Untangling latitudinal richness gradients at higher taxonomic levels: familial perspectives on the diversity of New World bat communities

Aims (i) To describe at the level of local communities latitudinal gradients in the species richness of different families of New World bats and to explore the generality of such gradients. (ii) To characterize the relative effects of changes in the richness of each family to the richness of entire communities. (iii) To determine differences in the rate and direction of latitudinal gradients in species richness within families. (iv) To evaluate how differences among families regarding latitudinal gradients in species richness influence the latitudinal gradient in species richness of entire communities. Location Continental New World ranging from the northern continental United States (Iowa, 42° N) to eastern Paraguay (Canindeyú, 24° S). Methods Data on the species composition of communities came from 32 intensively sampled sites. Analyses focused on species richness of five of nine New World bat families. Multivariate analysis of variance and discriminant function analysis determined and described differences among temperate, subtropical, and tropical climatic zones regarding the species richness of bat families. Simple linear regression described latitudinal gradients in species richness of families. Path analysis was used to describe: (i) the direct effect of latitude on species richness of communities, (ii) the indirect effects of latitude on the species richness of communities through its effect on the species richness of each family, (iii) the relative effects of latitude on the species richness of bat families, and (iv) the relative contribution of each family to variation in the species richness of communities. Results Highly significant differences among climatic zones existed primarily because of a difference between the temperate zone and the tropical and subtropical zones combined. This difference was associated with the high number of vespertilionids in the temperate zone and the high number of phyllostomids in the tropical and subtropical zones. Latitudinal gradients in species richness were contingent on phylogeny. Although only three of the five families exhibited significant gradients, all families except for the Vespertilionidae exhibited indistinguishable increases in species richness with decreases in latitude. The Emballonuridae, Phyllostomidae and Vespertilionidae exhibited significant latitudinal gradients whereby the former two families exhibited the classical increase in species richness with decreasing latitude and the latter family exhibited the opposite pattern. Variation in species richness of all families contributed significantly to variation in the species richness of entire communities. Nonetheless, the Phyllostomidae made a significantly stronger contribution to changes in species richness of communities than did all other families. Much of the latitudinal gradient in species richness of communities could be accounted for by the effects of latitude on the species richness of constituent families. Main conclusions Ecological and evolutionary differences among higher taxonomic units, particularly those differences involving life‐history traits, predispose taxa to exhibit different patterns of diversity along environmental gradients. This may be particularly true along extensive gradients such as latitude. Nonetheless, species rich taxa, by virtue of their greater absolute rates of change, can dominate and therefore define the pattern of diversity at a higher taxonomic level and eclipse differences among less represented taxa in their response to environmental gradients. This is true not only with respect to how bats drive the latitudinal gradient in species richness for all mammals, but also for how the Phyllostomidae drives the latitudinal gradient for all bats in the New World. Better understanding of the mechanistic basis of latitudinal gradients of diversity may come from comparing and contrasting patterns across lower taxonomic levels of a higher taxon and by identifying key ecological and evolutionary traits that are associated with such differences.     

A faunal survey of streblid flies (Diptera: Streblidae) associated with bats in Paraguay

An extensive survey of the ectoparasites infesting bats in Paraguay provides information regarding the taxonomy and host distribution of streblid bat flies at a geographic interface between subtropical and temperate habitats. Five families of bats representing 45 species, including Molossidae (5 genera and 15 species), Natalidae (1 genus and 1 species), Phyllostomidae (11 genera and 15 species), Noctilionidae (1 genus and 2 species), and Vespertilionidae (4 genera and 12 species) were collected from 24 localities across Paraguay and sampled for ectoparasites. In total, 2,467 bat flies were collected, representing 11 genera and 31 nominal species of Streblidae, of which 6 genera and 24 species are new records for Paraguay. No streblids were collected from vespertilionid bats; 23 species infested phyllostomids, 6 species noctilionids, 1 species a natalid, and 1 species molossids. Streblid bat flies were highly specific to certain host groups and individual host species, and their geographic distributions closely followed those of their host bats. Of 31 streblid species surveyed, 27 were monoxenous (i.e., associated with a single host species), and 4 were stenoxenous (i.e., associated with a group of phylogenetically related hosts). The number of streblid species is greatly reduced in the Chaco region west of the Paraguay River, largely because of the lack of phyllostomid host bats.     

