Subfamily Glossophaginae Research Articles

Comparative mitochondrial genomics of American nectar-feeding long-nosed bats Leptonycteris spp. with insights into the phylogeny of the family Phyllostomidae

Among leaf-nosed bats (family Phyllostomidae), the genus Leptonycteris (subfamily Glossophaginae), contains three migratory and obligate nectar-dwelling species of great ecological and economic importance; the greater long-nosed bat L. nivalis, the lesser long-nosed bat L. yerbabuenae, and the southern long-nosed bat L. curasoae. According to the IUCN, the three species are categorized as ‘vulnerable’, ‘endangered’, and ‘near threatened’, respectively. In this study, we assembled and characterized in detail the mitochondrial genome of Leptonycteris spp. and examined the phylogenetic position of this genus in the family Phyllostomidae based on protein coding genes (PCGs). The mitogenomes of L. nivalis, L. curasoae, and L. yerbabuenae are 16,708, 16,758, and 16,729 bp in length and each encode 13 PCGs, 22 transfer RNA genes, 2 rRNA genes, and a putative control region (CR). Mitochondrial gene order is identical to that reported before for the family Phyllostomidae. All tRNAs exhibit a ‘cloverleaf’ secondary structure, except tRNA-Serine-1 that is missing the DHU arm in the three species. All PCGs are exposed to purifying selection with atp8 experiencing the most relaxed purifying selection as the ω ratio was higher for this gene compared to the other PCGs in each species. The CR of each species contains three functional domains: extended termination associated sequence (ETAS), Central, and conserved sequence block domain (CSB). A phylomitogenomic analysis revealed that Leptonycteris is monophyletic and most closely related to the genus Glossophaga. The analysis also supported the monophyly of the family Glossophaginae in the speciose family Phyllostomidae. The mitochondria characterization of these species provides relevant information to develop molecular markers for conservation purposes.

Synthesis of knowledge of the plant diet of nectar-feeding bats of México

The interaction between bats and plants is key to the stability of ecosystems and economically important industries, such as tequila and mezcal in México. For these reasons, it is important to determine the current state of knowledge about the plant diet of nectar-feeding bats. In this study, we conducted a systematic review of the literature on plants that have been reported as sources of food for nectar-feeding bats (subfamily Glossophaginae) in México. Based on this information, we identified bat species with the best-documented knowledge of their diet, the most consumed plant genera, and the territories with information gaps in the country. The literature search on the diet of nectar-feeding bats was carried out in the Web of Science database, Google Scholar, and digital collections of universities. We constructed rarefaction curves for bat diet richness, a heat map of the plant genera consumed by each, and a map of food localities in the different biogeographical provinces of México. This information served to explore whether knowledge of the diet of bats was related to variables such as the presence of chiropterophilous plants or the richness of nectar-feeding bats. In México, nectar-feeding bats feed mainly on plants of the genera Agave spp., Pseudobombax spp., and Ceiba spp., which, according to the literature, provide food to more than 75 % of nectar-feeding bats in Mexican territory. Leptonycteris yerbabuenae is the species with the most information on its diet in México, while Lichonycteris obscura is the species with the least information. None of the bat species reached a value greater than 65 % of the expected richness. Localities where the plant diet of nectar-feeding bats has been studied correspond to provinces in the Neotropical region of México. Knowledge about the diet of nectar-feeding bats in México is far from complete. The distribution of diet localities is biased over a large part of its geographic range and is positively correlated with areas with higher nectar-feeding bat species richness.

Genome-wide ultraconserved elements resolve phylogenetic relationships and biogeographic history among Neotropical leaf-nosed bats in the genus Anoura (Phyllostomidae)

