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.     

Base-compositional biases and the bat problem. I. DNA-hybridization melting curves based on AT- and GC-enriched tracers.

We explored the interordinal relationships of mammals using DNA-DNA hybridization, with particular reference to the much-debated problem of whether the megabats and microbats are more closely related to each other than the megabats are to primates. To try to improve resolution when taxa are distantly related and the melting points of hybrids are low and difficult to distinguish, we increased the GC content of DNA by a fractionation method that used the same melting-point apparatus used in the hybridization studies. When we used GC-rich DNA as the tracer to make hybrids, the melting point of the self-hybrid shifted to a higher temperature as expected, but the behaviour of heterologous hybrids varied with the taxa being compared. When the melting point of the heterologous hybrid also shifted to a higher temperature so that the two compared taxa maintained the same or proportional distance, we called this ''following behaviour'', because the heterologous hybrid made with GC-rich tracer ''followed'' the GC-rich self-hybrid to higher temperatures. We also commonly saw anomalous behaviour, where the melting point of the heterologous hybrid shifted to a lower temperature when compared with an AT-rich hybrid. In these anomalous cases, the distance measured between the taxa increased markedly as a result of GC-enrichment, indicating that an underestimate of distance may have resulted from AT bias in DNA. This inference was supported by the finding that it was rare to observe a decrease in measured distance between taxa using GC-rich DNA, but very common to find an increase as would be expected from the generally higher AT contents of eutherian DNAs. Moreover, the most extreme cases, where distances changed most using GC-rich DNA, were usually those involving comparisons between taxa known to have the most extreme AT-biases among mammals, such as the megabats and rhinolophoid (including megadermatid) microbats. Our results show consistent underestimates of measured differences between eutherian taxa with extreme AT-biases.     

The food habits of a Malagasy Giant: Hipposideros commersoni (E. Geoffroy, 1813)

Hipposideros commersoni is a large microchiropteran bat endemic to Madagascar. We analysed fragments of its prey from faeces and from underneath feeding perches to describe its diet from four sites. Diet was similar across sites and Coleoptera was the main prey item by percentage volume (75%), followed by Hemiptera (13%). Carabidae and Scarabidae were the most frequent coleopterans found in the diet. Direct observations (n = 3) were made of bats flying short distances from perches along forest trails to prey on Cicadidae ( c. 20 mm in length) located on tree trunks. There were differences in the composition of faecal samples collected form netted bats and pellets collected under feeding perches, with the latter consisting of more Blattoptera (Blattellidae 'cockroaches'). Hipposideros commersoni appears to have a unique foraging behaviour and diet among Malagasy microchiropterans and its preference for certain Coleoptera and other large invertebrates may account for reported seasonal variation in body fattening and activity.     

Vanishing GC-rich isochores in mammalian genomes

To understand the origin and evolution of isochores-the peculiar spatial distribution of GC content within mammalian genomes-we analyzed the synonymous substitution pattern in coding sequences from closely related species in different mammalian orders. In primate and cetartiodactyls, GC-rich genes are undergoing a large excess of GC --> AT substitutions over AT --> GC substitutions: GC-rich isochores are slowly disappearing from the genome of these two mammalian orders. In rodents, our analyses suggest both a decrease in GC content of GC-rich isochores and an increase in GC-poor isochores, but more data will be necessary to assess the significance of this pattern. These observations question the conclusions of previous works that assumed that base composition was at equilibrium. Analysis of allele frequency in human polymorphism data, however, confirmed that in the GC-rich parts of the genome, GC alleles have a higher probability of fixation than AT alleles. This fixation bias appears not strong enough to overcome the large excess of GC --> AT mutations. Thus, whatever the evolutionary force (neutral or selective) at the origin of GC-rich isochores, this force is no longer effective in mammals. We propose a model based on the biased gene conversion hypothesis that accounts for the origin of GC-rich isochores in the ancestral amniote genome and for their decline in present-day mammals.     

