Can wing morphology inform conservation priorities for Southeast Asian cave bats?

Adaptations for foraging in the complex airspaces of forest interiors may make bat species in the Asian tropics particularly susceptible to forest loss. However, ecomorphological analysis of Vietnamese bat assemblages challenges the hypothesis that, due to their greater vagility, cave‐roosting bats are less vulnerable to habitat fragmentation than foliage‐roosting species. Of the 13 most highly adapted forest‐interior species in our study, eight were cave‐roosting members of the Rhinolophidae and Hipposideridae and had wing morphologies closely resembling five foliage‐roosting members of the Murininae and Kerivoulinae—species typically thought to have low vagility. Overall, both cave‐roosting and foliage‐roosting bats exhibited a wide range of flight indices and species' wing designs corresponded with preferred foraging habitats, suggesting that foraging strategy may outweigh roost preference as a determinant of bat wing morphology and flight performance. Consequently, where such variation occurs, cave‐roosting bat ensembles are likely to include species with low vagility and similar sensitivity to habitat fragmentation. This could have important conservation implications as Asian karst formations support high cave densities and important bat diversity yet increasingly represent forest refugia in anthropogenic landscapes. We, therefore, advocate greater consideration of species vagility in determining conservation priorities for the region's bat fauna.     

EFFECT OF HABITAT AND FORAGING HEIGHT ON BAT ACTIVITY IN THE COASTAL PLAIN OF SOUTH CAROLINA

Abstract: We compared bat activity levels in the Coastal Plain of South Carolina among 5 habitat types: forested riparian areas, clearcuts, young pine plantations, mature pine plantations, and pine savannas. We used time-expansion radio-microphones and integrated detectors to simultaneously monitor bat activity at 3 heights (30, 10, 2 m) in each habitat type. Variation in vegetative clutter among sampling heights and among habitat types allowed us to examine the differential effect of forest vegetation on the spatial activity patterns of clutter-adapted and open-adapted bat species. Moreover, monitoring activity at 30, 10, and 2 m permitted us to also compare bat activity above and below the forest canopy. We detected calls of 5 species or species groups: eastern red/Seminole bats (Lasiurus borealis/L. seminolus), eastern pipistrelles (Pipistrellus subflavus), evening bats (Nycticeius humeralis), big brown bats (Eptesicus fuscus), and hoary bats (Lasiurus cinerius). At 2 and 10 m, bat activity was concentrated in riparian areas, whereas we detected relatively low levels of bat activity in upland habitats at those heights. Activity was more evenly distributed across the landscape at 30 m. Bat activity levels above the forest canopy were almost 3 times greater than within or below the canopy. We detected significantly greater activity levels of 2 open-adapted species (hoary and big brown bats) above rather than within or below the forest canopy. However, activity levels of 2 clutter-adapted species (eastern red/Seminole bats and eastern pipistrelles) did not differ above, within, or below the forest canopy. Despite classification as a clutter-adapted species, evening bat activity was greater above rather than within or below the forest canopy. We believe our results highlight the importance of riparian areas as foraging habitat for bats in pine-dominated landscapes in the southeastern United States. Although acoustical surveys conducted below forest canopies can provide useful information about species composition and relative activity levels of bats that forage in cluttered environments, our results showing activity above canopy suggest that such data may not accurately reflect relative activity of bats adapted to forage in more open conditions, and therefore may provide an inaccurate picture of bat community assemblage and foraging habitat use.     

Palate Variation and Evolution in New World Leaf-Nosed and Old World Fruit Bats (Order Chiroptera)

Two bat families, the leaf-nosed (Phyllostomidae) and fruit bats (Pteropodidae), have independently evolved the ability to consume plant resources. However, despite their similar ages, species richness and the strong selective pressures placed on the evolution of skull shape by plant-based foods, phyllostomids display more craniofacial diversity than pteropodids. In this study, we used morphometrics to investigate the distribution of palate variation and the evolution of palate diversity in these groups. We focused on the palate because evolutionary alterations in palate morphology are thought to underlie much feeding specialization in bats. We hypothesize that the distribution of palate variation differs in phyllostomids and pteropodids, and that the rate of palate evolution is higher in phyllostomids than pteropodids. The results suggest that the overall level of palate integration is higher in adult populations of pteropodids than phyllostomids but that the distribution of palate variation is otherwise generally conserved among phyllostomids and pteropodids. Furthermore, the results are consistent with these differences in palate integration likely having a developmental basis. The results also suggest that palate evolution has occurred significantly more rapidly in phyllostomids than pteropodids. These findings are consistent with a scenario in which the greater integration of the pteropodid palate has limited its evolvability.     

