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The majority (90%) of native terrestrial mammal species living in the Dominican Republic are bats, and two-thirds of these species are endemic to the Caribbean. However, recent molecular studies using DNA barcoding of the mitochondrial cytochrome c oxidase subunit 1 gene have suggested at least a 25% underestimation of biodiversity in bats throughout the world. A recent survey of bats in the Dominican Republic documented 15 of the 18 known species on the island of Hispaniola. Phylogenetic analysis of 132 individuals resulted in well-supported monophyletic species-level clades (maximal bootstrap values) with intraspecific variation ranging from 0% to 4.7% and interspecific variation ranging from 14.1% to 32.5%. A phylogeographic pattern separating the northern and southern Dominican Republic was recovered in 3 species of bats (Macrotus waterhousii, Pteronotus parnellii, and Pteronotus quadridens). The inclusion of broader geographic sampling across the Neotropics indicated that 3 widely distributed species (Eptesicus fuscus, Molossus molossus, and Monophyllus redmani) have high sequence divergence among insular or between insular and continental populations. Further systematic study is needed to identify morphologically cryptic species and their implications for conservation priorities in the Caribbean.

Pteronotus quadridens Pteronotus quadridens fuliginosus: United States: Puerto Rico, San Juan, Pueblo Viejo, Cueva de Fari (AMNH 39353–59, 39363–67); Trujillo Alto (39396–03); Dominican Republic: between Cabra and Nagua, Cueva de Murcielagos (AMNH 238138); La Altagracia, Boca de Yuma (AMNH 238139); Santo Domingo, Cana Honda (AMNH 25760); Haiti: Cerca La Source (USNM 253631–32); Sud, Sapoti (19 km SW Beaumont) (AMNH 236654); Jamaica: St. Elizabeth, Balaclava, Oxford Cave (AMNH 45248– 55); St. Elizabeth, Accompong Town (UF-M 12684). Pteronotus quadridens torrei: Cuba: [no additional information] (AMNH 60637–38); Oriente, 8 mi E Baracoa (USNM 113786, 121061); La Habana, Bararanao, Cueva de El Fantasma (AMNH 176061); Las Villo, Punta Caquanes, Yaguajay “Cueva de Colon” (AMNH 186970–71). Pteronotus personatus species group Pteronotus psilotis: Mexico: Colima, Ixtlahuacán, 1.6 km NE Las Conchas (OMNH 27773); Colima, Manzanillo, Playa de Oro, Cueva la Murcielaguera (OMNH 38769); Morelos, Tequesquitengo (AMNH 180443–44, 180446); Nayarit, Nayarit-Sinaloa state line, Rio Canas, near la Concha (USNM 508415); Oaxaca, Juchitan, Tapanatpec (AMNH 175018–19, 177672–93, 177695–96, 178468); Sonora, La Aduana (MSB 55419); Vera Cruz, Agua Fria (MSB 75556). Pteronotus personatus: Colombia: Bolivar ( USNM 430190, 433531); Bolivar, Cartagena (USNM 433517, 433531); Panama: San Blas, Armila (Quebrada Venado) (USNM 335085–86); Venezuela: Sucre, La Brea (AMNH 33300); Yara- cui, Aroa, Bolivar R.R. (33112, 32123–24, 32127– 32133, 32136–37); Trinidad and Tobago: St. George, Tacariquas; St. Augustine (AMNH 175560)., Published as part of Morgan, Gary S., Czaplewski, Nicholas J. & Simmons, Nancy B., 2019, A New Mormoopid Bat From The Oligocene (Whitneyan And Early Arikareean) Of Florida, And Phylogenetic Relationships Of The Major Clades Of Mormoopidae (Mammalia: Chiroptera), pp. 1-141 in Bulletin of the American Museum of Natural History 2019 (434) on page 138, DOI: 10.1206/0003-0090.434.1.1, http://zenodo.org/record/4613816