AnouraGray, 1838 are Neotropical nectarivorous bats and the most speciose genus within the phyllostomid subfamily Glossophaginae. However, Anoura species limits remain debated, and phylogenetic relationships remain poorly known, because previous studies used limited Anoura taxon sampling or focused primarily on higher-level relationships. Here, we conduct the first phylogenomic study of Anoura by analyzing 2039 genome-wide ultraconserved elements (UCEs) sequenced for 42 individuals from 8 Anoura species/lineages plus two outgroups. Overall, our results based on UCEs resolved relationships in the genus and supported (1) the monophyly of small-bodied Anoura species (previously genus Lonchoglossa); (2) monotypic status of A. caudifer; and (3) nested positions of “A. carishina”, A. caudifer aequatoris, and A. geoffroyi peruana specimens within A. latidens, A. caudifer and A. geoffroyi, respectively (suggesting that these taxa are not distinct species). Additionally, (4) phylogenetic networks allowing reticulate edges did not explain gene tree discordance better than the species tree (without introgression), indicating that a coalescent model accounting for discordance solely through incomplete lineage sorting fit our data well. Sensitivity analyses indicated that our species tree results were not adversely affected by varying taxon sampling across loci. Tree calibration and Bayesian coalescent analyses dated the onset of diversification within Anoura to around ∼ 6–9 million years ago in the Miocene, with extant species diverging mainly within the past ∼ 4 million years. We inferred a historical biogeographical scenario for Anoura of parapatric speciation fragmenting the range of a wide-ranging ancestral lineage centered in the Central to Northern Andes, along with Pliocene–Pleistocene dispersal or founder event speciation in Amazonia and the Brazilian Atlantic forest during the last ∼ 2.5 million years.

Differential cellular proliferation underlies heterochronic generation of cranial diversity in phyllostomid bats

BackgroundSkull diversity in the neotropical leaf-nosed bats (Phyllostomidae) evolved through a heterochronic process called peramorphosis, with underlying causes varying by subfamily. The nectar-eating (subfamily Glossophaginae) and blood-eating (subfamily Desmondontinae) groups originate from insect-eating ancestors and generate their uniquely shaped faces and skulls by extending the ancestral ontogenetic program, appending new developmental stages and demonstrating peramorphosis by hypermorphosis. However, the fruit-eating phyllostomids (subfamilies Carollinae and Stenodermatinae) adjust their craniofacial development by speeding up certain developmental processes, displaying peramorphosis by acceleration. We hypothesized that these two forms of peramorphosis detected by our morphometric studies could be explained by differential growth and investigated cell proliferation during craniofacial morphogenesis.ResultsWe obtained cranial tissues from four wild-caught bat species representing a range of facial diversity and labeled mitotic cells using immunohistochemistry. During craniofacial development, all bats display a conserved spatiotemporal distribution of proliferative cells with distinguishable zones of elevated mitosis. These areas were identified as modules by the spatial distribution analysis. Ancestral state reconstruction of proliferation rates and patterns in the facial module between species provided support, and a degree of explanation, for the developmental mechanisms underlying the two models of peramorphosis. In the long-faced species, Glossophaga soricina, whose facial shape evolved by hypermorphosis, cell proliferation rate is maintained at lower levels and for a longer period of time compared to the outgroup species Miniopterus natalensis. In both species of studied short-faced fruit bats, Carollia perspicillata and Artibeus jamaicensis, which evolved under the acceleration model, cell proliferation rate is increased compared to the outgroup.ConclusionsThis is the first study which links differential cellular proliferation and developmental modularity with heterochronic developmental changes, leading to the evolution of adaptive cranial diversity in an important group of mammals.

Interactions between bats and floral resources in a premontane forest, Valle del Cauca, Colombia

The study of interaction networks between species is a subject that has drawn increasing attention in recent decades, especially in investigations involving relationships between plants and pollinators or seed dispersers. In the Neotropics, bats of the subfamily Glossophaginae show morphological modifications for a specialized diet consisting of nectar and pollen, but opportunistic species belonging to other subfamilies that consume floral resources (nectar and pollen) have also been identified. This study describes for the first time the interactions between nectarivorous bats in the Andean region of Colombia from the identification of pollen associated with the bat species inhabiting the protected area “Reserva Forestal Bosque de Yotoco (RFBY)”. Bats were captured with mist nets; a pollen sample was collected from each specimen by contact with glycerin gelatin, and pollen samples were mounted on slides. In addition, plant material of the blooming species that displayed the syndrome of chiropterophyly was collected to build a reference pollen collection. For the analysis, we used Levin’s standardized niche breadth (BA), the relative frequency of resources (Fi), and the Resource Importance Value Index (RIVI). The interaction network was built and the NODF nesting algorithm was calculated to establish the degree of coupling within the network. In 44 loads of contact pollen for six species, three belonging to the subfamily Glossophaginae (Anoura caudifer, Glossophaga soricina and Choeroniscus godmani), we identified six palynomorphs from 7,971 pollen grains. The highest RIVI values correspond to three species of the genus Inga. The Levin’s standardized niche breadth (BA) highlights Anoura caudifer and Glossophaga soricina as generalist species that take advantage of the availability of floral resources for consumption (Tables 2, 3). The interaction network reveals a high nesting degree according to the value of the NODF algorithm of 86.67 (Figure 3). With four bat species, the RFBY has a high diversity of nectarivorous species relative to other localities across the Andean region of Colombia, despite the study area being a fragmented landscape; hence, this area is of critical importance for the local bat species, as it fulfills their food requirements. The bat species using the largest amount of resources in the community is Anoura caudifer; this finding differs from the results of other studies, where the genus Glossophaga has been reported as the dominant taxon in terms of resource use. Plants of the genus Inga and O. pyramidale are key resources for the nectarivorous bat species recorded, and this is probably also the case for other animals in the area; therefore, the inclusion of these species in the active restoration process will likely promote the establishment of a larger number of bats.

