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Research Article
Islands in the stream: Distribution of Myotis attenboroughi (Chiroptera, Vespertilionidae) in Grenada and mainland South America illuminates the evolutionary history of Caribbean Myotis
expand article infoRoberto Leonan M. Novaes‡, Vinícius C. Cláudio‡§, Natasha A. Bertocchi‡, Edson F. Abreu|, Don E. Wilson, Jesús E. Maldonado#, Ricardo Moratelli‡
‡ Fundação Oswaldo Cruz, Rio de Janeiro, Brazil
§ JGP Consultoria e Participações Ltda., São Paulo, Brazil
| Angelo State University, San Angelo, United States of America
¶ National Museum of Natural History, Washington DC, United States of America
# Smithsonian’s National Zoological Park and Conservation Biology Institute, Washington, DC, United States of America

Corresponding author: Roberto Leonan M. Novaes ( robertoleonan@gmail.com )

Academic editor: Danilo Harms
© 2025 Roberto Leonan M. Novaes, Vinícius C. Cláudio, Natasha A. Bertocchi, Edson F. Abreu, Don E. Wilson, Jesús E. Maldonado, Ricardo Moratelli.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation: Novaes RLM, Cláudio VC, Bertocchi NA, Abreu EF, Wilson DE, Maldonado JE, Moratelli R (2025) Islands in the stream: Distribution of Myotis attenboroughi (Chiroptera, Vespertilionidae) in Grenada and mainland South America illuminates the evolutionary history of Caribbean Myotis. Evolutionary Systematics 9(1): 107-122. https://doi.org/10.3897/evolsyst.9.146107
ZooBank: urn:lsid:zoobank.org:pub:1A53FE99-9955-4426-99AB-EC0C8687CBB0
Open Access

Abstract

Myotis attenboroughi was recently described based on specimens from Tobago Island and considered an endemic species until now. Based on morphological and molecular data, we extended its occurrence to Grenada and Suriname. The presence of M. attenboroughi in Grenada, Tobago and mainland South America can be explained by the intermittent connections between these islands and the continent during the Plio-Pleistocene glaciations. However, our analyses recovered an Myotis individual from Grenada phylogenetically related to Myotis nyctor from Barbados, although it is morphologically like the other specimens of M. attenboroughi, revealing a surprising incongruence between genotype and phenotype. Barbados is a geologically recent island and historically unconnected with other Caribbean islands or with South America. We hypothesized that the invasion of the M. nyctor lineage from Barbados to Grenada was the result of an extreme overwater dispersal, perhaps driven by wind streams during tropical storms. Subsequently, introgression of lineages may have occurred through hybridization, which can explain the discordance between the phenotype (like M. attenboroughi) and the genotype (like M. nyctor) of this specimen. Additional comments on the formation of the Caribbean Myotis assemblage are presented from the perspective of new evolutionary discoveries for this genus.

Key Words

Barbados, Caribbean biogeography, historical DNA, Myotis nyctor, Suriname, taxonomy

Introduction

Myotis has a worldwide distribution, occurring in every region of the globe except for parts of Australasia and the polar regions and is the most diverse genus within the order Chiroptera, comprising 139 valid species (Moratelli et al. 2019; Simmons and Cirranello 2024). Thirty-six neotropical species are currently recognized (Novaes et al. 2024), seven of which are found on the Caribbean islands (Novaes et al. 2021), being: M. nesopolus Miller, 1900 in Bonaire and Curaçao; M. dominicensis Miller, 1902 in Dominica and Guadeloupe; M. martiniquensis LaVal, 1973 in Martinique; M. nyctor LaVal & Schwartz, 1974 in Barbados and Grenada; M. pilosatibialis LaVal, 1973 (possibly M. armiensis Carrión-Bonilla & Cook, 2021), M. riparius Handley, 1960 in Trinidad; and M. attenboroughi Moratelli et al., 2017 in Tobago (LaVal 1973; Larsen et al. 2012; Moratelli et al. 2017; Novaes et al. 2021).

Miller and Allen (1928) recognized only M. nigricans (Schinz, 1821), with three subspecies in the Caribbean islands. Populations from Grenada and Trinidad and Tobago were assigned to the nominative subspecies, those from Dominica to M. nigricans dominicensis Miller, 1902; and those from Curaçao to M. nigricans nesopolus Miller, 1900. Later, LaVal (1973) elevated dominicensis to the species level and described M. martiniquensis from Martinique and Barbados. LaVal and Schwartz (1974) subsequently recognized Barbados’ populations as distinct and described a new subspecies, M. martiniquensis nyctor. Genoways and Williams (1979) elevated ne­sopolus to the species level and reported its occurrence in Bonaire, while Genoways et al. (1998) reported M. nigricans from Grenada. Masson and Breuil (1992) reported the occurrence of one Myotis species from Guadeloupe, presumably assigned to dominicensis. Larsen et al. (2012) raised M. martiniquensis nyctor to the species level, noting its presence in Barbados, with pending records from Grenada based on three museum specimens that were not examined by the authors. Moratelli et al. (2017) examined these specimens, assigning USNM 254717 to Peropteryx (Emballonuridae), and CM 83427 and USNM 252600 to M. nyctor. In addition, these authors revised the population of Myotis nigricans from Tobago and, based on morphological and molecular evidence, assigned this population to a new species, Myotis attenboroughi (see Moratelli et al. 2017). Recently, Novaes et al. (2021) reviewed the Venezuelan and Caribbean Myotis, suggesting that populations from Barbados and Grenada may represent distinct taxa, and emphasized the need for additional morphological and molecular data to support this hypothesis.

Based on novel molecular and morphological evidence, we extend the occurrence of M. attenboroughi to Grenada (based on the reexamination of the specimen USNM 252600, collected in 1938 and previously assigned to M. nyctor) and to mainland South America (based on the specimen CM 77705 previously assigned to Myotis cf. nigricans). Our reassessment of the taxonomic identity of these and other historical specimens from Tobago (i.e., USNM 540692) and Grenada (i.e., CM 83427) also allowed us to further discuss the evolution of Myotis in the Caribbean.

Material and methods

Our research group conducted a comprehensive review of neotropical Myotis by analyzing over 7,500 specimens and hundreds of DNA sequences. This extensive analysis resulted in the recognition of 11 new species and several other nomenclatural acts (e.g., Moratelli et al. 2011, 2013; Novaes et al. 2022a, b). The present study was based on the analysis of datasets comprising molecular, morphometric, and discrete morphological characters. We adopted the Phylogenetic Species Concept (Wheeler 1999), considering monophyly and diagnosability as criteria for species recognition (Gutiérrez and Garbino 2018).