Se estudiaron los patrones de emergencia y estados reproductivos de las poblaciones de murciélagos en cuatro cuevas de la República Dominicana. Se capturó un total de 12 especies: Mormoops blainvillei, Pteronotus parnellii, Pteronotus quadridens, Chilonatalus micropus, Natalus major, Artibeus jamaicensis, Brachyphylla pumila, Erophylla bombifrons, Macrotus waterhousii, Monophyllus redmani, Phyllonycteris poeyi y Eptesicus fuscus. Los murciélagos se capturaron de forma más o menos continua entre las 18:15 y 23:00h. De un total de 1,445 ejemplares a los cuales se les determinó el sexo, 969 fueron machos (67.1%). Los resultados obtenidos con respecto a la actividad reproductiva muestran cómo las especies presentes en las cuevas se encuentran en estados reproductivos diferentes, registrándose ejemplares preñadas o lactantes, pertenecientes a dos familias dominantes: Mormopidae y Phyllostomidae. Además, que se contrastaron reportes de Cuba y Puerto Rico donde se muestra que algunas especies podrían tener épocas reproductivas más largas que las reportadas actualmente.

Molecular approaches, such as polymerase chain reaction (PCR), allow biologists to analyze the diet of mammals in more detail than conventional techniques, by targeting DNA of prey found within feces. Diets of the Antillean ghost-faced bat (Mormoops blainvillei) and sooty mustached bat (Pteronotus quadridens) on Puerto Rico were determined via PCR, and resulting sequences of DNA were compared with those in the Barcode of Life Data Systems (BOLD). We identified 21 species consumed by M. blainvillei and 4 by P. quadridens, although an additional 12 sequences were identified to genus or family. Seven taxa in the diet of these bats were of agricultural or human-health concern. All sequences, including those that did not match a reference insect in BOLD, were collapsed into molecular operational taxonomic units (MOTUs) and used to demonstrate that dietary breadth (Levin's measure) of M. blainvillei was 18% less than that of P. quadridens and that dietary overlap (Schoener's index) between species was ...

Dietary differentiation can be a key mechanism for the coexistence of syntopic species with similar niches. On the Caribbean island of Puerto Rico, three species of bat from the family Mormoopidae — the Antillean ghost-faced bat (Mormoops blainvillei), sooty mustached bat (Pteronotus quadridens), and Parnell's mustached bat (Pteronotus parnellii portoricensis) — are aerial insectivores that roost in the same caves. To investigate the possibility of dietary differentiation, we estimated the percent volume and percent frequency of occurrence of the orders of arthropods consumed by these three species of bat, using standard fecal analysis. We also compared dietary diversity among species, as well as the amount of dietary overlap, with respect to season and habitat. Lastly, this study used canonical correspondence analysis (CCA), a method of ordination, to assess the effects of species, sex, age, reproductive condition, season, and habitat on intraspecific differences in the diet of the Puerto Rican Mormoopidae. Eight orders of arthropods were found in the diet of these mormoopids, with Lepidoptera, Hymenoptera, and Coleoptera being major staples. The CCA revealed differences in diet among the three species, suggesting that dietary differentiation is at least one mechanism for coexistence. In addition, the variables habitat and season correlated significantly with the diet of M. blainvillei and P. quadridens, whereas habitat and sex correlated with the diet of P. p. portoricensis. Thus, our study shows dietary differences among the three species of Mormoopidae living in the same caves on Puerto Rico, as well as intraspecific differences within the diet of each species.