Compositional Bias May Affect Both DNA-Based and Protein-Based Phylogenetic Reconstructions

It is now well-established that compositional bias in DNA sequences can adversely affect phylogenetic analysis based on those sequences. Phylogenetic analyses based on protein sequences are generally considered to be more reliable than those derived from the corresponding DNA sequences because it is believed that the use of encoded protein sequences circumvents the problems caused by nucleotide compositional biases in the DNA sequences. There exists, however, a correlation between AT/GC bias at the nucleotide level and content of AT- and GC-rich codons and their corresponding amino acids. Consequently, protein sequences can also be affected secondarily by nucleotide compositional bias. Here, we report that DNA bias not only may affect phylogenetic analysis based on DNA sequences, but also drives a protein bias which may affect analyses based on protein sequences. We present a striking example where common phylogenetic tools fail to recover the correct tree from complete animal mitochondrial protein-coding sequences. The data set is very extensive, containing several thousand sites per sequence, and the incorrect phylogenetic trees are statistically very well supported. Additionally, neither the use of the LogDet/paralinear transform nor removal of positions in the protein alignment with AT- or GC-rich codons allowed recovery of the correct tree. Two taxa with a large compositional bias continually group together in these analyses, despite a lack of close biological relatedness. We conclude that even protein-based phylogenetic trees may be misleading, and we advise caution in phylogenetic reconstruction using protein sequences, especially those that are compositionally biased. Received: 19 February 1998 / Accepted: 28 August 1998     

Estimating density of a forest-dwelling bat: a predictive model for Rafinesque’s big-eared bat

Data on species distribution and abundance are the foundation of population ecology. However, due to difficulties in surveying bats, abundance estimates for tree-roosting microchiropterans are non-existent. Therefore, our objective was to develop methods for estimating colony abundance and density, taking as our model Rafinesque’s big-eared bat Corynorhinus rafinesquii, a species of conservation concern found in cypress-gum swamps of the southeastern United States. We searched 123 transects at eight study sites in the Coastal Plain of Georgia, USA to locate and characterize diurnal summer roosts of C. rafinesquii. We modeled the relationship between the number of bat colonies and landscape-scale habitat variables with zero-inflated negative binomial regression and used Akaike’s information criterion to select the most parsimonious models. We generated a predictive density map to identify areas of high colony density and to estimate overall abundance. Colony density was predicted by the duration of wetland flooding, wetland width, and study site. Application of the regression model to a GIS indicated there were 3,734 colonies containing 6,910 adult bats on the eight study sites. Predicted density ranged from 0.07 colonies/ha and 0.07 adult bats/ha in saturated wetlands to 0.47 colonies/ha and 1.18 adult bats/ha in semi-permanently flooded wetlands. This study is the first to estimate density and abundance of forest-dwelling microchiropterans over a large area. Such data can serve as a baseline for future work on population trends in C. rafinesquii. In addition, our approach could be replicated for other bat species with moderately cryptic roosts.     

Characterization and phylogenetic utility of the mammalian protamine p1 gene

We sequenced the protamine P1 gene (ca. 450 bp) from 20 bats (order Chiroptera) and the flying lemur (order Dermoptera). We compared these sequences with published sequences from 19 other mammals representing seven orders (Artiodactyla, Carnivora, Cetacea, Perissodactyla, Primates, Proboscidea, and Rodentia) to assess structure, base compositional bias, and phylogenetic utility. Approximately 80% of second codon positions were guanine, resulting in protamine proteins containing a high frequency of arginine residues. Our data indicate that codon usage for arginine differs among higher mammalian taxa. Parsimony analysis of 40 species representing nine orders produced a well-resolved tree in which most nodes were supported strongly, except at the lowest taxonomic levels (e.g., within Artiodactyla and Vespertilionidae). These data support monophyly of several taxa proposed by morphologic and molecular studies (all nine orders: Laurasiatheria, Cetartiodactytla, Yangochiroptera, Noctilionoidea, Rhinolophoidea, Vespertilionoidea, Phyllostomidae, Natalidae, and Vespertilionidae) and, in agreement with recent molecular studies, reject monophyly of Archonta, Volitantia, and Microchiroptera. Bats were sister to a clade containing Perissodactyla, Carnivora, and Cetartiodactyla, and, although not unequivocally, rhinolophoid bats (traditional microchiropterans) were sister to megachiropterans. Sequences of the protamine P1 gene are useful for resolving relationships at and above the familial level in bats, and generally within and among mammalian orders, but with some drawbacks. The coding and intervening sequences are small, producing few phylogenetically informative characters, and aligning the intron is difficult, even among closely related families. Given these caveats, the protamine P1 gene may be important to future systematic studies because its functional and evolutionary constraints differ from other genes currently used in systematic studies.     