The thermal niche of Neotropical nectar-feeding bats: Its evolution and application to predict responses to global warming.

The thermal niche of a species is one of the main determinants of its ecology and biogeography. In this study, we determined the thermal niche of 23 species of Neotropical nectar‐feeding bats of the subfamily Glossophaginae (Chiroptera, Phyllostomidae). We calculated their thermal niches using temperature data obtained from collection records, by generating a distribution curve of the maximum and minimum temperatures per locality, and using the inflection points of the temperature distributions to estimate the species optimal (STZ) and suboptimal (SRZ) zones of the thermal niche. Additionally, by mapping the values of the STZ and SRZ on a phylogeny of the group, we generated a hypothesis of the evolution of the thermal niches of this clade of nectar‐feeding bats. Finally, we used the characteristics of their thermal niches to predict the responses of these organisms to climate change. We found a large variation in the width and limits of the thermal niches of nectar‐feeding bats. Additionally, while the upper limits of the thermal niches varied little among species, their lower limits differ wildly. The ancestral reconstruction of the thermal niche indicated that this group of Neotropical bats evolved under cooler temperatures. The two clades inside the Glossophaginae differ in the evolution of their thermal niches, with most members of the clade Choeronycterines evolving “colder” thermal niches, while the majority of the species in the clade Glossophagines evolving “warmer” thermal niches. By comparing thermal niches with climate change models, we found that all species could be affected by an increase of 1°C in temperature at the end of this century. This suggests that even nocturnal species could suffer important physiological costs from global warming. Our study highlights the value of scientific collections to obtain ecologically significant physiological data for a large number of species.

Chromosomal evolution among leaf-nosed nectarivorous bats – evidence from cross-species chromosome painting (Phyllostomidae, Chiroptera)

BackgroundNew World leaf-nosed bats, Phyllostomidae, represent a lineage of Chiroptera marked by unprecedented morphological/ecological diversity and extensive intergeneric chromosomal reorganization. There are still disagreements regarding their systematic relationships due to morphological convergence among some groups. Their history of karyotypic evolution also remains to be documented.ResultsTo better understand the evolutionary relationships within Phyllostomidae, we developed chromosome paints from the bat species Macrotus californicus. We tested the potential of these paints as phylogenetic tools by looking for chromosomal signatures in two lineages of nectarivorous phyllostomids whose independent origins have been statistically supported by molecular phylogenies. By examining the chromosomal homologies defined by chromosome painting among two representatives of the subfamily Glossophaginae (Glossophaga soricina and Anoura cultrata) and one species from the subfamily Lonchophyllinae (Lonchophylla concava), we found chromosomal correspondence in regions not previously detected by other comparative cytogenetic techniques. We proposed the corresponding human chromosomal segments for chromosomes of the investigated species and found two syntenic associations shared by G. soricina and A. cultrata.ConclusionComparative painting with whole chromosome-specific paints of M. californicus demonstrates an extensive chromosomal reorganization within the two lineages of nectarivorous phyllostomids, with a large number of chromosomes shared between M. californicus and G. soricina. We show that the evolution of nectar-feeding bats occurs mainly by reshuffling of chiropteran Evolutionarily Conserved Units (ECUs). Robertsonian fusions/fissions and inversions seem to be important modifiers of phyllostomid karyotypes, and autapomorphic character states are common within species. Macrotus californicus chromosome paints will be a valuable tool for documenting the pattern of karyotypic evolution within Phyllostomidae radiation.