Molecular data and analyses

Molecular analyses were based on 122 sequences of the mitochondrial cytochrome b gene (cytb, ca. 1,140 bp) from New World Myotis species and three outgroups (Appendix 1). Most sequences (123 out of 125, including outgroups) were obtained from NCBI’s GenBank, inclu­ding the sequences from CM 77705 (Suriname) and CM 83427 (Grenada). The remaining two sequences were generated in this study. Tissue samples of Myotis from Grenada (USNM 252600) and Tobago (USNM 540692) were obtained from toe clips from historical specimens deposited in the Smithsonian National Museum of Na­tural History, Washington D.C., USA, following rigorous sampling procedures described in Abreu et al. (2020). DNA extractions were performed in an isolated historical DNA facility at the Smithsonian Center for Conservation Genomics (CCG), using a standard phenol-chloroform protocol (McDonough et al. 2018), including a long (48–72 hours) lysis step. We did not perform specific amplifications for the cytb gene. Fragments of mtDNA were obtained as a byproduct (off-target sequences) of the capture and enrichment of Ultraconserved Elements sequenced for a parallel study conducted by our team. Sequencing was performed on Illumina Hi-Seq 4000 150 PE at the Vincent J. Coates Genomics Sequencing Laboratory at the University of California, Berkeley. A detailed description of the procedures used in the preparation of genomic libraries, DNA quantification, and sequencing is available in Abreu et al. (2020). To obtain the cytb gene sequences, we mapped the clean Illumina reads against a reference mitochondrial genome from GenBank, using the “Map to Reference” tool in Geneious R11 (Kearse et al. 2012). Mapping and sequence assembly parameters followed Abreu et al. (2020).

The cytb dataset was aligned using the UPGMA clustering method implemented in the MUSCLE algorithm (Edgar 2004) in the MEGA X software (Kumar et al. 2018) with default settings. The evolutionary model of nucleotide substitution was chosen for phylogenetic analyses using the software JModelTest 2 (Darriba et al. 2012), employing the Bayesian Information Criterion (BIC). The Hasegawa–Kishino–Yano model (Hasegawa et al. 1985) yielded the best fit to our dataset regarding the substitution of nucleotides, corrected for rate heterogeneity with gamma distribution and proportion of invariant sites parameters (i.e., HKY + Γ + I). Phylogenetic reconstruction was performed using the Bayesian Inference (BI) probabilistic method (Huelsenbeck et al. 2001) in the software MrBayes v. 3.4 (Ronquist and Huelsenbeck 2003) using the coupled Markov Chain Monte Carlo (MCMC). Four simultaneous Markov chains were performed for 100,000,000 generations with trees sampled every 10,000 generations. The first 26,000 trees were discarded as burn-in. Posterior probabilities were calculated from the consensus of the remaining trees. The confidence of the Bayesian sampling was verified for the free parameters using the effective sample size (ESS) statistic implemented in the software Tracer v. 1.5 (Rambaut and Drummond 2009). Convergence was checked by plotting log-likelihood va­lues against the generation time for each model, with all parameters showing ESS greater than 300 and asymptotically convergence indicating reliable performance.

Pairwise genetic distances within and among Myotis species were estimated using the HKY model implemented in ‘ape 5.0’ package for R software (Paradis and Schliep 2019), which measures the distance between pairs of sequences by estimating the proportion of different nucleotides between them.

Morphological data and analyses

For the morphological analyses and comparisons, we examined 14 specimens of M. attenboroughi (13 from Tobago, including holotype and paratypes; and one from Grenada); eight of M. nyctor from Barbados (including a paratypes); and one specimen from Grenada tentatively identified as M. cf. nyctor (Appendix 2). Quantitative morphological data were based on 16 skull dimensions, representing different axes of the length and width of the skull, rostrum, and mandible, and three external measurements (Table 1). Measurements were taken using digital calipers accurate to 0.01 mm, exclusively from individu­als classified as adults based on closed epiphyses (see Brunet-Rossini and Wilkinson 2009). Additionally, measurements of total length, tail length, hindfoot length and body weight were recorded from the specimen’s tag and used for comparative purposes only.

Table 1.
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Skull dimensions used to perform morphological comparisons in Neotropical Myotis species. Measurements were taken in millimeters.

Measurement Description
Forearm Length (FL) From the elbow to the distal end of the forearm including carpals
Third metacarpal length (3ML) From the distal end of the forearm including carpals to the distal end of the 3rd metacarpal.
Lenght of dorsal fur (LDF) Length of the longest hairs at the midpoint of the scapulae
Length of ventral fur (LVF) Length of the longest hairs at the midpoint of the sternum
Mandibular length (MAL) From the mandibular symphysis to the condyloid process
Mandibular toothrow length (MAN) From the lower canine to third molar
Greatest length of skull (GLS) From the apex of upper internal incisors to the occiput
Condylo-canine length (CCL) From the anterior surface of upper canines to a line connecting the occipital condyles
Condylo-incisive length (CIL) From the apex of upper internal incisors to a line connecting the occipital condyles
Condylo-basal length (CBL) From the anterior region of premaxilla to a line connecting the occipital condyles
Basal length (BAL) Least distance from the apex of upper internal incisors to the anterior margin of the foramen magnum
Zygomatic breadth (ZYG) Greatest breadth across the outer margins of the zygomatic arches
Mastoid breadth (MAB) Greatest breadth across the mastoid region
Braincase breadth (BCB) Greatest breadth of the globular part of the braincase
Interorbital breadth (IOB) Least breadth between the orbits.
Postorbital constriction (POB) Least breadth across frontals posterior to the postorbital bulges
Breadth across canines (BAC) Greatest breadth across outer edges of the crowns of upper canines including cingulae
Breadth across molars (BAM) Greatest breadth across outer edges of the crowns of upper molars
Maxillary toothrow length (MTL) From the upper canine to third molar
Upper molar toothrow length (M1M3) From M1 to M3

Principal Component Analysis (PCA) was used to identify general trends in variation in size and shape variation of the skull between forms of Barbados, Grenada, and Tobago. This analysis was performed in R using the MASS (Vanables and Ripley 2002) and Lattice packages (Sarkar 2008). As PCA requires a complete dataset without missing data, cranial measurements that could not be taken from the specimen due to skull fractures and losses of parts were estimated from the log-transformed dataset using the EM algorithm implemented in the R package Amelia II (Honaker et al. 2011).

Qualitative morphological analyses were based on six cranial and external characters traditionally used in Neotropical Myotis taxonomy (q.v., LaVal 1973; Moratelli et al. 2013; Novaes et al. 2022a). Fur color was also used to describe the variation, following the nomenclature used in the color catalog of Ridgway (1912).

Results

Phylogenetic inference and genetic distances

The specimen of Myotis from Grenada (USNM 252600) was recovered within the M. attenboroughi clade, which also includes the paratype USNM 540692 (Fig. 1). The specimen from Paramaribo, Suriname (CM 77705), was recovered as sister to all Caribbean samples (Tobago and Grenada) of M. attenboroughi. The specimen CM 83427 from Grenada was recovered within a clade composed by specimens of M. nyctor from Barbados, and here it is being treated as M. cf. nyctor. Both clades corresponding to M. attenboroughi and M. nyctor are closely related to other Myotis species from the Caribbean and northern South America, all included in the albescens species group (Fig. 1).

Figure 1.

Phylogenetic tree based on Bayesian Inference of cytochrome-b sequences of species of Neotropical Myotis. Nodal support was calculated by posterior probabilities. Caribbean species are highlighted by gray bars.

The estimated HKY genetic distances indicated that M. attenboroughi from Suriname (CM 77705) diverges by approximately 1.5% from samples from Tobago and Grenada. The average divergence within the clade composed of Tobago and Grenada samples of M. attenboroughi was less than 0.1%. Genetic distances among M. attenboroughi and phylogenetic closely species range from 6 to 8% (Table 2). Myotis attenboroughi from Grenada (USNM 252600) diverges by about 10% from M. nyctor from Barbados, alike M. attenboroughi from Grenada diverges in about 10% from M. cf. nyctor from Grenada (CM 83427).