Secondary forest buffers the effects of fragmentation on aerial insectivorous bat species following 30 years of passive forest restoration

Passive forest restoration can buffer the effects of habitat loss on biodiversity. We acoustically surveyed aerial insectivorous bats in a whole-ecosystem fragmentation experiment in the Brazilian Amazon over a 2-year period, across 33 sites, comprising continuous old-growth forest, remnant fragments, and regenerating secondary forest matrix. We analyzed the activity of 10 species/sonotypes to investigate occupancy across habitat types and responses to fragment size and interior-edge-matrix (IEM) disturbance gradients. Employing a multiscale approach, we investigated guild (edge foragers, forest specialists, flexible forest foragers, and open space specialists) and species-level responses to vegetation structure and forest cover, edge, and patch density across six spatial scales (0.5–3 km). We found species-specific habitat occupancy patterns and nuanced responses to fragment size and the IEM disturbance gradient. For example, Furipterus horrens had lower activity in secondary forest sites and the interior and edge of the smallest fragments (1 and 10 ha) compared to continuous forest, and only two species (Pteronotus spp.) showed no habitat preference and no significant responses across the IEM and fragment size gradients. Only the Molossus sonotype responded negatively to vegetation structure. We uncovered no negative influence of forest cover or edge density at guild or species-level. Our results indicate that reforestation can buffer the negative effects of fragmentation and although these effects can still be detected in some species, generally aerial insectivorous bats appear to be in recovery after 30 years of passive forest restoration. Our findings reinforce the need to protect regenerating forests while conserving vast expanses of old-growth forest.     

Sources of Assimilated Proteins in Old and New World Phytophagous Bats

Dietary proteins are considered crucial for growth and maintenance in mammals, but many fruit‐eating mammals feed largely on a protein‐poor diet. In the Chiroptera, frugivory has evolved twice, in the Old World Pteropodidae and in the New World Phyllostomidae, especially the Stenodermatinae. Recent studies based on the analysis of nitrogen isotope ratios (δ¹⁵N) suggest that phyllostomids feed to varying degrees on arthropods to meet their nitrogen (N) requirements. Arthropod feeding has rarely been observed in Pteropodidae. Thus, we asked whether pteropodids meet their N requirements by feeding exclusively on plant matter. We predicted that tissue from pteropodid wing membranes should be depleted in ¹⁵N relative to those of obligate insectivorous rhinolophoid, vespertilionid or emballonurid bats, if pteropodids acquire proteins exclusively from fruits, leaves or nectar. We found that δ¹⁵N in pteropodids were significantly lower than in obligate insectivorous bats. In addition, mean δ¹⁵N of Old World pteropodids was similar to that of obligate frugivorous stenodermatines of the New World tropics. We infer from these data that pteropodids are predominantly phytophagous bats. From a nutritional perspective, pteropodids and stenodermatines are very similar, suggesting that they share convergent physiological adaptations to compensate for the lack of dietary nitrogen.     

Use of Forest Edges by Bats in a Managed Pine Forest Landscape

ABSTRACT Forest edges often have increased species richness and abundance (edge effect) and affect spatial behaviors of species and dynamics of species interactions. Landscapes of intensively managed pine (Pinus spp.) stands are characterized by a mosaic of patches and linear forest edges. Managed pine forests are a primary landscape feature of the southeastern United States, but the effects of intensive management on bat communities are poorly understood. Insectivorous bats are important top predators in nocturnal forest food webs. We examined bat foraging behavior along forest edges and in 4 structurally distinct stand types (open-canopy pine, prethinned pine, thinned pine, and unmanaged forest) within a managed pine forest in the coastal plain of North Carolina, USA. During May-August, 2006 and 2007, we recorded echolocation calls using Pettersson D240X bat detectors linked to digital recorders at 156 sites. We also sampled nocturnal flying insects at each site using Malaise insect traps. We used negative binomial count regression models to describe bat foraging behavior relative to forest edges, stand types, and prey availability. Although some species showed affinities for certain stand types and prey items, bat activity patterns were most strongly related to forest edges. Edges were used extensively by 6 aerial-hunting bat species, but avoided by Myotis species. Forest edges function similarly to natural forest gaps, by providing foraging opportunities for aerial-hunting bat species. Therefore, the maintenance of forest edges in managed pine landscapes may enhance foraging habitat for aerial-hunting bat species.     