New Genus and Species of Nectar-Feeding Bat from the Atlantic Forest of Southeastern Brazil (Chiroptera: Phyllostomidae: Glossophaginae)

The lowland Atlantic Forest of southeastern Brazil is well known for its biological diversity and numerous endemic taxa. Among bats collected recently at the Reserva Natural Vale and previously in the Floresta Nacional de Goytacazes, in the state of Espírito Santo, are specimens easily recognized as nectar-feeding glossophagines, but that exhibit a suite of morphological traits that preclude assignment to any of the 10 genera currently recognized in the subfamily Glossophaginae. Here we describe a new genus and species based on this material. This new taxon, named Dryadonycteris capixaba, is diagnosed based on both external and craniodental features, including traits not seen in other nectar-feeding phyllostomids, such as similar-sized calcar and foot and inflated maxillary bones. The combination of character states seen in Dryadonycteris suggests that it belongs in the Tribe Choeronycterini, subtribe Choeronycterina, but the mosaic nature of primitive and derived states seen in this taxon precludes easy assessment of its relationships to other choeronycterine genera. Future explicit phylogenetic analyses of morphological data and DNA sequencing studies will be necessary to resolve its phylogenetic position within Choeronycterini.

Nectar bat stows huge tongue in its rib cage

Bats of the subfamily Glossophaginae (family Phyllostomidae) are arguably the most specialized of mammalian nectarivores, and hundreds of neotropical plants rely on them for pollination. But flowers pollinated by bats are not known to specialize for bat subgroups (unlike flowers that have adapted to the length and curvature of hummingbird bills, for example), possibly because the mouthparts of bats do not vary much compared with the bills of birds or the probosces of insects. Here I report a spectacular exception: a recently-described nectar bat that can extend its tongue twice as far as those of related bats and is the sole pollinator of a plant with corolla tubes of matching length.

NEW GENUS AND SPECIES OF NECTAR-FEEDING BAT IN THE TRIBE LONCHOPHYLLINI (PHYLLOSTOMIDAE: GLOSSOPHAGINAE) FROM NORTHEASTERN BRAZIL

Abstract A new genus and species in the tribe Lonchophyllini, subfamily Glossophaginae (Chiroptera: Phyllostomidae) is described based on the analysis of 4 specimens collected in 3 different localities in a semiarid area of northeastern Brazil. The presence of a groove and filiform papillae on the lateral side of the tongue places this taxon as a member of the tribe Lonchophyllini. Fourteen cranial and dental characters distinguish the new genus from all other genera in the tribe (Lonchophylla, Lionycteris, and Platalina). The new genus is characterized by the presence of inward-facing molars on the palate; the loss of the cingulum in P3 and P4; the reduction or lack of the metastyle and preprotocrest in M1, ectoflexus, preprotocrest, and postprotocrest in M2, and postprotocrest in M3; and loss of hypoflexid, metacristid, entocristid, and entoconid on m1–3. A wider gap also occurs between lower incisors and canines than in other genera of Lonchophyllini. The palate in the new genus presents an unusual mor...

Cytogenetic analysis in species of the Subfamily Glossophaginae (Phyllostomidae, Chiroptera) supports a polyphyletic origin

Abstract In this work the karyotypes of 20 specimens of bats belonging to the subfamily Glossophaginae (Phyllostomidae-Chiroptera) were analyzed, species Glossophaga soricina (2n=32, FN=60), Lionycteris spurrelli (2n=28, FN=50), Lonchophylla thomasi (2n=36, FN=48) and Choeronicus minor (2n=20, FN=36), whose data were compared with each other and with data from the literature. The analyses were done using G, C, G/C sequential and NORs banding techniques and hybridization in situ with telomeric probes. Published data were also used for Artibeus lituratus (Stenodermatinae). The species G. soricina, L. spurrelli and L. thomasi presented few homeologies among themselves and none with C. minor; the latter has a pair of chromosomes shared with the species Artibeus lituratus (Stenodermatinae), indicating a possible convergent evolution in the subfamily Glossophaginae. Additionally, by comparing our results with those in the literature, interpopulational karyotypic variability was observed in the species L. thomasi.