A family matter: conclusive resolution of the taxonomic position of the long-fingered bats, miniopterus

The long-fingered bats (Miniopterus sp.) are among the most widely distributed mammals in the world. However, despite recent focus on the systematics of these bats, their taxonomic position has not been resolved. Traditionally, they are considered to be sole members of Miniopterinae, 1 of 5 subfamilies within the largest family of bats, the Vespertilionidae. However, this classification has increasingly been called into question. Miniopterines differ extensively from other vespertilionids in numerous aspects of morphology, embryology, immunology, and, most recently, genetics. Recent molecular studies have proposed that the miniopterines are sufficiently distinct from vespertilionids that Miniopterinae should be elevated to full familial status. However, controversy remains regarding the relationship of the putative family, Miniopteridae to existing Vespertilionidae and to the closely related free-tailed bats, the Molossidae. We report here the first conclusive analysis of the taxonomic position of Miniopterus relative to all other bat families. We generated one of the largest chiropteran data sets to date, incorporating approximately 11 kb of sequence data from 16 nuclear genes, from representatives of all bat families and 2 Miniopterus species. Our data confirm the distinctiveness of Miniopterus, and we support previous recommendations to elevate these bats to full familial status. We estimate that they diverged from all other bat species approximately 49-38 MYA, which is comparable to most other bat families. Furthermore, we find very strong support from all phylogenetic methods for a sister group relationship between Miniopteridae and Vespertilionidae. The Molossidae diverged from these lineages approximately 54-43 MYA and form a sister group to the Miniopteridae-Vespertilionidae clade.     

澳门翼手目物种多样性补充调查及其保护现状

<正>翼手目为哺乳动物中的第二大目,中国的翼手目多样性也在不断更新,Smith和解焱(2009)统计中国翼手目118种,蒋志刚等(2015)则认为中国翼手目有7科34属134种。广东省翼手目有6科24属55种(邹发生和叶冠锋,2016),此后也仍陆续有新纪录,如长指鼠耳蝠(Myotis longipes)(张琴等,2017)、卡氏伏翼(Hypsugo cadornae)(Xie et al., 2021);我们前期对澳门翼手目的调查,发现5科8属10种(黄继展等,2013)。     

Eimeria from bats of Bolivia: two new species from vespertilionid bats.

Between 1985 and 1987, fecal samples were collected from 71 bats representing 14 species (Desmodontidae, Molossidae, Noctilionidae, Phyllostomidae, Vespertilionidae) from 8 localities in 3 states (Beni, Pando, Santa Cruz) in Bolivia, South America. Of these, 2 black myotid bats (Vespertilionidae), Myotis nigricans, and 1 tent-making bat (Phyllostomidae), Uroderma magnirostrum, had oocysts in their feces that represent undescribed species of Eimeria. The new species from M. nigricans (2/4, 50%) has sporulated oocysts that are subspheroidal, 18.9 x 16.9 (17-23 x 14-20) microm, without a micropyle; oocyst residuum of 6-8 spheroidal globules and 1 highly refractile polar granule are present. The oocyst wall has 2 layers (approximately 1.3 microm thick), with a rough outer layer. Ovoidal sporocysts are 10.1 x 7.4 (7-14 x 5-10) microm, with a Stieda body, substieda body, and a sporocyst residuum. The new eimerian species from U. magnirostrum (1/2, 50%) has sporulated oocysts that are subspheroidal to ellipsoidal, 23.8 x 20.8 (20-26 x 19-24) microm, without micropyle or oocyst residuum, but 1-3 polar granules are present. The oocyst wall has 2 layers (approximately 1.5 microm thick), with a rough, mammilated outer layer. Ovoidal sporocysts are 11.6 x 8.6 (10-12 x 7-10) microm, with a Stieda body, substieda body and a sporocyst residuum.     