Table 2.
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Average Hasegawa–Kishino–Yano genetic distances within (boldface along diagonal) and among (below diagonal) Myotis species based on cytochrome-b gene sequences.

Species 1 2 3 4 5 6 7
1. M. attenboroughi [0.008]
2. M. larensis 0.073 [0.006]
3. M. nesopolus 0.080 0.051 [0.002]
4. M. diminutus 0.069 0.020 0.044 [0.009]
5. M. caucensis 0.090 0.041 0.040 0.037 [0.031]
6. M. oxyotus 0.059 0.063 0.072 0.044 0.069 [0.015]
7. M. nyctor 0.111 0.132 0.145 0.116 0.137 0.124 [0.006]

Morphological description and comparisons

The specimen of M. attenboroughi from Grenada (USNM 252600) is an adult male preserved as dry skin and skull (Figs 2, 3). It is a small-sized specimen (forearm length 32.6 mm; other measurements in Table 3), with medium-sized ears (14 mm). The fur is silky in texture and medium-sized (LDH 5.6 mm, LVH 4.5 mm). The dorsal fur is bicolor, with dark brown bases (ca. 1/2 of the total length of the fur), and Mummy Brown tips (ca. 1/2), without a defined contrast between the bands. Ventral fur is strongly bicolor, with blackish bases and Light-Buff tips (Fig. 2). Membranes are Mummy Brown. Uropatagium is attached to the foot by a broad band of membrane. Dorsal surfaces of elbow and tibia are naked. The uropatagium lacks the fringe of hairs along the trailing edge.

Table 3.
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Selected measurements of Myotis attenboroughi and M. nyctor. Descriptive statistics include the mean, range (in parentheses), and sample size. Measurements are rounded to the nearest 0.1 mm. See Table 1 for abbreviations.

Measurements M. attenboroughi Grenada USNM 252600 M. attenboroughi Tobago M. cf. nyctor Grenada CM 83427 M. nyctor Barbados
FL 32.6 32.3 (31.4–33.3) 5 33.5 35.3 (34.6–35.9) 7
3ML 30.1 29.5 (28.5–30.3) 4 – 32.2 (31.4–32.7) 7
LDF 5.6 7.0 (6.0–8.0) 4 5.9 6.2–6.5 (2)
LVF 4.5 5.0 (5.0–6.0) 4 3.7 4.0–5.0 (2)
MAL 9.2 9.2 (8.9–9.5) 10 9.1 10.4 (10.2–10.6) 7
MAN 5.1 5.2 (5.1–5.3) 10 5.1 5.9 (5.4–6.0) 7
GLS – 12.9 (12.5–13.1) 10 13.2 14.3 (14.1–14.4) 7
CCL 11.6 11.5 (11.1–11.7) 10 11.7 12.8 (12.6–13.1) 7
CBL – 12.1 (11.8–12.4) 10 12.2 13.5 (13.3–13.7) 7
CIL – 12.3 (12.0–12.6) 10 12.4 13.8 (13.4–14.0) 7
BAL – 11.0 (10.8–11.4) 10 11.2 12.5 (12.1–12.7) 7
ZYG 8.2 7.9 (7.8–8.1) 6 8.3 (8.1–8.3) 2
MAB 6.7 6.5 (6.4–6.7) 10 6.7 7.2 (6.9–7.3) 7
BCB 6.2 6.0 (5.9–6.2) 10 6.2 6.6 (6.4–6.7) 7
IOB 4.2 4.2 (4.1–4.3) 10 4.0 4.3 (4.0–4.6) 7
POB 3.6 3.2 (3.2–3.5) 10 3.4 3.3 (3.3–3.4) 7
BAC 3.1 3.1 (2.9–3.3) 10 3.2 3.6 (3.6–3.8) 7
BAM 5.0 5.2 (5.1–5.3) 10 5.2 5.4 (5.4–5.5) 7
MTL 4.8 4.9 (4.8–5.0) 10 4.8 5.6 (5.4–5.7) 7
M1M3 2.7 2.8 (2.7–2.8) 10 2.7 3.0 (2.9–3.0) 7
Figure 2.

Dorsal (left) and ventral (right) view of the specimen of Myotis attenboroughi (USNM 252600) from Grenada Island.

Figure 3.

Skull profiles of Myotis attenboroughi (USNM 252600) from Grenada Island in dorsal, ventral, and lateral views.

Like in other neotropical Myotis, the dental formula is 2/3, 1/1, 3/3, 3/3 = 38. The skull is comparatively small, lacking a sagittal crest; lambdoidal crests are present, but very low; parietals are slightly inclined (Fig. 3). The occipital region is rounded, projecting behind the posterior surfaces of occipital condyles. The second upper premolar (P3) is aligned in the toothrow, smaller than P2 and P4, and visible in labial view. The set of qualitative morphological characters of the Grenada specimen (USNM 252600) is in accordance with the holotype of M. attenboroughi from Tobago (USNM 540693) and with the diagnosis presented in the original description (i.e., Moratelli et al. 2017). The morphometric measurements of this specimen are within the known range for M. attenboroughi from Tobago (including holotype and paratypes; Table 3).

Myotis attenboroughi and M. nyctor are phenotypically quite similar but they can be distinguished by a set of morphological characters (Figs 4, 5). Myotis nyctor has silky, medium-sized fur (LDH 6.3 mm, LVH 4.5 mm). Dorsal fur bicolored, with Cinnamon Brown to Mummy Brown tips (ca. 1/2 of the total length of the fur) and darker ba­ses, without well-marked limits between bands. Ventral fur strongly bicolored, with dark brown bases and Light-Buff tips. Myotis nyctor differs from M. attenboroughi in general size (Table 3) and craniodental characters, which include its longer and narrow rostrum; larger canines; less globose braincase; and narrower interorbital constriction.

Figure 4.

Skins of the paratype of Myotis nyctor (KU 109473) from Barbados (A, B) and holotype of Myotis attenboroughi (USNM 540693) from Tobago (C, D).

Figure 5.

Skulls of the paratype of Myotis nyctor (KU 109473) from Barbados, holotype of Myotis attenboroughi (USNM 540693) from Tobago, and a specimen of Myotis attenboroughi (USNM 252600) from Grenada.

The specimen from Grenada (CM 83427), phylogeneti­cally grouped with M. nyctor samples from Barbados, presented external and skull morphology virtually identical to M. attenboroughi specimen (USNM 252600) also from Grenada. The specimen CM 83427 is an adult male preserved as dry skin and skull. Despite exhibiting no discrete distinguishable character, these two individuals from Grenada (CM 83427 and USNM 252600) were also recovered closely positioned within the morphospace based on the PCA analysis. Both specimens presented skull dimensions similar to M. attenboroughi, being recovered close to the cluster formed by samples of M. attenboroughi from Tobago (Fig. 6). In this analysis, the first principal component (PC1) accounted for almost 98% of the variation, which was mainly driven by the variation found in measurements associated with condylo-incisive length, condylo-basal length, greatest length of skull, and condylo-canine length (Table 4).

Table 4.
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Vector correlation loadings with original variables of principal components analysis for Myotis attenboroughi and M. nyctor samples.