Behaviors of Western Spruce Budworm Moths (Choristoneura occidentalis) as Defences Against Bat Predation

We investigated potential defense behaviors of adult western spruce budworm (Choristoneura occidentalis), a non-auditive lepidopteran, against bat predation. Although western spruce budworm moths started to fly before sunset, earlier than many species of moths, temporal isolation of flying moths from foraging bats was incomplete as moths were most active after sunset once bats were foraging. Flying C. occidentalis were most active close to their host trees, and thus were isolated from some bat activity because vegetation limits foraging by some bats. Moths mostly flew near the tops of trees, an area that may have a high predation pressure from bats. Resting western spruce budworm spent little time fluttering their wings or crawling, behaviors that are used as cues by gleaning bats. The outbreak nature of this species, in which large numbers of moths are active at one time, may allow dilution effects to reduce predation risk.     

Carpal development and morphology in archontan mammals

Carpal morphology and development in bats, colugos, tree shrews, murids, and sciurids were studied in order to homologize carpal elements. Prenatal coalescence of discrete cartilaginous templates with a loss of a center of ossification appears to be the most common method of reducing carpal elements in these mammals. Only bats and colugos showed postnatal ossification between discrete elements as a method of reducing carpal elements. Carpal morphology of tree shrews is more diverse than previously reported. Ptilocercus shows a highly derived carpal morphology that may be related to its relatively greater arboreality. Dendrogale exhibits what is most likely the ancestral tupaiid carpal morphology. Carpal morphologies of Tupaia, Urogale, and Anathana are identical to each other. Carpal morphology differs between megachiropterans and microchiropterans. These differences may be related to different aerodynamic constraints between the suborders. The carpal morphology of microchiropterans is diverse and may reflect different adaptive regimes between microchiropteran families. Carpal morphology of the colugos shows both megachiropteran and microchiropteran characters. The function of these characters in colugos and bats (stabilization of the carpus in dorsiflexion) is proposed to be similar, although the locomotor roles may be quite different between these taxa. J. Morphol. 235:135–155, 1998. © 1998 Wiley-Liss, Inc.     

Base compositional bias and phylogenetic analyses: a test of the "flying DNA" hypothesis

Phylogenetic methods can produce biased estimates of phylogeny when base composition varies along different lineages. Pettigrew (1994, Curr. Biol. 4:277-280) has suggested that base composition bias is responsible for the apparent support for the monophyly of bats (Chiroptera: megabats and microbats) from several different nuclear and mitochondrial genes. Pettigrew's "flying DNA" hypothesis makes several predictions: (1) that metabolic constraints associated with flying result in elevated levels of adenine and thymine throughout the genome of both megabats and microbats, (2) that the resulting base compositional bias in bats is sufficient to mislead phylogenetic methods and account for the support for bat monophyly from several nuclear and mitochondrial genes, and (3) that phylogenetic analysis using pairwise distances corrected for compositional bias should eliminate the support for bat monophyly. We tested these predictions by analyzing DNA sequences from two nuclear and three mitochondrial genes. The predicted base compositional bias does not appear to exist in some of the genes, and in other genes the differences in AT content are very small. Analyses under a wide diversity of criteria and models of evolution, including analyses that take base composition into account (using log-determinant distances), all strongly support bat monophyly. Moreover, simulation analyses indicate that even extreme bias toward AT-base composition in bats would be insufficient to explain the observed levels of support for bat monophyly. These analyses provide no support for the "flying DNA" hypothesis, whereas the monophyly of bats appears to be well supported by the DNA sequence data.     