Bat flies (Diptera: Streblidae, Nycteribiidae) parasitic on bats (Mammalia: Chiroptera) at Parque Estadual da Cantareira, São Paulo, Brazil: parasitism rates and host-parasite associations

A total of 443 bat flies belonging to the families Nycteribiidae and Strelidae, were collected on 22 species of bats (Molossidae, Phyllostomidae, and Vespertilionidae) from Parque Estadual da Cantareira (S?o Paulo, Brazil), between January, 2000 and January, 2001. Eighteen new occurrences of bat flies were recorded on Anoura geoffroyi (Anastrebla caudiferae), Glossophaga soricina (A. caudiferae), Sturnira lilium (Trichobius phyllostomae, T. furmani, and Paraeuctenodes similis), Artibeus lituratus (A. caudiferae), A. fimbriatus (Megistopoda proxima), A. obscurus (Metelasmus pseudopterus), Myotis nigricans (M. proxima, M. aranea, Paratrichobius longicrus), M. ruber (Anatrichobius passosi, Joblingia sp.), M. levis (A. passosi), M. albescens (A. passosi, Basilia andersoni), and Histiotus velatus (M. aranea). Seven new occurrences were recorded for the state of S?o Paulo, increasing the range for T. tiptoni, T. furmani, M. proxima, Aspidoptera falcata, A. caudiferae, A. modestini and B. andersoni. The relationships between parasitism and host sex, reproductive stage, age hyperparasitism by fungi are discussed.     

Bats, clocks, and rocks: diversification patterns in Chiroptera

Identifying nonrandom clade diversification is a critical first step toward understanding the evolutionary processes underlying any radiation and how best to preserve future phylogenetic diversity. However, differences in diversification rates have not been quantitatively assessed for the majority of groups because of the lack of necessary analytical tools (e.g., complete species-level phylogenies, estimates of divergence times, and robust statistics which incorporate phylogenetic uncertainty and test appropriate null models of clade growth). Here, for the first time, we investigate diversification rate heterogeneity in one of the largest groups studied thus far, the bats (Mammalia: Chiroptera). We use a recent, robust statistical approach (whole-tree likelihood-based relative rate tests) on complete dated species-level supertree phylogenies. As has been demonstrated previously for most other groups, among-lineage diversification rate within bats has not been constant. However, we show that bat diversification is more heterogeneous than in other mammalian clades thus far studied. The whole-tree likelihood-based relative rates tests suggest that clades within the families Phyllostomidae and Molossidae underwent a number of significant changes in relative diversification rate. There is also some evidence for rate shifts within Pteropodidae, Emballonuridae, Rhinolophidae, Hipposideridae, and Vespertilionidae, but the significance of these shifts depends on polytomy resolution within each family. Diversification rate in bats has also not been constant, with the largest diversification rate shifts occurring 30-50 million years ago, a time overlapping with the greatest number of shifts in flowering plant diversification rates.     

Two new species of yellow-shouldered bats,genus Sturnira Gray, 1842 (Chiroptera,Phyllostomidae) from Costa Rica,Panama and western Ecuador

Two new species of yellow-shouldered bats Sturnira Gray, 1842 (Chiroptera, Phyllostomidae) from Central America and western South America are described using molecular and morphological data. The two new species, which occur in Costa Rica and Panama and in western Ecuador, were previously confused with S. ludovici, and S. lilium and S. luisi, respectively. Sturnira now includes 22 described species, making it the most speciose genus in the Neotropical family Phyllostomidae.     

Ectoparasitic insects (Diptera: Streblidae and Siphonaptera: Ischnopsyllidae) of bats from Iquitos and surrounding areas (Loreto, Peru)

Based on specimens collected from bats of different families, we add new species and extend the known ecological distribution and host associations of insect ectoparasites of bats in Peru. New information is provided for the distribution of 26 species of parasites (25 Diptera and 1 Siphonaptera: Ischnopsyllidae). Four species (Neotrichobius ectophyllae, Strebla galindoi, Strebla paramirabilis and Myodopsylla wolffsohni wolffsohni) are new for Peru and 16 represent new records for the department of Loreto. Also, we found 17 new host-ectoparasite relationships. Of note, we found remarkable new association between Neotrichobius delicatus and bat species from the families Molossidae and Noctilionidae and a novel association between Paradyschiria parvula and a species of Vespertilionidae. Host-ectoparasite specificity was recorded with 14 species as monoxenous, three oligoxenous, seven pleioxenous and two polyxenous.     