Measurements PC 1 PC 2
MAL 0.331 0.260
MAN 0.194 0.243
GLS 0.369 -0.298
CCL 0.366 0.037
CBL 0.386 -0.178
CIL 0.408 -0.063
BAL 0.388 -0.143
MAB 0.168 -0.061
BCB 0.123 -0.163
IOB 0.032 0.650
POB 0.011 -0.157
BAC 0.130 0.215
BAM 0.124 0.317
MTL 0.182 0.242
M1M3 0.064 0.202
Figure 6.

Dispersion points of the Principal Component Analysis based on skull measurements Myotis attenboroughi from Tobago (red dots), Myotis nyctor from Barbados (gray dots), the newly discovered M. attenboroughi from Grenada (red square; USNM 252600), and the specimen of M. cf. nyctor from Grenada (gray square; CM 83427).

Discussion

Our results unequivocally supported the identification of the specimen USNM 252600 from Grenada as M. attenboroughi. This specimen was captured by S. Gates in March 1938 and originally identified as M. nigricans. Genoways et al. (1998) examined this specimen and collected an additional individual during an expedition to Grenada in June 1986 (CM 83427), confirming their identity as M. nigricans and arguing that the specimens from Grenada are very similar to M. nigricans from mainland Venezuela. Later, Larsen et al. (2012) sequenced the cytb gene from the specimen CM 83427 and found this speci­men within a clade composed of M. nyctor from Barbados. However, Larsen et al. (2012) noted that cranial and external measurements of specimens from Grenada were outside the range observed for M. nyctor from Barbados, which corroborated the Genoways et al. (1998) observations. Based on qualitative morphological analyses, Moratelli et al. (2017) supported the results of Larsen et al. (2012) and reidentified the specimens USNM 252600 and CM 83427 as M. nyctor, thus rejecting the hypothesis of M. nigricans occurring in Grenada. In a critical review of Caribbean Myotis, Novaes et al. (2021) suggested that the taxonomic status of populations from Grenada still needed to be assessed, considering the presence of morphological disparities and possible retention of ancestral polymorphism resulting from recent speciation.

The present record of M. attenboroughi on the island of Grenada extends the geographic range of this species to two localities in the Caribbean islands (Grenada and Tobago) and one locality in the mainland South America (Suriname; Fig. 7). It is likely that this species has an even wider distribution in northern South America and may occur in the lowlands along coastal rainforests of the Guiana Shield and northeastern Venezuela, which corroborates the observations made by Genoways et al. (1998).

Figure 7.

Occurrences localities of Myotis attenboroughi in Caribbean islands of Grenada (1) and Tobago (2); and mainland South America in Paramaribo, Suriname (3).

The specimen CM 83427 morphologically matches with the diagnosis of M. attenboroughi (see Moratelli et al. 2017), being virtually identical to USNM 252600 from Grenada. However, our phylogenetic inference recovered this spe­cimen as belonging to the M. nyctor clade from Barbados, as previously shown by Larsen et al. (2012), revealing a surprising incongruence between genotype and phenotype. One could argue that the discrepancy between morpholo­gical and molecular analyses may be due to issues related with the tissue sampling for DNA analysis and/or with the lab routine for molecular data generation. Although this hypothesis cannot be completely ruled out without the re-sequencing these specimens, at least we can attest to the best practices while working with historical DNA for the individual USNM 252600. Sampling followed rigorous procedures described and tested in McDonough et al. (2018) and Abreu et al. (2020), and data generation (DNA extraction and genomic library preparation) took place in state-of-the-art facilities with protocols and equipment exclusively used for ancient and historical samples. Moreover, for specimen USNM 252600 we also generated thousands of nuclear ultraconserved elements (data not shown), and these data also corroborate the phylogenetic placement found here for this specimen. Therefore, if data generation issues were ruled out, this unexpected result should be explained in light of the complex evolutionary history of Caribbean Myotis.

The Caribbean Myotis assemblage originated from multiple overwater dispersals from northern South America to the Lesser Antilles in the Plio-Pleistocene interval (3.2–1.4 mya), which included posterior reverse colonization from the Caribbean to mainland Central and South Americas (Baker and Genoways 1978; Stadelmann et al. 2007; Larsen et al. 2012; Novaes et al. 2021). This pattern can also be observed in other insectivorous bats (e.g., Dávalos 2005, 2006; Genoways et al. 2005; Pavan et al. 2013). Dispersions may have been mediated by the migration arc formed by the expansion of land area of the Lesser Antilles islands due to sea level retreat during the Quaternary glaciations (Koopman 1958; Genoways et al. 2010; Dávalos and Russell 2012; Allen et al. 2019; Hoffman et al. 2019). This was especially important for Grenada, Trinidad, Tobago, and northern South America, which allowed a great exchange of species (Koopman 1958; Genoways et al. 2010). Thus, the presence of M. attenboroughi in Grenada, Tobago, and Suriname – and probably along a broader area in northern South America – can be explained by the intermittent connections between these islands and the continent during the Plio-Pleistocene glaciations. In fact, some studies consider the fauna of Grenada and Grenadines (united as a single island during the Last Glacial Maximum) as small representation of the South American bat fauna (e.g., Koopman 1958; Genoways et al. 2010; Pavan et al. 2013).

On the other hand, the presence of a Grenadian Myotis genotypically closer to M. nyctor from Barbados cannot be explained by this scenario. Barbados and Grenada are sepa­rated by ca. 240 km in the Tobago Basin, with an ocean depth of more than 2,000 m and no oceanic ridges that may have facilitated the connection of these two populations during glaciation periods (Speed 1981; Humphrey 1997). In addition, Barbados was completely or partially under­water until 1 mya, and perhaps as recent as 700,000 years ago (Speed and Keller 1993; Lovette et al. 1999). So, the arrival of the M. nyctor ancestral lineage in Barbados is quite recent and probably resulted from an extreme overwater dispersal event from South America (Larsen et al. 2012a). Therefore, it is possible to assume that other similar events, perhaps driven by wind streams during tropical storms (Hurme et al. 2025), may have facilitated the invasion of M. nyctor from Barbados into Grenada. Subsequently, introgression of lineages may have occurred through hybridization, which can explain the discordance between the phenotype (like M. attenboroughi) and the genotype (like M. nyctor) of the specimen CM 83427 (JN020562) from Grenada. Recent studies reveal a complex evolutionary histo­­ry of New World Myotis, which include several events of historical and modern introgression of lineages, hybridization, incomplete lineage sorting, and gene flow in phenotypically similar non-sister species (Morales and Carstens 2018; Platt et al. 2018; Korstian et al. 2022, 2024).

The hypotheses above are speculative, and new studies are necessary to understand the history of the Caribbean Myotis assemblage and the evolutionary processes linked to colonization and diversification. Genome-wide sequen­ces for the Barbados and Grenada specimens should advance our ability to answer these questions since the cytb-only dataset seems to have limited resolution power for the Caribbean clade, restricting data interpretation (Novaes et al. 2021). Furthermore, recent studies have shown discordance between the mitochondrial and nuclear genomes of Neotropical Myotis, resulting from the aforementioned evolutionary phenomena (Platt et al. 2018; Korstian et al. 2024). Still, there is robust evidence indicating that the Caribbean is an important diversification center for bats, and that the fauna of the Lesser Antilles was formed from multiple overwater dispersal from northern South America, with reverse colonization of species that invaded the continent after speciation (Hedges et al. 1992; Dávalos 2004, 2007; Pavan et al. 2013; Rojas et al. 2016; Tavares et al. 2018). In this way, the high richness of species, endemism, and presence on the different islands makes Myotis an important model for research into biogeography and evolution of the Caribbean.

