > 100 (7) > 100 (2) R (9) 1000 1600 1600b Trachops cirrhosus (C) 58.7 ± 1.4 (5) 58.65 ± 1.76 (12) 31.5 ± 0.71 (2) 35 ± 3.64 (12) R (18) 1250 1650 1650b VESPERTILIONIDAE Bauerus dubiaquercus † * (I) 53.6 ± 2.51 (53) 53.16 ± 2.07 (53) 20.09 ± 3.3 (45) 19.04 ± 2.39 (53) C (118) 700 1950 2300 Eptesicus brasiliensis (I) 42.45 ± 1.77 (13) 42.73 ± 0.83 (8) 10.63 ± 0.74 (8) 11.81 ± 3.64 (8) U (27) 750 1950 3000 Eptesicus furinalis (I) 41.28 ± 0.95 (7) 40.83 ± 1.88 (8) 9.92 ± 1.11 (6) 10.21 ± 0.7 (7) U (22) 1000 1650 1800 Eptesicus fuscus (I) 50 ± 3.63 (3) 49.73 ± 3.18 (4) 16.5 ± 4.82 (3) 18.13 ± 4.55 (4) R (7) 1050 1950 2700 Lasiurus frantzii (I) N/A 38.67 ± 1.15 (3) N/A 8.33 ± 0.58 (3) R (3) 1600 1650 2500 Lasiurus ega (I) 46 (1) 45 (1) 12 (1) 13 (1) R (2) 1600 1650 1650b Myotis albescens (I) 37.7 (1) 35.47 ± 1.69 (11) N/A 5.45 ± 1.57 (11) R (18) 700 1600 1500 Myotis pilosatibialis (I) 36.87 ± 1.09 (218) 36.11 ± 1.19 (169) 5.34 ± 1.47 (203) 5.24 ± 1.76 (167) A (513) 700 1950 2500 Myotis nigricans (I) 37.3 ± 1.57 (4) 36.07 ± 1.28 (15) 4.75 ± 0.5 (4) 5.17 ± 0.82 (15) U (22) 750 1900 3150 Table 2 continued. Family Scientific name Forearm ♀ (mm) Forearm ♂ (mm) Mass ♀ (g) Mass ♂ (g) Captures Min elevation in CNP Max elevation in CNP Known max elevation Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 15 Myotis velifer (I) 42.73 ± 1.32 (4) N/A 9.63 ± 2.06 (4) N/A R (5) 750 1950 3300 Perimyotis subflavus (I) 35.5 (1) 32 (1) 5 ± 1.41 (2) 6 (2) R (4) 800 1650 2600 MOLOSSIDAE Molossus rufus (I) 50.89 ± 1.22 (72) 51.25 ± 2.14 (12) 39 ± 3.9 (64) 41.08 ± 4.98 (12) U (92) 800 1600 1500 Molossus sinaloe (I) 46.83 ± 1.01 (4) N/A 21 ± 2 (4) N/A R (4) 800 800 2400 MORMOOPIDAE Mormoops megalophylla (I) 54.5 ± 0.71 (2) 55.2 ± 1.13 (2) 16.5 ± 2.12 (2) 16.5 ± 0.71 (2) R (4) 1600 1650 2300 Pteronotus davyi (I) 46.38 ± 1.02 (15) 46.63 ± 1.53 (9) 9.54 ± 0.88 (13) 9 ± 1.6 (9) U (25) 1000 1950 2300 Pteronotus gymnonotus (I) N/A 53.5 (1) N/A 14 (1) R (1) 1600 1600 1600 Pteronotus mesoamericanus * (I) 59.6 ± 1.17 (35) 59.26 ± 1.39 (24) 22.59 ± 3.22 (28) 22.6 ± 1.75(23) U (65) 700 1950 2200 Pteronotus personatus (I) 45(1) N/A 11 (1) N/A R (1) 1650 1650 1600b NATALIDAE Natalus lanatus (I) N/A 41 (2)× N/A 8 (2) × R (3) 1650 1700 2400 Natalus mexicanus (I) 39 (1) 37.5 ± 2.12(2) × 8 (1) 7.5 ± 0.71 (2)× R (2) 1400 1650 2400 THYROPTERIDAE Thyroptera tricolor (I) N/A 35.20 (1)× N/A 16 (1)× R (1) 800 800 1300 Table 2 continued. Family Scientific name Forearm ♀ (mm) Forearm ♂ (mm) Mass ♀ (g) Mass ♂ (g) Captures Min elevation in CNP Max elevation in CNP Known max elevation Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 16 the bat assemblage here remained until now unpublished. From these described areas, La Mosquitia National Park and Fonseca Gulf harbour 58 and 49 bat species, respectively (Hernández 2015), which means that when taking our results into account, CNP possesses the most diverse chiropteran community in Honduras. This is a result of the strong altitudinal gradient and associated diversity of habitats found in CNP. However, it is likely that other areas in Honduras, which at present remain poorly explored, have similar or greater bat diversity than CNP, which reinforces the importance of long-term monitoring surveys within the country. Since studies of bat diversity in this area are scarce, several of the species reported here are the first records for the area. Moreover, some species have been reported only a few times in the country. For example, Micronycteris minuta has been reported once in Atlántida department (McCarthy et al.1993) and has recently been reported in Guatemala, about 35 km from CNP (Pérez-Consuegra et al. 2017). According to geographical range maps in IUCN (2019) and Reid (2009), Enchisthenes hartii, Myotis velifer, and Dermanura azteca present a minor extension range. These species have been previously reported in El Paraiso, La Esperanza, and Francisco Morazán departments in southern Honduras (Goodwin 1942, LaVal 1969, McCarthy et al. 1993), and apparently these species occur primarily at higher elevations (McCarthy et al. 1993). Generating more information about the diversity of bats in Honduras could greatly help to identify more conservation priority areas and improve regional conservation strategies. Reports of new country records and major extension ranges in Honduras have been increasing in the last decades (e.g., Espinal and Mora 2012, Medina-Fitoria and Turcios-Casco 2019, Mora and Lopez 2010, Turcios-Casco et al. 2019, 2020). Here we corroborate the presence of Natalus lanatus in Honduras, expanding both its known spatial and elevation range. Our records of E. brasiliensis in the