Habitat fragmentation and the prevalence of parasites (Diptera,Streblidae) on three Phyllostomid bat species

Ectoparasitism in bats seems to be influenced strongly by the type of roost preferred by the hosts, and group size; however, the effect of habitat loss and fragmentation on the prevalence of ectoparasites in bats has scarcely been studied. In northeastern Yucatan, Mexico, we estimated the prevalence of infestation by Streblidae flies in three phyllostomid bat species with different roost preferences (caves, trees, or both) in two types of landscape matrices (tropical semi‐deciduous forest and man‐made pastures) that differed in area of forest cover and the number of forest fragments. Habitat fragmentation and the presence of a contrasting matrix may limit the availability of roosts (trees) and the movement of bats across the landscape. Accordingly, we hypothesized higher prevalence of Streblidae infestation in the pasture matrix and in the group of bats that roost in trees. Bat abundance was higher in the pasture matrix; however, the prevalence of infestation was significantly higher in the continuous forest matrix and in bats that roosted in caves. The prevalence of some species of Streblidae was affected by habitat fragmentation in species that roost in caves, such as Desmodus rotundus, as well as those using foliage and caves, such as Artibeus jamaicensis. Our results provide evidence that some species of Streblidae may respond differently to habitat fragmentation than their hosts, generating changes to bat‐ectoparasite interactions in fragmented areas. Environmental variations involving roosts, not evaluated in this study, may influence our results, since these factors affect ectoparasite abundance and reproduction.     

Thermoregulation in the Angolan free-tailed bat Mops condylurus: A small mammal that uses hot roosts.

The Angolan free-tailed bat (Mops condylurus) uses roosts that often exceed 40 degrees C, an ambient temperature (Ta) that is lethal to many microchiropterans. We measured the physiological responses of this species at Ta's from 15 degrees to 45 degrees C. Torpor was commonly employed during the day at the lower Ta, but the bats generally remained euthermic at night, with a mean body temperature (Tb) of 35.2 degrees C. Metabolic rate reflected the pattern of Tb, increasing with falling Ta at night but decreasing during the day. Metabolic rate and evaporative losses were lower in torpid than in euthermic bats. Body temperature increased at each Ta >35 degrees C and was 43 degrees C at Ta of 45 degrees C. At Ta of 40 degrees C bats increased dry thermal conductance and evaporative heat loss compared to lower Ta. At 45 degrees C dry thermal conductance was lower than at 40 degrees C and evaporative heat loss was 132% of metabolic heat production. At high Ta there was only a slight increase in metabolic rate despite the employment of evaporative cooling mechanisms and an increase in Tb. Collectively our results suggest that M. condylurus is well suited to tolerate high Ta, and this may enable it to exploit thermally challenging roost sites and to colonise habitats and exploit food sources where less stressful roosts are limiting.     

Base Compositional Bias and Phylogenetic Analyses: A Test of the “Flying DNA” Hypothesis

Phylogenetic methods can produce biased estimates of phylogeny when base composition varies along different lineages. Pettigrew (1994,Curr. Biol.4:277–280) has suggested that base composition bias is responsible for the apparent support for the monophyly of bats (Chiroptera: megabats and microbats) from several different nuclear and mitochondrial genes. Pettigrew's “flying DNA” hypothesis makes several predictions: (1) that metabolic constraints associated with flying result in elevated levels of adenine and thymine throughout the genome of both megabats and microbats, (2) that the resulting base compositional bias in bats is sufficient to mislead phylogenetic methods and account for the support for bat monophyly from several nuclear and mitochondrial genes, and (3) that phylogenetic analysis using pairwise distances corrected for compositional bias should eliminate the support for bat monophyly. We tested these predictions by analyzing DNA sequences from two nuclear and three mitochondrial genes. The predicted base compositional bias does not appear to exist in some of the genes, and in other genes the differences in AT content are very small. Analyses under a wide diversity of criteria and models of evolution, including analyses that take base composition into account (using log-determinant distances), all strongly support bat monophyly. Moreover, simulation analyses indicate that even extreme bias toward AT-base composition in bats would be insufficient to explain the observed levels of support for bat monophyly. These analyses provide no support for the “flying DNA” hypothesis, whereas the monophyly of bats appears to be well supported by the DNA sequence data.     