Author contributions

RLMN, VCC, and RM conceptualized the study; RLMN, VCC, EFA, DEW, JEM, and RM collected and generated the data; RLMN and NAB analyzed the data; RLMN and VCC drafted the manuscript; all authors contributed to the theoretical conception of the study and contribute to the final version of the manuscript.

Data availability

All the data that support the findings of this study are available in the main text. DNA sequences generated in this study have been deposited in NCBI’s GenBank. Morphological data matrices for all specimens are available upon request from the corresponding author.

Acknowledgements

RLMN has received financial support from Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ, Brazil; E-26/204.243/2021; E26/200.631/2022 and E26/200.395/2022). RM received financial support from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil; 315184/2021-3) and FAPERJ (E-26/203.274/2017, E-26/210.254/2018, E-202.487/2018, E-26/200.967/2021, E-26/210.071/2023). EFA was supported by CNPq (147145/2016-3, 203692/2017-9, 165553/2017-0).

References

  • Abreu EF, Pavan SE, Tsuchiya MTN, Wilson DE, Percequillo AR, Maldonado JE (2020) Museomics of tree squirrels: a dense taxon sampling of mitogenomes reveals hidden diversity, phenotypic convergence, and the need of a taxonomic overhaul. BMC Evolutionary Biology 20: a77. https://doi.org/10.21203/rs.3.rs-28614/v1
  • Allen RW, Collier JS, Stewart AG, Henstock T, Goes S, Rietbrock A, the VoiLA Team (2019) The role of arc migration in the development of the Lesser Antilles: A new tectonic model for the Cenozoic evolution of the eastern Caribbean. Geology 47: 891–895. https://doi.org/10.1130/G46708.1
  • Baker RJ, Genoways HH (1978) Zoogeography of Antillean bats. In: Gill FG (Ed.) Zoogeography in the Caribbean, The 1975 Leidy Medal Symposium. Philadelphia Academy of Natural Science, Philadelphia, 1–128.
  • Brunet-Rossini AK, Wilkinson GS (2009) Methods for age estimation and study of senescence in bats. In: Kunz TH, Parsons S (Eds) Ecological and behavioral methods for the study of bats. The Johns Hopkins University Press, Baltimore, 315–325.
  • Caraballo DA, Montani ME, Martínez LM, Antoniazzi LR, Sambrana TC, Fernández C, Cisterna DM, Beltrán FJ, Colombo VC (2020) Hete­rogeneous taxonomic resolution of cytochrome b gene identification of bats from Argentina: Implications for field studies. PLoS One 15(12): e0244750. https://doi.org/10.1371/journal.pone.0244750
  • Carrión-Bonilla CA, Cook JA (2020) A new bat species of the genus Myotis with comments on the phylogenetic placement of M. keaysi and M. pilosatibialis. Therya 11: 508–532. https://doi.org/10.12933/therya-20-999
  • Carrión-Bonilla CA, Ron S, Cook JA (2024) Species richness in Myotis (Chiroptera: Vespertilionidae): species delimitation and expanded geographic sampling reveal high Neotropical diversity. Journal of Mammalogy 105: 241–258. https://doi.org/10.1093/jmammal/gyad124
  • Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: a772. https://doi.org/10.1038/nmeth.2109
  • Dávalos LM (2005) Molecular phylogeny of funnel–eared bats (Chiroptera: Natalidae). With notes on biogeography and conservation. Molecular Phylogenetics and Evolution 37: 1–103. https://doi.org/10.1016/j.ympev.2005.04.024
  • Genoways HH, Baker RJ, Bickham JW, Phillips CJ (2005) Bats of Jamaica. Special Publications, Museum of Texas Tech University 48: 1–154.
  • Genoways HH, Phillips CJ, Baker RJ (1998) Bats of the Antillean island of Grenada: a new zoogeographic perspective. Occasional Papers, Museum of Texas Tech University 177: 1–28. https://doi.org/10.5962/bhl.title.143306
  • Genoways HH, Kwiecinski GG, Larsen PA, Pedersen SC, Larsen RJ, Hoffman JD, Silva M, Phillips CJ, Baker RJ (2010) Bats of the Grenadine Islands, West Indies, and placement of Koopman’s line. Chiroptera Neotropical 16: 520–540.
  • Genoways HH, Williams SL (1979) Notes on bats (Mammalia: Chiroptera) from Bonaire and Curaçao, Dutch West Indies. Annals of the Carnegie Museum 48: 311–321. https://doi.org/10.5962/p.330831
  • Gutiérrez EE, Garbino GST (2018) Species delimitation based on diagnosis and monophyly, and its importance for advancing mammalian taxonomy. Zoological Research 39: 1–8.
  • Hasegawa M, Kishino H, Yano T (1985) Dating of human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution 22: 160–174. https://doi.org/10.1007/BF02101694
  • Hedges SB, Hass CA, Maxson LR (1992) Caribbean biogeography: molecular evidence for dispersal in West Indian terrestrial vertebrates. Proceedings of the National Academy of Sciences USA 89: 1909–1913. https://doi.org/10.1073/pnas.89.5.1909
  • Hoffman JD, Kadlubar GD, Pedersen SC, Larsen RJ, Larsen PA, Phillips CJ, Kwecinski GG, Genoways HH (2019) Predictors of bat species richness within the islands of the Caribbean basin. Special Publications, Museum of Texas Tech University 71: 337–377.
  • Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP (2001) Bayesian Inference of phylogeny and its impact on evolutionary biology. Science 294: 2310–2314.
  • Humphrey JD (1997) Geology and hydrogeology of Barbados. In: Vacher HL, Quinn T (Eds) Geology and hydrogeology of Carbonate islands: developments in sedimentology. Elsevier Science B.V., San Francisco, 381–406. https://doi.org/10.1016/S0070-4571(04)80033-5
  • Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrok S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T, Ashton B, Meintjes P, Drummond A (2012) Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1647−1649. https://doi.org/10.1093/bioinformatics/bts199
  • Koopman KF (1958) Land bridges and ecology in bat distribution on islands of the northern coast of South America. Evolution 12: 429–439. https://doi.org/10.2307/2405956
  • Korstian JM, Paulat NS, Platt RN, Stevens RD, Ray DA (2022) SINE-Based phylogenomics reveal extensive introgression and incomplete lineage sorting in Myotis. Genes 13: 339. https://doi.org/10.3390/genes13030399
  • Korstian JM, Stevens RD, Lee TE, Baker RJ, Ray DA (2024) Unraveling the Myotis morass: ultraconserved-element analysis reveals introgression, cryptic diversity, and taxonomic trouble. Journal of Mammalogy 105: 259–276. https://doi.org/10.1093/jmammal/gyad119
  • Kumar S, Stecher G, Li M, Knyaz C, Tamuda K (2018) MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms. Molecular Biology and Evolution 35: 1547−1549. https://doi.org/10.1093/molbev/msy096
  • Larsen RJ, Larsen PA, Genoways HH, Catzeflis FM, Geluso K, Kwiecinski KK, Pedersen SC, Simal F, Baker RJ (2012a) Evolutionary history of Caribbean species of Myotis, with evidence of a third Lesser Antillean endemic. Mammalian Biology 77: 124–134. https://doi.org/10.1016/j.mambio.2011.11.003