Free-roaming cats are known to adversely impact native faunas in the areas where they have been introduced, an impact that is even greater on islands. We examine the predation of bats by cats at Culebrones cave, Puerto Rico, West Indies. Culebrones cave is a hot cave located in the karst region of northern Puerto Rico. The temperature gradient inside the cave sustains a multi-species assemblage of bats consisting of approximately 300,000 individuals of six species, namely: Brachyphylla cavernarum, Erophylla bombifrons, Monophyllus redmani, Mormoops blainvillei, Pteronotus quadridens and Pteronotus parnellii. Even though rats are often their primary prey, cats will use alternative prey, which enables them to maintain their abundance when one prey is not available. In Puerto Rico, birds and reptiles are known to be preyed upon by cats. Although cats are commonly observed in or around bat caves in Puerto Rico, this is the first systematic attempt to evaluate their role as bat predators. We made observations of the hunting strategy of cats using an infrared camera and recorded the number of wings left as remains of these hunting bouts. Wings were identified to species. Cat scats were also recovered and examined to identify prey species. Our results suggest that captures of different species of bats is not a function of their abundance in the cave. While M. blainvillei (11 g) and P. quadridens (5 g) are the most abundant species in the cave, B. cavernarum (50 g) and M. redmani (11 g) are captured in greater numbers by the cats.

Bats are receiving increasing attention in the parasitological world due to their potential role as reservoir hosts for zoonotic pathogens. However, details of the macroparasitic fauna (helminths and ectoparasites) are currently recorded and studied. Here, we start to address this paucity of data by detailing a study where we have documented the macroparasite fauna of a sample of tropical bats (Mormoops blainvillei, Pteronotus quadridens, and Monophyllus redmani) from Puerto Rico. Additionally, we investigated the possible host characteristics influencing the prevalence and intensity of macroparasite infection. Macroparasites were collected and identified from three species of bat, which were thoroughly washed and dissected. The overall parasite community of all three bat species consisted of a range of ectoparasites as well as the cestode Vampirolepis christensoni and the nematode Capillaria pusilla, although there was considerable variation in the parasite community of each individual species. We discovered bat flies of a previously undescribed species of the Nycterophilia genus as well as new parasite records for all three species of bats. All parasites had an aggregated distribution within the host population. Differences were observed in the intensity of the helminths between bat species, but not for ectoparasite prevalence. As the helminth intensity increased so the ectoparasite intensity decreased. Overall, the helminth intensity was female-biased and increased, for both sexes with increasing body mass; no sex-bias or body mass effects were associated with ectoparasite prevalence.

Echolocation calls were recorded from Pteronotus quadridens flying in the field and in an enclosed space. In the field, search calls contained 1 or 2 harmonics. Patterns of call design show a segment of quasi–constant frequency (QCF2nd-harmonic at 81–84 kHz), followed by a downward frequency-modulated (FM) component. The 2nd harmonic was always more intense than the 1st. Search, approach, and terminal phases of calls were described during hunting sequences of P. quadridens. The transition between call phases was characterized by monotonic variations in some acoustic parameters, including a decrease in call duration and an increase in repetition rate, bandwidth, and slope of the FM component. We also analyzed calls emitted by bats flying in confined spaces that consistently contained 3 harmonics, of which the 2nd harmonic contained the greatest energy. The values of call duration were shorter and bandwidth was higher than values characterizing calls emitted during the search phase in the field.

The temperature preferences of the mormoopid and phyllostomid bats Pteronotus quadridens and Erophylla sezekorni, from the West Indies, were determined in the laboratory and compared to field observations. Pteronotus quadridens was invariably found within the deepest and hottest parts of caves, at temperatures between 28°C and 35°C, while E. sezekorni was found at temperatures from 25°C to 28°C. Temperatures selected by each species in a thermopreferendum chamber were similar to their respective roosting temperatures in the caves. These inter-specific differences are statistically significant. Our results support the hypothesis that roost temperature and differences in temperature preferences among species, are important in explaining multispecies associations and the spatial segregation within caves.