广东丹霞山国家级自然保护区 蝙蝠物种多样性调查

于2017年至2018年对广东丹霞山国家级自然保护区的翼手目动物进行调查,主要采用日栖息地与夜栖息地、捕食区网捕等调查方法,共调查到翼手目5科13属23种,其中蝙蝠科(Vespertilionidae)9属15种,菊头蝠科(Rhinolophidae)1属4种,蹄蝠科(Hipposideridae)1属2种,假吸血蝠科(Megadermatidae)1属1种,犬吻蝠科(Molossidae)1属1种。从区系组成来看,以东洋界为主(19种),其次为广布种(3种),古北界仅1种。从栖息类型上看,分为洞栖型、树栖型及建筑物栖息型,以洞栖型为主(15种),建筑物栖息型其次(12种),树栖型最少(5种),但其中9个物种的栖息地类型同时包含了上述3种栖息类型中的2种。本研究在广东丹霞山发现中国蝙蝠新分布记录1种,为卡氏伏翼(Hypsugo cadornae);而中印鼠耳蝠(Myotis indochinensis)为中国分布的再次确认。     

The status of the Japanese and East Asian bats of the genus Myotis (Vespertilionidae) based on mitochondrial sequences

The genus Myotis includes the largest number of species in the family Vespertilionidae (Chiroptera), and its members are distributed throughout most of the world. To re-evaluate the phylogenetic position of East Asian Myotis species with respect to Myotis species worldwide, we analyzed mitochondrial gene sequences of NADH dehydrogenase subunit 1 and cytochrome b from 24 East Asian individuals as well as 42 vespertilionid bats determined previously. The results suggest that: (1) some individuals having the same species name in Europe and Japan do not form a monophyletic clade, indicating that some bat species exhibit morphological convergence, (2) Japanese Myotis mystacinus forms a sister relationship with Myotis brandtii (Palaearctic), and both species are included in the American clade implying that an ancestor of these species originated in North America, and (3) the Black whiskered bat, Myotis pruinosus, is endemic to Japan and forms sister relationships with Myotis yanbarensis and Myotis montivagus collected from Okinawa (Japan) and Selangor (Malaysia), respectively, implying that M. pruinosus originated from the south. The systematics of Japanese and East Asian Myotis bats were revisited by considering their phylogenetic relationships. Our study provides the first extensive phylogenetic hypothesis of the genus Myotis that includes East Asian and Japanese species.     

Bat assemblage structure in two dry forests of Colombia: Composition, species richness, and relative abundance

We studied the composition, species richness, and relative abundance of bat assemblages in the Colombian dry forests of Chicamocha and Patía. In Chicamocha, 11 bats of the family Phyllostomidae were captured with mist-nets, corresponding to 85–100% of the potential phyllostomids species in the area. Two bats of the family Vespertilionidae were also captured in Chicamocha. In Patía, 12 species were captured with mist-nets, all Phyllostomidae, representing 72–100% of the estimated total number of species in the zone. Minor differences in number of species and composition were detected among sampling periods in Chicamocha. The most common species in this dry forest were Glossophaga longirostris and Sturnira lilium. In Patía, notable differences in the number of species and composition were observed among sampling periods, and the most common species were Artibeus jamaicensis, Carollia perspicillata and Phyllostomus discolor. Arid-zone dwelling bats were absent in Patía and we suggest that this absence may be associated with the isolation of Patía from other northern dry zones of Colombia since Quaternary times. There was also low abundance of bats in Patía, which appears to be related to human disturbance. The most abundant phyllostomid bat species in the two dry forests studied are those that include fruit and/or nectar-pollen from columnar cacti as an important proportion of their diets.     