  • Larsen RJ, Knapp MC, Genoways HH, Khan FAA, Larsen PA, Wilson DE, Baker RJ (2012b) Genetic diversity of Neotropical Myotis (Chiroptera: Vespertilionidae) with an emphasis on South American species. PLoS One 7: e46578. https://doi.org/10.1371/journal.pone.0046578
  • LaVal RK (1973) A revision of the neotropical bats of the genus Myotis. Natural History Museum, Los Angeles County, Science Bulletin 15: 1–54.
  • LaVal RK, Schwartz A (1974) A new bat of the genus Myotis from Barbados. Caribbean Journal of Science 14: 189–192.
  • Lovette IJ, Seutin G, Ricklefs RE, Bermingham E (1999) The assembly of an island fauna by natural invasion: sources and temporal patterns in the avian colonization of Barbados. Biological Invasions 1: 33–41. https://doi.org/10.1023/A:1010090414598
  • Masson D, Breuil M (1992) Un Myotis (Chiroptera, Vespertilionidae) en Guadeloupe (Petites Antilles). Mammalia 56: 473–475.
  • McDonough MM, Parker LD, McInerney NR, Campana MG, Maldonado JE (2018) Performance of commonly requested destructive museum samples for mammalian genomic studies. Journal of Mammalogy 99: 789−802. https://doi.org/10.1093/jmammal/gyy080
  • Morales AE, Carstens BC (2018) Evidence that Myotis lucifugus “subspecies” are five nonsister species, despite gene flow. Systematic Biology 67: 756–769. https://doi.org/10.1093/sysbio/syy010
  • Moratelli R, Peracchi AL, Dias D, Oliveira JA (2011) Geographic variation in South American populations of Myotis nigricans (Schinz, 1821) (Chiroptera, Vespertilionidae), with the description of two new species. Mammalian Biology 76: 592−607. https://doi.org/10.1016/j.mambio.2011.01.003
  • Moratelli R, Gardner AL, Oliveira JA, Wilson DE (2013) Review of Myotis (Chiroptera, Vespertilionidae) from northern South America, including description of a new species. American Museum Novitates 3780: 1−36. https://doi.org/10.1206/3780.2
  • Moratelli R, Wilson DE, Novaes RLM, Helgen KW, Gutiérrez EE (2017) Caribbean Myotis (Chiroptera, Vespertilionidae), with description of a new species from Trinidad and Tobago. Journal of Mammalogy 98: 994–1008. https://doi.org/10.1093/jmammal/gyx062
  • Moratelli R, Burgin C, Cláudio VC, Novaes RLM, López-Baucells A, Haslauer R (2019) Family Vespertilionidae (Vesper Bats). In: Wilson DE, Mittermeier RA (Eds) Handbook of the Mammals of the World. Volume 9 – Bats. Lynx Editions, Barcelona, 716–981.
  • Novaes RLM, Cláudio VC, Larsen RJ, Wilson DE, Weksler M, Moratelli R (2021) The taxonomic status of Myotis nesopolus larensis (Chiroptera, Vespertilionidae) and new insights on the diversity of Caribbean Myotis. ZooKeys 1015: a145. https://doi.org/10.3897/zookeys.1015.59248
  • Novaes RLM, Cláudio VC, Carrión-Bonilla C, Abreu EF, Wilson DE, Maldonado JE, Weksler M (2022a) Variation in the Myotis keaysi complex (Chiroptera, Vespertilionidae), with description of a new species from Ecuador. Journal of Mammalogy 103: 540–559. https://doi.org/10.1093/jmammal/gyab139
  • Novaes RLM, Rodríguez-San Pedro A, Saldarriaga-Córdoba MM, Aguilera-Acuña O, Wilson DE, Moratelli R (2022c) Systematic review of Myotis (Chiroptera, Vespertilionidae) from Chile based on molecular, morphological, and bioacoustic data. Zootaxa 5188: 430–452. https://doi.org/10.11646/zootaxa.5188.5.2
  • Novaes RLM, Cláudio VC, Wilson DE, Weksler M, Moratelli R (2024) Taxonomic status of Myotis extremus (Chiroptera, Vespertilionidae) from Mesoamerica, with comments on distribution and taxonomy of Myotis nigricans. Zoologischer Anzeiger 308: 99–110. https://doi.org/10.1016/j.jcz.2023.11.009
  • Patrick LE, Stevens RD (2014) Investigating sensitivity of phylogenetic community structure metrics using North American desert bats. Journal of Mammalogy 95: 1240–1253. https://doi.org/10.1644/14-MAMM-A-007
  • Patterson BD, Webala PW, Peterhans JCK, Goodman SM, Bartonjo M, Demos TC (2019) Genetic variation and relationships among Afrotropical species of Myotis (Chiroptera: Vespertilionidae). Journal of Mammalogy 100: 1130–1143. https://doi.org/10.1093/jmammal/gyz087
  • Pavan AC, Martins FM, Morgante JS (2013) Evolutionary history of bulldog bats (genus Noctilio): recent diversification and the role of the Caribbean in Neotropical biogeography. Biological Journal of the Linnean Society 108: 210–224. https://doi.org/10.1111/j.1095-8312.2012.01979.x
  • Platt RN II, Faircloth BC, Sullivan KAM, Kieran TJ, Glenn TC, Vandewege MW, Lee TE, Baker RJ, Stevens RD, Ray DA (2018) Conflicting evolutionary histories of the mitochondrial and nuclear genomes in New World Myotis bats. Systematic Biology 67: 236–249. https://doi.org/10.1093/sysbio/syx070
  • Rojas D, Warsi OM, Dávalos LM (2016) Bats (Chiroptera: Noctilionoidea) challenge a recent origin of extant Neotropical diversity. Systematic Biology 65: 432–448. https://doi.org/10.1093/sysbio/syw011
  • Ruedi M, Mayer F (2001) Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggests deterministic ecomorphological convergences. Molecular Phylogenetics and Evolution 21: 463−448. https://doi.org/10.1006/mpev.2001.1017
  • Ruedi M, Csorba G, Liang-Kong L, Cheng-Han C (2015) Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920: 301–342. https://doi.org/10.11646/zootaxa.3920.2.6
  • Sedlock JL, Heaney LR, Balete DS, Ruedi M (2020) Philippine bats of the genus Kerivoula (Chiroptera: Vespertilionidae): overview and assessment of variation in K. pellucida and K. whiteheadi. Zootaxa 4755: 454–490. https://doi.org/10.11646/zootaxa.4755.3.2
  • Simmons NB, Cirranello AL (2024) Bat species of the world: a taxonomic and geographic database. Version 1.6. https://batnames.org/home.html [Accessed 30 September 2024]
  • Speed RC (1981) Geology of Barbados: implications for an accretionary origin. Acta Oceanologica 81: 259–265.
  • Speed RC, Keller CA (1993) Synopsis of the geological evolution of Barbados. Journal of the Barbados Museum of History and Society 41: 113–139.
  • Stadelmann B, Lin LK, Kunz TH, Ruedi M (2007) Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes. Molecular Phylogenetics and Evolution 43: 32–48. https://doi.org/10.1016/j.ympev.2006.06.019
  • Tavares VC, Warsi OM, Balseiro F, Mancina CA, Dávalos LM (2018) Out of the Antilles: Fossil phylogenies support reverse colonization of bats to South America. Journal of Biogeography 45: 859–873. https://doi.org/10.1111/jbi.13175
  • Wheeler QD (1999) Why the phylogenetic species concept? – Elementary. Journal of Nematology 31: 134–141.