In this investigation, we use variation in wing morphology, echolocation behaviour, patterns of habitat use and molecular diet analysis to demonstrate that six species of sympatric insectivorous bats in Jamaica show significant differences that could explain resource partitioning among the species. High-intensity echolocating species that used shorter, broadband signals and had shorter, broader wings (Pteronotus macleayii, Pteronotus quadridens, Mormoops blainvillii) foraged most in edge habitats, but differed in timing of peak activity. P. macleayii and M. blainvillii differed in diet, but low sample size precluded diet analysis for P. quadridens. High-intensity echolocating species that used longer, more narrowband signals and had longer, narrower wings (Molossus molossus, Tadarida brasiliensis) foraged most in open areas and differed in diet from the other species. Two disparate species were most active in clutter (dense vegetation). Pteronotus parnellii used high-duty-cycle echolocation apparently specialized for detecting fluttering targets in clutter. Macrotus waterhousii used low-intensity, broadband echolocation calls and presumably uses prey-generated sounds when foraging. These two species also differed in diet. Our data show that differences in morphology and echolocation behaviour coincide with differences in habitat use and diet, resulting in minimal overlap in resource use among species.

Naked-backed bats of the genus Pteronotus (family Mormoopidae) occur in the Neotropics from Mexico through northern South America. These are relatively small-sized insectivorous species that frequently roost in caves. Eight specimens of naked-backed bats (Pteronotus parnellii) were livetrapped in Suriname and one in Cuba (P. quadridens). Their parotid glands were fixed in an aldehyde mixture designed for field work and postfixed in the laboratory with osmium tetroxide. Tissues were further prepared for electron microscopy by conventional means. The parotid glands of the two species of Pteronotusclosely resemble each other except for the substructure of their serous secretory granules. Serous granules in P. parnellii are bizonal, with a moderately dense inner matrix and an outer, denser corona or crescent. The matrix is occupied by laminae, flakes, and filaments in random array. In contrast, serous granules in P. quadridens consist of a uniform matrix that contains dense, usually stacked toroids or tubules either in random array or packed in bundles. A parotid gland from one specimen of P. parnellii contained an endpiece that consisted of cells that contained giant (up to 9 µm in diameter) serous granules. Serous cells in both species contain aggregates of small, uniformly dense, rod-like, membrane-delimited organelles as well as occasional bundles of cytofilaments. The endpieces are separated from intercalated ducts by a ring of granulated cells that contain secretory granules that often have a bull’s-eye configuration. Intercalated and striated ducts are typical in appearance, except that many of the cells in the latter contain small, dense secretory granules in their apical cytoplasm. The parotid glands in the two species of naked-baked bats differ slightly in terms of acinar secretory granule ultrastructure, but otherwise are fairly conservative. It is thought that the glands in these particular bats might represent the ‘‘basal’’ condition of the salivary glands of insectivorous bats and thus can serve as a reference point for making comparisons to the highly diversified (in terms of diet) phyllostomid bats. Anat Rec 255:105‐115, 1999. r 1999 Wiley-Liss, Inc.

I surveyed use of caves by bats in Puerto Rico. The exact location of each species of bat within the roost was determined for 17 caves. A comparison of variance was performed on the data to test the null hypothesis that occurrence of each species was independent of the others. Nonrandom associations were common among cave-dwelling bats in Puerto Rico. The species showing the strongest associations— Monophyllus redmani, Mormoops blainvillii , and Pteronotus quadridens and Erophylla sezekorni and P. parnellii —were associated with hot caves.

Echolocating bats use the time from biosonar pulse emission to the arrival of echo (defined as echo delay) to calculate the space depth of targets. In the dorsal auditory cortex of several species, neurons that encode increasing echo delays are organized rostrocaudally in a topographic arrangement defined as chronotopy. Precise chronotopy could be important for precise target-distance computations. Here we show that in the cortex of three echolocating bat species (Pteronotus quadridens, Pteronotus parnellii and Carollia perspicillata), chronotopy is not precise but blurry. In all three species, neurons throughout the chronotopic map are driven by short echo delays that indicate the presence of close targets and the robustness of map organization depends on the parameter of the receptive field used to characterize neuronal tuning. The timing of cortical responses (latency and duration) provides a binding code that could be important for assembling acoustic scenes using echo delay information from objects with different space depths.