Genetic Relationships Between Brazilian Species of Molossidae and Phyllostomidae (Chiroptera, Mammalia)

A comparative analysis of G-banded karyotypes was performed for seven species of Chiroptera, representing two families (Phyllostomidae and Molossidae). Despite the differences in diploid and fundamental numbers, extensive homologies between six karyotypes were identified: A . planirostris, P. lineatus, S. lilium, G. soricina, P. hastatus (Phyllostomidae) and M. rufus (Molossidae). Robertsonian rearrangements and pericentric inversions account for the differences between the karyotypes of phyllostomid and molossid species. The homologies and rearrangements observed reinforce the monophiletic origin of phyllostomids and the inclusion of species in different subfamilies. In situ hybridization with genomic DNA revealed considerable conservation of the karyotypes, including C. perspicillata, that did not show G-band homologies with the other species analyzed. For the first time, chromosomal evidence is presented of a common origin for Phyllostomidae and Molossidae.     

基于核基因RAG2部分序列探索菊头蝠总科、蝙蝠科和鞘尾蝠科物种的系统发育关系

采用PCR技术获得了翼手目(菊头蝠总科、蝙蝠科以及鞘尾蝠科)16种19个个体的核基因重组激活基因2(Recombination activating gene,RAG2)部分序列,长度为730～760 bp.结合从Genbank中提取的翼手目3科14个体的RAG2序列,通过构建贝叶斯(Bayesian inference,BI)和最大似然树对翼手目种属间的系统进化关系进行研究.研究表明:在菊头蝠总科中,菊头蝠科和蹄蝠科是两个独立的科,且在蹄蝠科中,大蹄蝠、中蹄蝠、普氏蹄蝠3种之间的亲缘关系非常近;蝙蝠科中的南蝠属与棕蝠属是姊妹群;长翼蝠亚科应提升为长翼蝠科.此外,鞘尾蝠科与犬吻蝠科形成姊妹群关系.     

Neotropical bats in the canopy: diversity,community structure,and implications for conservation

We compare results of parallel ground and canopy netting of bats (Microchiroptera) in three adjacent forest sites near Belém, Brazil, to document possible differences in vertical distribution of species. We caught 1871 individuals representing 49 species of three families (Emballonuridae, Phyllostomidae, Vespertilionidae). Capture effort, totaling 1955.5 mistnet hours in several cycles over a two-year period, was similar for ground and canopy nets. The canopy rigs yielded more species (n = 41) than the ground nets (n = 35), but both samples were characterized by rank abundance curves with similar shape and with a dominance of frugivores (Phyllostomidae). Nearly half (n = 24) of the species were captured in numbers too small (n < 6) to allow firm classification, but differences in capture frequencies of some of the better-sampled species in high and low nets reveal vertical stratification. Species-specific differences in diet, foraging strategies, roost sites, and sampling bias contribute to this pattern. As a result of the differential use of space among bats, alterations of forest structure are likely to result in changes in structure and function of local bat communities, but our limited knowledge of natural history and ecology of many species limits definition of changes. We see a critical need for further research into the extent to which habitat complexity influences species richness and abundance of bats. This information is especially important in view of the need to develop and apply conservation-oriented programs to maintain biodiversity. A review of recent improvements in techniques for inventorying bats shows that a combination of methods, including mistnetting and acoustic monitoring, is mandatory for such studies.     

Mitochondrial genome of <Emphasis Type="Italic">Pteronotus personatus</Emphasis> (Chiroptera: Mormoopidae): comparison with selected bats and phylogenetic considerations

We described the complete mitochondrial genome (mitogenome) of the Wagner’s mustached bat, Pteronotus personatus, a species belonging to the family Mormoopidae, and compared it with other published mitogenomes of bats (Chiroptera). The mitogenome of P. personatus was 16,570 bp long and contained a typically conserved structure including 13 protein-coding genes, 22 transfer RNA genes, two ribosomal RNA genes, and one control region (D-loop). Most of the genes were encoded on the H-strand, except for eight tRNA and the ND6 genes. The order of protein-coding and rRNA genes was highly conserved in all mitogenomes. All protein-coding genes started with an ATG codon, except for ND2, ND3, and ND5, which initiated with ATA, and terminated with the typical stop codon TAA/TAG or the codon AGA. Phylogenetic trees constructed using Maximum Parsimony, Maximum Likelihood, and Bayesian inference methods showed an identical topology and indicated the monophyly of different families of bats (Mormoopidae, Phyllostomidae, Vespertilionidae, Rhinolophidae, and Pteropopidae) and the existence of two major clades corresponding to the suborders Yangochiroptera and Yinpterochiroptera. The mitogenome sequence provided here will be useful for further phylogenetic analyses and population genetic studies in mormoopid bats.     