Park also provide a major extension range of this species in Honduras. Numerous novel altitudinal records have also been reported for other taxa within CNP, most notably in birds (Martin et al. 2016). The frequency of these records may, in part, be due to the lack of scientific research completed in Mesoamerican cloud forests (Bubb et al. 2004), highlighting the need for further fieldwork in these ecosystems, although as >80% of species detected in CNP are not Mesoamerican endemics, this does not offer a complete explanation. The finding that some bat species are found in CNP at higher altitudes than reported elsewhere could also relate to the Park’s proximity to the Caribbean coast, where the mild maritime climate and the influence of the Alizé trade winds create comparatively warm, humid conditions at middle and high altitudes in CNP (Martin et al. 2016). These climatic conditions could allow some species to persist at higher elevations than is possible in the cooler, drier non-coastal mountains. Another hypothesis could be that the altitudinal records in our dataset are due to the effects of climate change facilitating upslope elevational shifts in the bat communities here (LaVal 2004). Indeed, such upslope movements have been previously documented in CNP’s bird community (Neate-Clegg et al. 2019). Such upslope movements may ultimately have serious conservation consequences for cloud-forest endemics, representing an “escalator to extinction” (Marris 2007) where the extent of suitable habitats for these species becomes progressively limited. Both bat diversity and capture rate showed a negative correlation with elevation, where the diversity was higher at lower elevations. Patterns of a greater richness at low and mid-elevations have been reported in bats (Carvalho et al. 2019, Cisneiros et al. 2014), possibly due to higher food availability. Moreover, forest assemblages at higher elevations have more variable abundances and are numerically dominated by a small number of species. The highest elevational range we examined was dominated by few species, particularly S. hondurensis, Myotis pilosatibialis and Dermanura tolteca (73%), species reported as common at high elevation. In some cases, even small changes in elevation affected the assemblages (Capaverde et al. 2018). Nevertheless, bat assemblages cannot be completely explained by elevation. Several studies have reported that variation in temperature, Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 17 precipitation, habitat structure, floristic composition, food availability, and roost areas are some of the factors that also play an important role in the structure and maintenance of particular species in an area (Arita 1993, Cisneros et al. 2014, McCain and Grytnes 2010, Oliveira et al. 2018). Moreover, all these factors can vary among years, causing movements among the bat species, not only between seasons but also between years. Frugivores constituted the trophic guild best represented in our bat assemblages and were present in all elevational ranges of the survey; however, they were mainly found at lower elevations (>1300 m asl). Only seven species of frugivorous bats (out of 20 species) were captured at the highest elevation of the study (1900 m asl). Insectivorous species were, in general, broadly distributed in all elevational ranges in CNP but mainly occurred at higher altitudes, primarily due to the large number of captures of M. pilosatibialis. Both patterns were found in several studies in the neotropics (e.g. Calderon et al. 2013, Castaño et al. 2018, Pineda et al. 2005, Sanchez-Cordero 2001). However, new molossid and emballonurid species are particularly likely to be detected if more surveys were completed in the lowlands using acoustic recorders. Similarly, nectivorous species were broadly distributed in all elevation ranges; however, only Anoura geoffroyi and H. underwoodi were present in all elevation ranges. Anoura geoffroyi is a species with a broad occupancy of altitudinal habitats and is common in high elevations (Calvalho et al. 2019), but was rarely captured in CNP, whereas H. underwoodi is a species commonly captured in the park, but rarely captured in other areas in Honduras (Turcios-Casco and Medina-Fitoria 2018). On the other hand, the sanguivorous species, Desmodus rotundus, was most likely concentrated at lower altitudes due to these predominately agricultural landscapes supporting large numbers of livestock on which these species feed, whereas Diphylla ecaudata was mainly captured in the core-zone restricted by the availability of blood from wild birds (Greenhall et al. 1984, Uieda 1994). The finding that