Chiropteran tendon locking mechanism

Rather than the usual mammalian scheme in which tendon and sheath surfaces provide as little friction as possible, the tendons and sheaths of many bats have a locking segment on the manual and pedal flexor tendon complex. This tendon locking mechanism (TLM) exists opposite the proximal phalanges of each toe and pollex of many bats. Its structure, similar to a ratchet mechanism, assists bats in hanging with little muscular effort. The third digit of the pelvic limb and the pollex of species representing 15 chiropteran families were studied to determine the presence or absence, morphology, and function of the TLM. Most of the species studied have a TLM consisting of a patch of tubercles on the ventral surface of the flexor tendon associated with the proximal phalanx of each pollex or toe. The sheath adjacent to this portion of the flexor tendon has a series of transverse folds or ridges, which, when engaged with the tubercles on the tendon, lock the tendon in place. The TLM is similar in megachiropterans and microchiropterans possessing it. The TLM is absent, however, in some of the microchiropterans studied, most notably in the phyllostomids. Since many birds have a TLM similar to that of bats, it is an excellent example of the convergent evolution of a feature brought about by similar functional pressures on birds and bats. © 1993 Wiley-Liss, Inc.     

Character analysis and the integration of molecular and morphological data in an understanding of sand dollar phylogeny.

Reconciling discordant morphological and molecular phylogenies remains a problem in modern systematics. By examining conflicting DNA-hybridization and morphological phylogenies of sand dollars, I show that morphological criteria may be used to help evaluate the reliability of molecular phylogenies where they differ from morphological trees. All available criteria for assessing the reliability of DNA-hybridization phylogenies suggest that the sand dollar DNA-hybridization phylogeny is robust. Standard homology-recognition criteria are used to assess the a priori reliabilities of the morphological attributes associated with the node drawn into question by the DNA data, and it is shown that these attributes are among the least phylogenetically informative of all the morphological characters. Moreover, the questioned node has the smallest number of supporting characters, and most of these characters are associated with the food grooves, which suggests that they may be functionally correlated. Thus, on the basis of the analysis of the morphological data and given the robustness of the DNA tree, the DNA phylogeny is preferred. Further, paleobiogeographic data support the DNA tree rather than the morphological tree, and a plausible heterochronic mechanism has been proposed that may account for the homoplasious morphological evolution that must have occurred if the DNA tree is correct.     

Molecular phylogeny of plethodonine salamanders and hylid frogs: statistical analysis of protein comparisons.

The bootstrapping method of determining confidence in the topology of phylogenetic trees has been applied to electrophoretic protein data for two groups of amphibians: salamanders of two North American genera (Aneides and Plethodon) of the tribe Plethodontini and Holarctic hylid frogs. Some current methods of phylogenetic reconstruction for electrophoretic protein data have been evaluated by comparing the trees obtained from molecular data sets with available morphological data. Molecular data on the phylogenetic relationships of Aneides and Plethodon, data obtained from electrophoretic and immunological studies, indicate that Aneides probably was derived from western Plethodon subsequent to the separation of eastern and western Plethodon. Thus Plethodon very likely is a paraphyletic genus. The extremely low rate of morphological evolution in Plethodon compared with that in Aneides causes difficulty in indicating their evolutionary relationships taxonomically because there are no synapomorphic morphological characters that define either eastern or western Plethodon, whereas there are several for the genus Aneides. Thus molecular data alone probably indicate the evolutionary relationships of the species in these genera. Highton and Larson's (1979) arrangement of species of Plethodon into eight species groups is supported. The topologies of the unweighted pair-group method using arithmetic means (UPGMA) and distance Wagner trees were compared with independent morphological and molecular data on the relationships of the 28 plethodonine species. It was found that UPGMA trees indicate relationships that are more in agreement with other information than are those provided by distance Wagner trees. The use of the bootstrap technique indicates that the topologies of UPGMA trees are better supported statistically than are the topologies of distance Wagner trees. Moreover, different addition criteria produce a variety of distance Wagner trees with different topologies, each with several groupings that are not supported statistically. It is concluded that considerable caution should be used in interpreting the topology of distance Wagner trees. Very similar results were obtained with a second data set on 30 taxa of Holarctic hylid frogs. Trees obtained by the neighbor-joining method are more in agreement with UPGMA phenograms and other data, so this method of phylogenetic reconstruction may be useful to systematists not willing to assume constant rates of evolution.(ABSTRACT TRUNCATED AT 400 WORDS)     