Comparative Analysis of the Amino Acid Spectrum of Blood Plasma in Chiroptera (<Emphasis Type="Italic">Vespertilio murinus</Emphasis> L., 1758 and <Emphasis Type="Italic">Myotis dasycneme</Emphasis> B., 1825) in the Fauna of the Ural Mountains

The article presents the results of a new comparative analysis of free amino acids in the blood plasma of representatives of insectivorous Chiroptera (Mammalia: Vespertilionidae) in the fauna of the Ural Mountains: the pond bat (Myotis dasycneme Boie, 1825) and the parti-colored bat (Vespertilio murinus Linnaeus, 1758). This is the first study to show the species variability of free amino acids in resident and migratory species of bats from different ecosystems of the Ural Region.     

The diversity field of New World leaf‐nosed bats (Phyllostomidae)

Aim To analyse how the patterns of species richness for the whole family Phyllostomidae determine the structure of diversity fields (sets of species‐richness values) within the ranges of individual bat species. Location The range of the family Phyllostomidae in North and South America. Methods We generated a database of the occurrence of 143 phyllostomid bat species in 6794 quadrats, analysing the species‐richness frequency distribution for all sites, and for subsets of sites defined by the geographic ranges of species. Range–diversity plots, depicting simultaneously the size and the mean species richness of ranges, were built to explore the patterns of co‐occurrence in widespread and restricted species. We compared the empirical patterns against two null models: (1) with scattered (non‐cohesive) ranges, and (2) with cohesive ranges modelled with the spreading‐dye algorithm. Diversity fields were analysed with richness maps for individual species and with comparisons of species‐richness frequency distributions. Results Overall richness frequency distribution showed a multimodal pattern, whereas simulated distributions showed lower values of variance, and were unimodal (for model 1) and bimodal (for model 2). Range–diversity plots for the empirical data and for the cohesive‐ranges simulation showed a strong tendency of species to co‐occur in high‐diversity sites. The scattered‐ranges simulation showed no such tendency. Diversity fields varied according to idiosyncratic features of species generating particular geographic patterns and richness frequency distributions. Main conclusions Phyllostomid bats show a higher level of co‐occurrence than expected from null models. That tendency in turn implies a higher variance in species richness among sites, generating a wider species‐richness frequency distribution. The diversity field of individual species results from the size, shape and location of ranges, but also depends on the general pattern of richness for the whole family.     

Mitochondrial DNA phylogeography reveals marked population structure in the common vampire bat, Desmodus rotundus (Phyllostomidae)

Desmodus rotundus (Chiroptera: Phyllostomidae; Desmodontinae) is the most common vampire bat and has a broad distribution, ranging from southern Mexico to central Chile in the west, and Paraguay and northern Argentina in the east of South America ( Koopman 1988 ). Because of its feeding habit, this bat is considered the main source of rabies transmission to cattle. Although this species has a large spectrum of morphological variability throughout its range, thus far no study has examined the distribution of genetic lineages over its geographic range. Four geographically circumscribed clades of D. rotundus were described in the Brazilian territory on the basis of mitochondrial sequence analyses: southern Atlantic forest (SAF), northern Atlantic forest (NAF), Pantanal (PAN) and Amazon plus Cerrado (AMC) clade. The differentiation among these clades is strongly supported statistically, although the phylogenetic relationship between them remains uncertain. The extremely high levels of sequence divergence that were found between clades (ranging from 6% to 11%) are the highest ever described for a Neotropical bat species and cannot be explained by female philopatry alone. This indicates that D. rotundus comprises two or more distinct, possibly cryptic species. The biogeographic pattern described for this bat is similar to those described for other bats and terrestrial mammals, suggesting geographical congruence between historical vicariant processes, including likely vicariant events between north and south Atlantic Forest and between the Atlantic Forest and the Amazon.