carnivorous species, such as Chrotopterus auritus and Trachops cirrhosus, are present only in the core zone but not in higher elevations of the Park is probably because these sections of CNP are still predominately comprised of intact, closed-canopy forest habitats and thus provide a higher abundance of prey (amphibians, birds, rodents, etc) (Soriano 2000), corroborating the observation that they are more abundant in undisturbed than disturbed areas (Fenton et al. 1992, Medellín et al. 2000). Although all analyses were corrected by sampling effort, it should be highlighted that the particularly high diversity at Base camp may still be partially a result of significant sampling biases, given more trapping nights were completed here than any other site. Increasing the number of surveys increases not only the number of captures but also the possibility of recording rare species. On the other hand, species such as Phyllostomus hastatus, T. cirrhosus, and Micronycteris schmidtorum, were only recorded after ten years of surveys in this camp. These species are generally documented at lower elevations. Thus, the greater species richness at this site may also be due to the fact that lowland species may be starting to colonize higher elevation habitats (LaVal 2004); however, further research is needed for drawing conclusions. While the survey effort underpinning the results in this study is extensive, with an inventory completeness estimated at ca. 95%, further work remains to be conducted in CNP. Future fieldwork is particularly encouraged within the lower-lying (500-900 m) sections of the Park, as this elevation band is likely to represent the most species-rich part of CNP, yet is where we invested the least fieldwork effort. The cultivated agricultural habitats and remnant patches of lowland forest may well support new species for the Park, which to date have remained unrecorded by our surveys. It is also possible that our focus on mist-netting methodologies may have led to several high-flying, insectivorous species remaining undetected in the CNP, these species being difficult to capture in nets (Kalko and Handley 2001, MacSwiney et al. 2008). Further work using acoustical recorders, which are effective in detecting these species, may yield more species records for the Park, since some species have not been caught despite being potentially present, such as members of the family Neotropical Naturalist P. Medina-Van Berkum, K. Vulinec, D. Crace, Z. Lopez Gallego, and T. E. Martin 2020 No. 3 18 Emballonuridae and Molossidae. The introduction of canopy nets and active searches for roost sites may also allow for the detection of species rarely captured in understory mist-nest. However, such approaches are not always practical due to limitations in expertise, time requirements, and the difficulties of working in rugged terrain (Kalko and Hadley 2001). Moreover, to better understand the functional ecology of CNP’s bat assemblages, more specific research is needed, including targeted investigations into diet and habitat relationships. In summary, the results we present here represent the most comprehensive survey of a bat community within a northern Central America cloud forest. Our data provides a detailed 14- year overview of bat diversity and community structure within these poorly-studied cloud forest ecosystems, as well as describes how this diversity and structure changes across the elevation gradient of the Park. Our fieldwork has also yielded novel information for species rarely captured in Mesoamerica, as well as range and altitudinal records for several neotropical bat species. It highlights CNP as a regionally important center of bat diversity within the Mesoamerican biodiversity hotspot and an important conservation priority. Acknowledgements This project was supported by Operation Wallacea. We extend thanks to the many members of the bat survey teams who gathered the data used in this paper and the hundreds of Operation Wallacea students who have supported the project since 2006. We also thank Expediciones y Servicios Ambientales de Cusuco for logistical support and Instituto de Conservación Forestal for issuing annual research permits to support our work in Cusuco. KV thanks Delaware State University for support. We are especially grateful to our local guides for their assistance and input, in particular Joxan Orlando Ayala. Finally, we thank two anonymous reviewers for their constructive comments on an earlier version of the manuscript. Literature Cited Almazán-Núñez, R.C., E.A. Alvarez-Alvarez, F. Ruiz-Gutiérrez, P. Sierra-Morales, A. 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