Absorption of visible spectrum radiation by the wing membranes of living pteropodid bats

The wing membranes of bats present a large surface area upon which radiation might be taken up, increasing heat load to the animals. This, combined with the high amount of heat produced during flight, has been advanced as one hypothesis explaining the fact that bats are almost exclusively nocturnal. The proportion of short-wave (visible) radiation absorbed by bat wing membrane has previously been measured at between 0.7 and 0.92. These measurements were made on pieces of membrane taken from the wings of dead, mainly insectivorous bats from temperate regions. Here we examined the amount of light transmitted through and reflected off the wing membranes of four species of live pteropodid bats. There were significant differences in wing reflection between species. At 0.68, the average proportion of light absorbed into the wing membranes was lower than previously reported. This might be because we worked with live animals or because ours were tropical bats which are routinely exposed to tropical sun when roosting. Variation in wing tension strongly affected light absorption. It was predicted that the relaxed state of wing membrane through part of the wing beat cycle would increase the absorption of light into the wings of day-flying bats. The proportion of light absorbed into wings was shown to be an important factor in the heat balance of hypothetical bats flying during the day. Our results raise the predicted temperature at which bats flying during the day might experience hyperthermia by approximately 2 °C and suggest that variation in albedo of wings between species may make some species more susceptible to overheating than others. Accepted: 6 December 1998     

Trapped in the darkness of the night: thermal and energetic constraints of daylight flight in bats

Bats are one of the most successful mammalian groups, even though their foraging activities are restricted to the hours of twilight and night-time. Some studies suggested that bats became nocturnal because of overheating when flying in daylight. This is because--in contrast to feathered wings of birds--dark and naked wing membranes of bats efficiently absorb short-wave solar radiation. We hypothesized that bats face elevated flight costs during daylight flights, since we expected them to alter wing-beat kinematics to reduce heat load by solar radiation. To test this assumption, we measured metabolic rate and body temperature during short flights in the tropical short-tailed fruit bat Carollia perspicillata at night and during the day. Core body temperature of flying bats differed by no more than 2°C between night and daytime flights, whereas mass-specific CO(2) production rates were higher by 15 per cent during daytime. We conclude that increased flight costs only render diurnal bat flights profitable when the relative energy gain during daytime is high and risk of predation is low. Ancestral bats possibly have evolved dark-skinned wing membranes to reduce nocturnal predation, but a low degree of reflectance of wing membranes made them also prone to overheating and elevated energy costs during daylight flights. In consequence, bats may have become trapped in the darkness of the night once dark-skinned wing membranes had evolved.     

Evening Bat Summer Roost-Site Selection on a Managed Pine Landscape

ABSTRACT Creation and maintenance of forested corridors to increase landscape heterogeneity has been practiced for decades but is a new concept in intensively managed southern pine (Pinus spp.) forests. Additionally, more information is needed on bat ecology within such forest systems. Therefore, we examined summer roost-site selection by evening bats (Nycticeius humeralis) in an intensively managed landscape with forested corridors in southeastern South Carolina, USA, 2003–2006. We radiotracked 53 (26 M, 27 F) adult evening bats to 75 (31 M, 44 F) diurnal roosts. We modeled landscape-level roost-site selection with logistic regression and evaluated models using Akaike's Information Criterion for small samples. Model selection results indicated that mature (≥40 yr) mixed pine-hardwood stands were important roost sites for male and lactating female evening bats. Upland forested corridors, comprised of mature pine or mixed pine-hardwoods, were important roosting habitats for males and, to a lesser extent, lactating females. Male roosts were farther from open stands and lactating female roosts were farther from mid-rotation stands than randomly selected structures. Our results suggest roost structures (i.e., large trees and snags) in mature forests are important habitat components for evening bats. We recommend maintaining older (>40 yr old) stand conditions in the form of forest stands or corridors across managed landscapes to provide roosting habitat. Furthermore, our results suggest that an understanding of sex-specific roost-site selection is critical for developing comprehensive guidelines for creating and maintaining habitat features beneficial to forest bats.     