Appendix 1

Table A1.
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Cytochrome b sequences used in phylogenetic analyses. The information presented for the taxonomic terminals is the result of re-identification of the specimens and does not necessarily coincide with the original identifications provided by the authors and GenBank. Abbreviations for specimen deposit institutions are: Universidad Autónoma Metropolitana-Iztapalapa (UAMI, Ciudad de México, Mexico); Pontificia Universidad Católica del Ecuador (QCAZ, Quito, Ecuador), Carnegie Museum of Natural History (CM, Pittsburg, USA); Field Museum of Natural History (FMNH, Chicago, USA), Museum of Southwestern Biology, University of New Mexico (MSB, Albuquerque, USA), Museum of Vertebrate Zoology, University of California (MVZ, Berkeley, USA), University of Nebraska State Museum (UNSM-ZM, Lincoln, USA); Sam Noble Oklahoma Museum of Natural History (OMNH, Norman, USA); Smithsonian’s National Museum of Natural History (USNM, Washington, DC, USA); Texas Tech University (TTU, Lubbock, USA), Biology Department of Tunghai University (THUMB, Taichung, Taiwan). *The cytb sequence of M. attenboroughi from Grenada (USNM 252600) is available as Suppl. material 1.

Species #GenBank Voucher Locality Source
Myotis albescens JX130463 TTU 85088 Pastaza, Ecuador Larsen et al. (2012b)
Myotis albescens JX130522 TTU 85091 Pastaza, Ecuador Larsen et al. (2012b)
Myotis albescens AF376839 FMNH 162543 Tarija, Bolivia Ruedi and Mayer (2001)
Myotis albescens JX130503 TTU 99124 Boquerón, Paraguay Larsen et al. (2012b)
Myotis albescens JX130504 TTU 99818 Ñeembucú, Paraguay Larsen et al. (2012b)
Myotis arescens OP270161 – Araucanía, Chile Novaes et al. (2022c)
Myotis arescens OP270162 – Araucanía, Chile Novaes et al. (2022c)
Myotis arescens OP270166 – Coquimbo, Chile Novaes et al. (2022c)
Myotis arescens AM261888 – Santiago, Chile Stadelmann et al. (2007)
Myotis attenboroughi JN020573 UNSM-ZM 29470 St. George Parish, Tobago Larsen et al. (2012a)
Myotis attenboroughi JN020574 UNSM-ZM 29483 St. George Parish, Tobago Larsen et al. (2012a)
Myotis attenboroughi PQ757917 USNM 540692 St. George Parish, Tobago Present study
Myotis attenboroughi* – USNM 252600 St. David, Grenada Island Present study
Myotis attenboroughi JX130505 CM 77705 Paramaribo, Suriname Larsen et al. (2012b)
Myotis armiensis JX130435 TTU 39146 Chiriquí, Panama Larsen et al. (2012b)
Myotis armiensis MW025265 MSB 262089 Chiriquí, Panama Carrión-Bonilla and Cook (2020)
Myotis armiensis MW025266 MSB 262237 Chiriquí, Panamá Carrión-Bonilla and Cook (2020)
Myotis armiensis MW025267 MSB 262788 Chiriquí, Panamá Carrión-Bonilla and Cook (2020)
Myotis armiensis MW025268 MSB 262085 Chiriquí, Panamá Carrión-Bonilla and Cook (2020)
Myotis armiensis JX130514 TTU 85060 Tungurahua, Ecuador Larsen et al. (2012b)
Myotis armiensis MW025269 QCAZ 17245 Napo, Ecuador Carrión-Bonilla and Cook (2020)
Myotis armiensis MW025274 QCAZ 12461 Zamora Chinchipe, Ecuador Carrión-Bonilla and Cook (2020)
Myotis armiensis MZ345121 USNM 370890 Distrito Federal, Venezuela Novaes et al. (2022a)
Myotis atacamensis OP270158 – Arica, Chile Novaes et al. (2022c)
Myotis atacamensis OP270159 – Arica, Chile Novaes et al. (2022c)
Myotis atacamensis OP270160 – Arica, Chile Novaes et al. (2022c)
Myotis bakeri AM261882 MVZ 168933 Olmos, Peru Stadelmann et al. (2007)
Myotis caucensis JX130484 CM 98860 Huánuco, Peru Larsen et al. (2012b)
Myotis caucensis JX130538 TTU 46346 Huánuco, Peru Larsen et al. (2012b)
Myotis causensis JX130495 QCAZ 6313 Pastaza, Ecuador Larsen et al. (2012b)
Myotis chiloensis OP270163 – Los Lagos, Chile Novaes et al. (2022c)
Myotis chiloensis OP270164 – Los Lagos, Chile Novaes et al. (2022c)
Myotis chiloensis OP270165 – Los Lagos, Chile Novaes et al. (2022c)
Myotis clydejonesi JX130520 TTU 109227 Sipaliwini, Suriname Larsen et al. (2012b)
Myotis clydejonesi JX130453 CM 98859 Huánuco, Peru Larsen et al. (2012b)
Myotis diminutus JX130447 TTU 103805 Loja, Ecuador Larsen et al. (2012b)
Myotis diminutus JX130448 QCAZ 9601 Esmeraldas, Ecuador Larsen et al. (2012b)
Myotis diminutus JX130466 QCAZ 9154 Esmeraldas, Ecuador Larsen et al. (2012b)
Myotis diminutus JX130467 QCAZ 9155 Esmeraldas, Ecuador Larsen et al. (2012b)
Myotis dinellii JX130475 TTU 66489 Córdoba, Argentina Larsen et al. (2012b)
Myotis dinellii MT262853 MG-ZV-M 217 Zavalla, Argentina Caraballo et al. (2020)
Myotis dinellii MT262857 MG-ZV-M 233 Chanar Laneado, Argentina Caraballo et al. (2020)
Myotis dominicensis JN020555 TTU 31507 St. Joseph’s Parish, Dominica Larsen et al. (2012a)
Myotis dominicensis JN020556 TTU 31508 St. Joseph’s Parish, Dominica Larsen et al. (2012a)
Myotis dominicensis AF376848 – St. Joseph’s Parish, Dominica Ruedi and Mayer (2001)
Myotis elegans JX130479 TTU 84380 Atlantida, Honduras Larsen et al. (2012b)
Myotis elegans JX130480 TTU 84138 Atlantida, Honduras Larsen et al. (2012b)
Myotis extremus AF376852 – Yucatán, Mexico Ruedi and Mayer (2001)
Myotis extremus JX130449 TTU 47514 Yucatán, Mexico Larsen et al. (2012b)
Myotis extremus JX130525 – Yucatán, Mexico Larsen et al. (2012b)
Myotis extremus JX130489 CM 55764 Veracruz, Mexico Larsen et al. (2012b)