Pollen Carrying by the Nectar-Feeding Bat Glossophaga soricina in a Suburban Environment

Pollen Carrying by the Nectar-Feeding Bat Glossophaga soricina in a Suburban Environment

On the Phylogeny of the Glossophaginae and the Proper Use of Outgroup Analysis

In two recent papers on the phylogeny of phyllostomid bats, Hood and Smith (1982) and Haiduk and Baker (1982) commented unfavorably on my recent cladistic analysis of the New World nectar-feeding bats (Griffiths, 1982). Both pairs of authors suggested that (1) my outgroup comparisons were incorrect, and (2) my character analysis was incorrectly performed. Haiduk and Baker (1982) went so far as to redo the character analysis and claimed that my hyoid/tongue morphology data better supported the branching sequence suggested by them than the one I presented (Griffiths, 1982). Despite the alleged errors in my analysis, both Hood and Smith (1982) and Haiduk and Baker (1982) admitted that at least parts of my conclusions seemed to be correct. Neither paper mentioned that for the first time in the 15 years the nectar-feeding bat problem has been intensively examined, two phylogenies-produced independently and based on unrelated data sets-are remarkably congruent (Baker et al. [1981] and Haiduk and Baker [1982] using biochemical and karyotypic data; Griffiths [1982] using hyoid/ tongue morphology data). Except for very minor differences in the exact relationship of Hylonycteris and Choeroniscus and a minor disagreement on the relationships of the three brachyphylline genera, the only difference in the two cladograms produced is in the treatment of the genera Lonchophylla and Lionycteris. Haiduk and Baker (1982) indicated that the two genera are relatively derived members of the subfamily Glossophaginae. On the other hand, I suggested (Griffiths, 1982) that the two genera evolved nectivory independently of the true glossophagines. I indicated further that no synapomorphous character state was known that united Lonchophylla and Lionycteris with the true glossophagines and, until such a character state was found, Lonchophylla and Lionycteris (along with Platalina which Haiduk and Baker did not karyotype) might best be placed in a separate subfamily, the Lonchophyllinae. In this note, I respond to the allegations that my outgroup comparisons were incorrect. In addition, I show that Haiduk and Baker (1982) are quite incorrect in their assertion that my hyoid/tongue data support the placement of Lonchophylla and Lionycteris as relatively derived glossophagine bat genera. Outgroup comparisons are made to determine correct character polarity within the group one is studying and, most especially, to determine which character states are synapomorphies, which are autapomorphies, and which are symplesiomorphies. When studying the relationships of the genera within the taxonomic group A, it is obviously important to study also outgroup representatives from all closely related sister groups (B and C), as well as representatives from more primitive groups (D) that are thought to be derived from the direct ancestor of A, B, and C. Without so doing, the systematist fails to get an accurate idea of the polarity of characters and of the uniqueness of the synapomorphies in the group (A) in question. Unitl recently, in a study of the relationships within the taxon Glossophaginae, there was no way of determining which other groups were closely-related sister groups (B and C) and which were modern representatives of ancestral groups (D). This was because no study had been done that showed the relationships of the Glossophaginae to other bat taxa. In the absence -of a definitive study, Haiduk and Baker (1982) are entitled to their opinion of the proper outgroups to use, as am I. Such a definitive study (Hood and Smith, 1982) has now been done,

A NEW SPECIES OFPARATRICHOBIUS(DIPTERA: STREBLIDAE) FROM ARIZONA

AbstractParatrichobius americanusnew species is described and illustrated from specimens collected in Arizona, from the host batChoeronycteris mexicanaTschudi It is most closely related to theP.longicrus(Miranda Ribeiro) complex of species, but is easily differentiated from all the known species by details of the chaetotaxy of the profemur, mesonotum and gonapophyses, and by the small eyes. This is the first species of the genus to be recorded from the bat subfamily Glossophaginae, all others are known only from the Stenoderminae (family Phyllostomidae).

Three new species of Radfordiella (Acarina: Macronyssidae) parasitic in the mouth of phyllostomatid bats.

Three species of Radfordiella are described from the protonymphs: R. oricola from Leptonycteris nivalis and Anoura geoffroyi; R. anourae from Anoura geoffroyi; and R. monophylli from Monophyllus redmani. Protonymphs of the new species were found embedded in protected sites in tissue of the soft palate near the first premolar. Other stages are unknown. Hosts are all in the subfamily Glossophaginae of the chiropteran family Phyllostomatidae. Radfordiella is rediagnosed based on the protonymphs and a key is given to protonymphs of the 6 described species. The relationships of species in the genus and the morphology of the protonymphs are discussed. Comments are made on some aspects of the host-parasite relationships.