Use of the wings in manipulative and suspensory behaviors during feeding by frugivorous bats

Frugivory evolved independently in Old and New World fruit bats (Families Pteropodidae and Phyllostomidae, respectively) and anecdotal reports state that these bats use their wings in different ways for manipulating food items and postural support during feeding. However, these often-cited behavioral differences have not been documented systematically. Here we report observations of manipulative and suspensory behavior collected from 41 individuals representing five phyllostomid and six pteropodid species. During feeding, phyllostomids used both feet to suspend themselves and invariably manipulated food with the wrists and thumbs of both wings. Most pteropodids in our sample used their thumbs for suspension during feeding and none manipulated fruit with their wings. The suspensory and feeding behaviors of pteropodids varied widely and there were significant differences between species. Discrepancies between phyllostomids and pteropodids in the use of the wings during feeding are associated with previously reported differences in wrist morphology. Based on examination of manipulative and suspensory behaviors in a phylogenetic context, we suggest that differences between pteropodids and phyllostomids reflect the distinct ancestral conditions from which these bats evolved.     

Bats partition activity in space and time in a large,heterogeneous landscape

Diverse species assemblages theoretically partition along multiple resource axes to maintain niche separation between all species. Temporal partitioning has received less attention than spatial or dietary partitioning but may facilitate niche separation when species overlap along other resource axes. We conducted a broad‐scale acoustic study of the diverse and heterogeneous Great Smoky Mountains National Park in the Appalachian Mountains. Between 2015 and 2016, we deployed acoustic bat detectors at 50 sites (for a total of 322 survey nights). We examined spatiotemporal patterns of bat activity (by phonic group: Low, Mid, and Myotis) to test the hypothesis that bats partition both space and time. Myotis and Low bats were the most spatially and temporally dissimilar, while Mid bats were more general in their resource use. Low bats were active in early successional openings or low‐elevation forests, near water, and early in the evening. Mid bats were similarly active in all land cover classes, regardless of distance from water, throughout the night. Myotis avoided early successional openings and were active in forested land cover classes, near water, and throughout the night. Myotis and Mid bats did not alter their spatial activity patterns from 2015 to 2016, while Low bats did. We observed disparate temporal activity peaks between phonic groups that varied between years and by land cover class. The temporal separation between phonic groups relaxed from 2015 to 2016, possibly related to changes in the relative abundance of bats or changes in insect abundance or diversity. Temporal separation was more pronounced in the land cover classes that saw greater overall bat activity. These findings support the hypothesis that niche separation in diverse assemblages may occur along multiple resource axes and adds to the growing body of evidence that bats partition their temporal activity.     

Seasonal Shifts in Nocturnal Habitat Use by Coastal Bat Species

Sensitivity of bats to land use change depends on their foraging ecology, which varies among species based on ecomorphological traits. Additionally, because prey availability, vegetative clutter, and temperature change throughout the year, some species may display seasonal shifts in their nocturnal habitat use. In the Coastal Plain of South Carolina, USA, the northern long-eared bat (Myotis septentrionalis), southeastern myotis (Myotis austroriparius), tri-colored bat (Perimyotis subflavus), and northern yellow bat (Lasiurus intermedius) are species of conservation concern that are threatened by habitat loss. Our objective was to identify characteristics of habitat used by these species during their nightly active period and compare use between summer and winter. We conducted acoustic surveys at 125 sites during May–August and at 121 of the same 125 sites December–March 2018 and 2019 in upland forests, bottomland forests, fields, ponds, and salt marsh and used occupancy models to assess habitat use. The northern long-eared bat and southeastern myotis (i.e., myotis bats) used sites that were closer to hardwood stands, pine stands, and fresh water year-round. We did not identify any strong predictors of tri-colored bat habitat use in summer, but during winter they used bottomland forests, fields, and ponds more than salt marsh and upland forests. During summer and winter, northern yellow bats used sites close to fresh water and salt marsh. Additionally, during summer they used fields, ponds, and salt marsh more than upland and bottomland forests, but in winter they used bottomland forests, fields, and ponds more than upland forest and salt marsh. Our results highlight important land cover types for bats in this area (e.g., bottomland forests, ponds, and salt marsh), and that habitat use changes between seasons. Accounting for and understanding how habitat use changes throughout the year will inform managers about how critical habitat features may vary in their importance to bats throughout the year. © 2021 The Wildlife Society.     