Pteronotus quadridens (Gundlach, 1840) Figures 3, 5 MATERIAL EXAMINED: Cueva de Lily: 1 complete skull, 2 skull fragments, 1 dentary, 2 radii. Oleg’s Bat Cave: 1 complete skull, 1 dentary. EXTANT DISTRIBUTION: Cuba, Hispaniola, Jamaica, and Puerto Rico (Dávalos and Turvey, 2012; Simmons, 2005). FOSSIL RECORD: Pteronotus quadridens has previously been recovered from a Quaternary cave deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table 1). Additionally, P. quadridens has been found in Pleistocene or Holocene cave deposits in Cuba (Silva Taboada, 1974, 1979; Woloszyn and Silva Taboada, 1977) and the Bahamas (Andros, Great Abaco, and New Providence; Morgan, 2001). REMARKS: No consistent differences in cranial or postcranial morphology or size were found between our sample and the comparative material (appendix)., Published as part of Velazco, Paúl M., O'Neill, Hannah, Gunnell, Gregg F., Cooke, Siobhán B., Rimoli, Renato, Rosenberger, Alfred L. & Simmons, Nancy B., 2013, Quaternary Bat Diversity in the Dominican Republic, pp. 1-20 in American Museum Novitates 2013 (3779) on page 11, DOI: 10.1206/3779.2, http://zenodo.org/record/5364403, {"references":["Davalos, L. M., and S. Turvey. 2012. West Indian mammals: the old, the new, and the recent. In B. D. Patterson and L. P. Costa (editors), Bones, clones, and biomes. The history of recent Neotropical mammals: 157 - 202. Chicago: University of Chicago Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world: a taxonomic and geographic reference. 3 rd ed., 1: 312 - 529. Baltimore, MD: Johns Hopkins University Press.","Silva Taboada, G. 1974. Fossil Chiroptera from cave deposits in central Cuba, with description of two new species (genera Pteronotus and Mormoops) and the first West Indian record of Mormoops megalophylla. Acta Zoologica Cracoviensia 19: 33 - 73.","Silva Taboada, G. 1979. Los murcielagos de Cuba. La Habana: Editorial Academia, Academia de Ciencias.","Woloszyn, B. W., and G. Silva Taboada. 1977. Nueva especie fosil de Artibeus (Mammalia: Chiroptera) de Cuba, y tipificacion preliminar de los depositos fosiliferos cubanos contentivos de mamiferos terrestres. Poeyana 161: 1 - 17.","Morgan, G. S. 2001. Patterns of extinction in West Indian bats. In C. A. Woods and F. E. Sergile (editors), Biogeography of the West Indies: patterns and perspectives: 369 - 406. Boca Raton: CRC Press."]}

Echolocating bats use the time elapsed from biosonar pulse emission to the arrival of echo (defined as echo-delay) to assess target-distance. Target-distance is represented in the brain by delay-tuned neurons that are classified as either “heteroharmonic” or “homoharmormic.” Heteroharmonic neurons respond more strongly to pulse-echo pairs in which the timing of the pulse is given by the fundamental biosonar harmonic while the timing of echoes is provided by one (or several) of the higher order harmonics. On the other hand, homoharmonic neurons are tuned to the echo delay between similar harmonics in the emitted pulse and echo. It is generally accepted that heteroharmonic computations are advantageous over homoharmonic computations; i.e., heteroharmonic neurons receive information from call and echo in different frequency-bands which helps to avoid jamming between pulse and echo signals. Heteroharmonic neurons have been found in two species of the family Mormoopidae (Pteronotus parnellii and Pteronotus quadridens) and in Rhinolophus rouxi. Recently, it was proposed that heteroharmonic target-range computations are a primitive feature of the genus Pteronotus that was preserved in the evolution of the genus. Here, we review recent findings on the evolution of echolocation in Mormoopidae, and try to link those findings to the evolution of the heteroharmonic computation strategy (HtHCS). We stress the hypothesis that the ability to perform heteroharmonic computations evolved separately from the ability of using long constant-frequency echolocation calls, high duty cycle echolocation, and Doppler Shift Compensation. Also, we present the idea that heteroharmonic computations might have been of advantage for categorizing prey size, hunting eared insects, and living in large conspecific colonies. We make five testable predictions that might help future investigations to clarify the evolution of the heteroharmonic echolocation in Mormoopidae and other families.