Karyotypes of Bats of the Family Phyllostomidae and Their Taxonomic Implications

A total of 143 bats representing 27 species of the Family Phyllostomidae were analyzed for their karyotypes. Analysis of karyotypic variation revealed no population or geographic variation and, within a given genus, only variation in the Y chromosome(s) of Artibeus was found. Based on similarity of karyotypes the following seven groups are evident: (A) Pteronotus; (B) Choeronycteris, Choeroniscus and Carollia; (C) Leptonycteris, Glossophaga, Phyllostomus, Trachops and Macrotus; (D) Micronycteris; (E) Anoura; (F) Artibeus, Sturnira, Vampyrops, Chiroderma, Enchisthenes and Centurio; and (G) Uroderma. The Subfamily Sturnirinae is considered to be a synonym of Subfamily Stenoderminae. Subfamily Glossophaginae is considered to be of polyphyletic origin but available data are inadequate for interpretation of their phylogeny. The status of Uroderma as a genus distinct from Artibeus is strongly supported. This study was designed to evaluate current concepts concerning the phylogeny of bats of the family Phyllostomidae by utilizing data from studies of karyotypes. METHODS AND MATERIALS. The techniques herein used were essentially those described by Patton (1967), with the modifications described by Baker and Patton (1967). Phyllostomid bats (especially certain species) often died within 24 hours or less after capture, and attempts to transport live specimens to a laboratory were unproductive. For this reason, many cytological slides were prepared in the field where a portable electric generator was used to power the centrifuge. Most bats were collected during the night in mist nets, and processed the next day at or near the collection site. By processing specimens in the field, few bats were lost. The only 1 Supported in part by National Institute of Health Predoctoral Fellowship No. 1-F1-GM-28,182-01 and 02 and by Research Grant GB-1867 from National Science Foundation. 2 Present address: Department of Biology, Texas Technological College, Lubbock, Texas, 79409.

Muscular Adaptation of the Nectar-Eating Bats (Subfamily Glossophaginae)

Some possible correlations among bats between structures and functions connected with alimentation were suggested by Lautenschlager (1935) and later by Park and Hall (1951). It seemed to me that a comparison of the musculature of the throat among bats that had different food habits might show some modification of these muscles in correlation with certain specialized food habits. After a general survey of bats representing different food habits, it was found that only the nectar-eating bats of the Subfamily Glossophaginae, had a highly modified musculature of the throat. Some of these modifications are easily correlated with the specialized food habits of these bats. The most striking adaptations of the musculature are the new origin and insertion of the sternohyoideus, and a highly modified geniohyoideus. The specimens dissected were as follows: Leptonycteris nivalis (Saussure), Nos. 34181, 34198-206,34509-11,34515-20,1 34522. Five mi. W Chapala, 5000 ft., Jalisco, Mexico. July, 1949. Glossophaga elongata Miller, Nos. 105221, 105233. Mal Paies, Curacao, Dutch West Indies. Glossophaga soricina (Pallas), No. 257249. Chilibrillo, Panama. March, 1932. Glossophaga soricina (Pallas), No. 34510. Five mi. W Chapala, 5000 ft., Jalisco, Mexico. July, 1949. Lonchophylla robusta Miller, Nos.; 266808, 266909. Chilibrillo, Panama. March, 1932. Choeronycteris mexicana Tschudi, Nos. 31985 (21 mi. SW Guadalajara), 31988 (N side Rio Ayuguida, 7 mi. NE Autlan). State of Jalisco, Mexico. August, 1950. Anoura geoffroyi Gray, No. 92582. San Sebastian, Jalisco, Mexico. March 21, 1897. Monophyllus cubanus Miller, No. 113677. Baracoa, Eastern Cuba. February 3, 1902. Before describing the throat musculature of the nectar-eating bats, it is advisable to discuss some general points concerning the musculature of the throat of bats of the suborder Microchiroptera. Some of the muscles, such as hyoglossus, sternohyoideus, and geniohyoideus, that have connection with the hyoid bone in mammals of other orders are not connected directly with that bone in the Microchiroptera in