Patterns of island occupancy in bats: influences of area and isolation on insular incidence of volant mammals

Aim  The influence of landscape structure on the distribution patterns of bats remains poorly understood for many species. This study investigates the relationship between area and isolation of islands and the probability of occurrence of six bat species to determine whether persistence and immigration abilities vary among bat species and foraging guilds.
Location  Thirty-two islands in the Gulf of California near the Baja California peninsula in north-west Mexico.
Methods  Using logistic regression and Akaike information criterion (AIC) model selection, we compared five a priori models for each of six bat species to explain patterns of island occupancy, including random patterns, minimum area effects, maximum isolation effects, additive area and isolation effects and compensatory area and isolation effects.
Results  Five species of insectivorous bats ( Pipistrellus hesperus , Myotis californicus , Macrotus californicus , Antrozous pallidus and Mormoops megalophylla ) displayed minimum area thresholds on incidence. The probability of occurrence tended to decrease at moderate distances of isolation ( c . 10–15 km) for these species (excepting A. pallidus ). The distributions of two non-insectivorous species ( Leptonycteris curasoae and Myotis vivesi ) were not influenced by island size and isolation.
Main conclusions  Minimum area thresholds on incidence suggest that island occupancy by insectivorous bats may be limited by resource requirements. Islands smaller than 100 ha typically did not support occupancy or use by insectivorous bats, except at minimal isolation distances. Insectivorous bat species may also be more sensitive to moderate levels of habitat isolation in some landscape contexts than previously expected. Our results suggest that differences in foraging habits may have important implications for understanding the distribution patterns of bats.     

Optimizing passive acoustic sampling of bats in forests

Passive acoustic methods are increasingly used in biodiversity research and monitoring programs because they are cost‐effective and permit the collection of large datasets. However, the accuracy of the results depends on the bioacoustic characteristics of the focal taxa and their habitat use. In particular, this applies to bats which exhibit distinct activity patterns in three‐dimensionally structured habitats such as forests. We assessed the performance of 21 acoustic sampling schemes with three temporal sampling patterns and seven sampling designs. Acoustic sampling was performed in 32 forest plots, each containing three microhabitats: forest ground, canopy, and forest gap. We compared bat activity, species richness, and sampling effort using species accumulation curves fitted with the clench equation. In addition, we estimated the sampling costs to undertake the best sampling schemes. We recorded a total of 145,433 echolocation call sequences of 16 bat species. Our results indicated that to generate the best outcome, it was necessary to sample all three microhabitats of a given forest location simultaneously throughout the entire night. Sampling only the forest gaps and the forest ground simultaneously was the second best choice and proved to be a viable alternative when the number of available detectors is limited. When assessing bat species richness at the 1‐km² scale, the implementation of these sampling schemes at three to four forest locations yielded highest labor cost‐benefit ratios but increasing equipment costs. Our study illustrates that multiple passive acoustic sampling schemes require testing based on the target taxa and habitat complexity and should be performed with reference to cost‐benefit ratios. Choosing a standardized and replicated sampling scheme is particularly important to optimize the level of precision in inventories, especially when rare or elusive species are expected.     

Does interspecific competition drive patterns of habitat use in desert bat communities?

Bodies of water are a key foraging habitat for insectivorous bats. Since water is a scarce and limiting resource in arid environments, bodies of open water may have a structuring effect on desert bat communities, resulting in temporal or spatial partitioning of bat activity. Using acoustic monitoring, we studied the spatial and temporal activity patterns of insectivorous bats over desert ponds, and hypothesised that sympatric bat species partition the foraging space above ponds based on interspecific competitive interactions. We used indirect measures of competition (niche overlap and competition coefficients from the regression method) and tested for differences in pond habitat selection and peak activity time over ponds. We examined the effect of changes in the activity of bat species on their potential competitors. We found that interspecific competition affects bat community structure and activity patterns. Competing species partitioned their use of ponds spatially, whereby each species was associated with different pond size and hydroperiod (the number of months a pond holds water) categories, as well as temporally, whereby their activity peaked at different hours of the night. The drying out of temporary ponds increased temporal partitioning over permanent ponds. Differences in the activity of species over ponds in response to the presence or absence of their competitors lend further support to the role of interspecific competition in structuring desert bat communities. We suggest that habitat use and night activity pattern of insectivorous bats in arid environments reflect the trade-offs between selection of preferred pond type or activity time and constraints posed by competitive interactions.     