Delay tuning was studied in the auditory cortex of Pteronotus quadridens. All the 136 delay-tuned units that were studied responded strongly to heteroharmonic pulse-echo pairs presented at specific delays. In the heteroharmonic pairs, the first sonar call harmonic marks the timing of pulse emission while one of the higher harmonics (second or third) indicates the timing of the echo. Delay-tuned units are organized chronotopically along a rostrocaudal axis according to their characteristic delay. There is no obvious indication of multiple cortical axes specialized in the processing of different harmonic combinations of pulse and echo. Results of this study serve for a straight comparison of cortical delay-tuning between P. quadridens and the well-studied mustached bat, Pteronotus parnellii. These two species stem from the most recent and most basal nodes in the Pteronotus lineage, respectively. P. quadridens and P. parnellii use comparable heteroharmonic target-range computation strategies even though they do not use biosonar calls of a similar design. P. quadridens uses short constant-frequency (CF)/frequency-modulated (FM) echolocation calls, while P. parnellii uses long CF/FM calls. The ability to perform “heteroharmonic” target-range computations might be an ancestral neuronal specialization of the genus Pteronotus that was subjected to positive Darwinian selection in the evolution.

Data are presented on metabolic rates and thermal conductance for four species of neotropical bats, Pteronotus quadridens, Mormoops blainvillii (Mormoopidae), Monophyllus redmani and Erophylla bombifrons (Phyllostomidae). Each of these bats predominantly or exclusively roosts in hot caves (28–40°C) in Puerto Rico. Basal metabolic rates (BMR) for these four species were 55%, 48%, 66%, and 66% of values expected from the Kleiber curve, respectively. Thermal conductance was 93%, 55%, 175%, and 158% of values expected from the Aschoff curve, respectively. These data indicate that, in addition to food habits, variation in BMR is highly correlated to roost microclimate. Pteronotus and Mormoops, like other insectivorous species, have low BMRs. The low BMR of the nectar/fruit eating Monophyllus and Erophylla, as well as the low termal conductance in Mormoops, is consistent with their habit of roosting in hot caves, suggesting that roost microclimate is an important selective force in the physiological adaptation of these bats.

Pteronotus quadridens (Gundlach, 1840). Arch. Naturgesch., 6:357. TYPE LOCALITY: Cuba, Matanzas, Canimar. DISTRIBUTION: Cuba, Jamaica, Hispaniola, Puerto Rico. SYNONYMS: fuliginosus, inflata, torrei. COMMENTS: Subgenus Chilonycteris. Includes torrei; For use of quadridens in place of fuliginosus; see Silva-Taboada (1976:7). See Rodrlguez-Durän and Kunz (1992, Mammalian Species, 395)., Published as part of Karl F. Koopman, 1993, Order Chiroptera, pp. 137-241 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 177, DOI: 10.5281/zenodo.7353060, {"references":["Silva-Taboada, G. 1976. Historia y actualizacion taxonomica de algunas especies Antillanas de murcielagos de los generos Pteronotus, Brachyphylla, Lasiurus, y Antrozous (Mammalia: Chiroptera). Poeyana (Academia de Ciencias de Cuba), 153: 1 - 24.","Kunz, T. H., and I. M. Pena. 1992. Mesophylla macconnelli. Mammalian Species, 405: 1 - 5."]}