Enriching agroecosystems with fruit-producing tree species favors the abundance and richness of frugivorous and nectarivorous bats in Veracruz, Mexico

We evaluated how the abundance and richness of frugivorous and nectarivorous bat species differs among three types of common agroecosystems (diversified coffee plantations, simple coffee plantations and pastures) in Veracruz, Mexico, that represent a gradient of structural and floristic complexity. Using mixed effects models we demonstrated that both the richness and the total abundance of bats was higher in the diversified coffee plantations. We detected similar patterns on comparing the abundance of the four most abundant bat species. Neither season nor the season-agroecosystem interaction had any effect on the comparisons made. Using multiple regressions we found that the richness of plants that are useful to both people and bats had the most explanatory power for the richness and total abundance of frugivorous and nectarivorous bats, as well as for the abundance of Carollia sowelli, Glossophaga soricina and Sturnira spp. Our results indicate that agroecosystems value for conservation of fruit and nectar-eating bats increases as the fruit-bearing trees increases. For the effective conservation of these guilds of bats in tropical agroecosystems, a strategy of diversification with fruit-bearing species is highly recommended; such a strategy would benefit both agricultural producers and wildlife.     

Evolutionary Studies on an α-amylase Gene Segment in Bats and other Mammals

Comparative studies of salivary glands showed that they maybe related to the adaptive radiation of bats, especially in the family Phylostomidae. In this study we have been searching for a likely relationship between different feeding habits found in bats and possible adaptive changes in a coding segment of the α-amylase enzyme. We have also tested some hypothesis about the phylogenetic relationship of bats and other mammals. A 663 bp segment of the α-amylase gene, corresponding to the exon 4 and part of the intron c, was sequenced in nine bat species. The exon 4 was also sequenced in further ten mammalian species. The phylogenetic trees generated with different methods produced the same results. When the intron c and the exon 4 were independently analyzed, they showed distinct topologies involving the bat species Sturnira lilium, different from the traditional bat phylogeny. Phylogenetic analysis of bats, primates and rodents supports the Euarchontoglires-Laurasiatheria hypothesis about the relationship among these groups. Selection tests showed that the α-amylase exon 4 is under strong purifying selection, probably caused by functional constraints. The conflicting bat phylogenies could not be explained by evolutionary convergence due to adaptive forces, and the different topologies may be likely due to the retention of plesiomorphic characters or the independent acquisition by evolutionary parallelism.     

The evolution of flight and echolocation in bats: another leap in the dark

The earliest known complete bats, from the Eocene (49–53 Mya), were already capable of flapping flight and echolocation. In the absence of direct fossil evidence there have been many speculative scenarios advanced to explain the evolution of these behaviours and their distributions in extant bats. Theories assuming chiropteran monophyly have generally presumed the ancestral pre‐bat was nocturnal, arboreal and insectivorous. Following this assumption hypotheses can be divided into the echolocation first, flight first and tandem development hypotheses, all of which assume that flight evolved only once in the lineage. In contrast, the chiropteran diphyly hypothesis suggests that flight evolved twice. Evidence supporting and refuting the different hypotheses are reviewed. It is concluded that there are significant problems attached to all the current models. A novel hypothesis is advanced, which starts from the assumption that bats are monophyletic and the ancestral pre‐bat was arboreal, but diurnal and frugivorous. After the evolution of flight it is suggested that these animals were driven into the nocturnal niche by the evolution of raptorial birds, and different groups evolved either specialised nocturnal vision (megachiropterans) or echolocation (microchiropterans). A block on sensory modality transfer has retained this distribution of perceptual capabilities ever since, despite some Megachiroptera evolving rudimentary echolocation, and the dietary convergence of some Microchiroptera with the Megachiroptera. The new hypothesis overcomes many of the problems identified in previous treatments.