Use of Fruit Essential Oils to Assist Forest Regeneration by Bats

Frugivorous bats can be attracted with essential oils from ripe chiropterochoric fruit. We evaluated the efficiency of these oils to attract bats in degraded areas within the Atlantic Rain Forest, particularly pasture and agricultural land. We hypothesized that induction units (IUs), each containing a rubber septum impregnated with oil, would have more bat activity than their respective control units (CUs; without the oil). To test this hypothesis we monitored bat flight activity with night‐vision infrared visors in eight IU and CU from August 2006 to July 2007. We also verified the probability of arrival of chiropterochoric seeds by analyzing the diet of bats captured in a neighboring forest area. Our initial hypothesis that units with odor would lead to greater bat activity was confirmed. Results indicated a rich community of fruit‐eating bats, and dietary analysis revealed a huge potential for dispersion of a vast amount of seeds from different plant species at the IU. Although our study does not reveal with certainty which bat species are attracted to the oil, the flying patterns coincide with those described for the foraging behavior of fruit‐eating phyllostomids. Furthermore, the fact that the bats spend more time flying around the odor source compared to flying time around CU suggest an increase in seed rain. Taken together, these results suggest that the use of essential oils from chiropterochoric fruits induces a qualitative and quantitative increase in seed dispersal in areas that otherwise would not be frequently visited by frugivorous bats.     

Larger trees may support larger Indiana bat maternity colonies in a dynamic landscape

Indiana bats (Myotis sodalis), federally listed as endangered, are of management concern in eastern North America. While researchers quantified the habitat affinities of the species throughout the range, few studies have occurred in regions where populations are at high risk for wind energy development and changing climes. Central Illinois, USA, is a dynamic landscape where forest area has been increasing in recent decades (on public and private land) because of changing farming practices and increased habitat protections. The increasing availability of large diameter trees, increasing forest biomass, and changing forest compositions have the potential to influence Indiana bat roost habitat preferences. We assessed Indiana bat maternity roost selection at the tree and forest plot scale to characterize patterns of use in this region from 2017–2018. We predicted that large trees on the landscape would support large colonies of Indiana bats. We located bats in multiple species of trees including elm (Ulmus spp.), cottonwood (Populus deltoides), and shagbark hickory (Carya ovata). We documented larger maternity colonies sharing roosts than in previous studies from the 1980s in the same region. We suggest managers and regulatory agencies monitor Indiana bats in dynamic landscapes such as those with changing forest composition and biomass.     

Rain increases the energy cost of bat flight

Similar to insects, birds and pterosaurs, bats have evolved powered flight. But in contrast to other flying taxa, only bats are furry. Here, we asked whether flight is impaired when bat pelage and wing membranes get wet. We studied the metabolism of short flights in Carollia sowelli, a bat that is exposed to heavy and frequent rainfall in neotropical rainforests. We expected bats to encounter higher thermoregulatory costs, or to suffer from lowered aerodynamic properties when pelage and wing membranes catch moisture. Therefore, we predicted that wet bats face higher flight costs than dry ones. We quantified the flight metabolism in three treatments: dry bats, wet bats and no rain, wet bats and rain. Dry bats showed metabolic rates predicted by allometry. However, flight metabolism increased twofold when bats were wet, or when they were additionally exposed to rain. We conclude that bats may not avoid rain only because of sensory constraints imposed by raindrops on echolocation, but also because of energetic constraints.     

Base-compositional biases and the bat problem. III. The questions of microchiropteran monophyly.

Using single-copy DNA hybridization, we carried out a whole genome study of 16 bats (from ten families) and five outgroups (two primates and one each dermopteran, scandentian, and marsupial). Three of the bat species represented as many families of Rhinolophoidea, and these always associated with the two representatives of Pteropodidae. All other microchiropterans, however, formed a monophyletic unit displaying interrelationships largely in accord with current opinion. Thus noctilionoids comprised one clade, while vespertilionids, emballonurids, and molossids comprised three others, successively more closely related in that sequence. The unexpected position of rhinolophoids may be due either to the high AT bias they share with pteropodids, or it may be phylogenetically authentic. Reanalysis of the data with varying combinations of the five outgroups does not indicate a rooting problem, and the inclusion of many bat lineages divided at varying levels similarly discounts long branch attraction as an explanation for the pteropodid-rhinolophoid association. If rhinolophoids are indeed specially related to pteropodids, many synapomorphies of Microchiroptera are called into question, not least the unitary evolution of echolocation (although this feature may simply have been lost in pteropodids). Further, a rhinolophoid-pteropodid relationship--if true--has serious implications for the classification of bats. Finally, among the outgroups, an apparent sister-group relation of Dermoptera and Primates suggests that flying lemurs do not represent the ancestors of some or all bats; yet, insofar as gliding of the type implemented in dermopterans is an appropriate model for the evolution of powered mammalian flying, the position of Cynocephalus in our tree indirectly strengthens the argument that true flight could have evolved more than once among bats.     

Bats in a changing landscape: Linking occupancy and traits of a diverse montane bat community to fire regime

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