Myotis extremus MF143477 – Veracruz, Mexico Platt et al. (2018)
Myotis extremus MW025270 MVZ 226977 Alta Verapaz, Guatemala Carrión-Bonilla and Cook (2020)
Myotis fortidens JX130437 – Michoacán, Mexico Larsen et al. (2012b)
Myotis fortidens JX130439 – Michoacán, Mexico Larsen et al. (2012b)
Myotis fortidens KC747690 LACM 73713 Guerrero, Mexico Patrick and Stevens (2014)
Myotis keaysi JX130516 QCAZ 11380 Chimborazo, Ecuador Larsen et al. (2012b)
Myotis keaysi JX130517 QCAZ 11383 Chimborazo, Ecuador Larsen et al. (2012b)
Myotis keaysi MW025273 MSB 70381 Cochabamba, Bolivia Carrión-Bonilla and Cook (2020)
Myotis larensis JN020569 TTU 48161 Guárico, Venezuela Larsen et al. (2012b)
Myotis larensis JX130529 TTU 48162 Guárico, Venezuela Larsen et al. (2012b)
Myotis larensis JX130535 CM 78645 Guárico, Venezuela Larsen et al. (2012b)
Myotis lavali AF376864 MVZ 185681 Paraíba, Brazil Ruedi and Mayer (2001)
Myotis martiniquensis AM262332 – Martinique Island Stadelmann et al. (2007)
Myotis martiniquensis JN020557 MNHN 2005-895 GranďRivière, Martinique Larsen et al. (2012a)
Myotis martiniquensis JN020558 MNHN 2005-896 Le Morne Rouge, Martinique Larsen et al. (2012a)
Myotis midastactus MW323450 USNM 584502 Santa Cruz, Bolivia Novaes et al. (2022a)
Myotis moratellii JX130572 QCAZ 9179 El Oro, Ecuador Larsen et al. (2012b)
Myotis moratellii MZ345120 USNM 513482 Los Ríos, Ecuador Novaes et al. (2022a)
Myotis nesopolus JN020575 – Bonaire, Netherlands Antilles Larsen et al. (2012a)
Myotis nesopolus JN020576 – Bonaire, Netherlands Antilles Larsen et al. (2012a)
Myotis nesopolus JN020577 – Bonaire, Netherlands Antilles Larsen et al. (2012a)
Myotis nigricans OR187561 FMA 630 Rio de Janeiro, Brazil Novaes et al. (2024)
Myotis nigricans OR187562 FMA 957 Rio de Janeiro, Brazil Novaes et al. (2024)
Myotis nigricans PP584498 FMA 1525 Rio de Janeiro, Brazil Novaes et al. (2024)
Myotis nigricans PP584499 FMA 1534 Rio de Janeiro, Brazil Novaes et al. (2024)
Myotis cf. nyctor JN020562 CM 83427 St. David Parish, Grenada Larsen et al. (2012a)
Myotis nyctor JN020563 TTU 109225 St. Thomas Parish, Barbados Larsen et al. (2012a)
Myotis nyctor JN020564 TTU 109226 St. Thomas Parish, Barbados Larsen et al. (2012a)
Myotis nyctor JN020565 TTU 109229 St. Thomas Parish, Barbados Larsen et al. (2012a)
Myotis nyctor JN020566 TTU 109224 St. Thomas Parish, Barbados Larsen et al. (2012a)
Myotis nyctor JN020567 TTU 109230 St. Thomas Parish, Barbados Larsen et al. (2012a)
Myotis oxyotus JX130509 – Loja, Ecuador Larsen et al. (2012b)
Myotis oxyotus JX130585 – Loja, Ecuador Larsen et al. (2012b)
Myotis oxyotus MW089499 QCAZ 11739 Imbabura, Ecuador Carrión-Bonilla et al. (2024)
Myotis pilosatibialis JX130526 TTU 35360 San Luis Potosí, Mexico Larsen et al. (2012b)
Myotis pilosatibialis JX130518 TTU 35631 San Luis Potosí, Mexico Larsen et al. (2012b)
Myotis pilosatibialis MW025271 MVZ 226976 Alta Verapaz, Guatemala Carrión-Bonilla and Cook (2020)
Myotis pilosatibialis MW025272 MVZ 226973 El Quiche, Guatemala Carrión-Bonilla and Cook (2020)
Myotis pilosatibialis MW025275 MVZ 224798 Quezaltenango, Guatemala Carrión-Bonilla and Cook (2020)
Myotis pilosatibialis JX130519 TTU 60981 Santa Ana, El Salvador Larsen et al. (2012b)
Myotis riparius JX130492 TTU 102883 Esmeraldas, Ecuador Larsen et al. (2012b)
Myotis riparius JX130473 CM 68443 Para, Suriname Larsen et al. (2012b)
Myotis riparius JX130474 CM 78659 Bolívar, Venezuela Larsen et al. (2012b)
Myotis riparius MW089495 MSB 70383 Cochabamba, Bolivia Carrión-Bonilla et al. (2024)
Myotis riparius JX130485 TTU 99645 Paraguarí, Paraguay Larsen et al. (2012b)
Myotis riparius MW089493 OMNH 36220 Tucumán, Argentina Carrión-Bonilla et al. (2024)
Myotis ruber AF376867 MVZ 185999 São Paulo, Brazil Ruedi and Mayer (2001)
Myotis simus JX130481 TTU 46348 Huánuco, Peru Larsen et al. (2012b)
Myotis velifer AF376870 MVZ 146766 Sonora, Mexico Ruedi and Mayer (2001)
Myotis velifer JX130438 UAMI 15306 Michoacán, Mexico Larsen et al. (2012b)
Myotis velifer JX130589 UAMI 15305 Michoacán, Mexico Larsen et al. (2012b)
Myotis yumanensis AF376875 MVZ 15585 California, USA Stadelmann et al. (2007)
Myotis sp. 1 JN020570 CM 63933 Nickerie, Suriname Larsen et al. (2012a)
Myotis sp. 1 JN020571 CM 69053 Para, Suriname Larsen et al. (2012a)
Myotis sp. 1 JN020572 CM 77699 Para, Suriname Larsen et al. (2012a)
Myotis sp. 1 JX130476 CM 77692 Marowjine, Suriname Larsen et al. (2012b)
Myotis sp. 1 JX130534 CM 77694 Nickerie, Suriname Larsen et al. (2012b)
Myotis sp. 1 JX130536 CM 77700 Para, Suriname Larsen et al. (2012b)
Myotis sp. 2 AF376865 FMNH 129208 Lima, Peru Ruedi and Mayer (2001)
Myotis sp. 3 MT262866 MFA-ZV 1425 Esperanza, Argentina Caraballo et al. (2020)
Myotis sp. 3 JX130450 TTU 34952 Puerto Linares, Bolívia Larsen et al. (2012b)
Myotis sp. 3 JX130528 TTU 34953 Puerto Linares, Bolívia Larsen et al. (2012b)
Myotis sp. 3 PP584500 UFMT 4946 Mato Grosso, Brazil Novaes et al. (in press)
Myotis sp. 3 PP584501 MZUFV 5180 Mato Grosso, Brazil Novaes et al. (in press)
Myotis sp. 3 JX130498 TTU 99046 Alto Paraguai, Paraguay Larsen et al. (2012b)
Myotis sp. 3 JX130455 TTU 95992 Alto Paraguai, Paraguay Larsen et al. (2012b)
Myotis sp. 3 JX130540 TTU 99151 Boquerón, Paraguay Larsen et al. (2012b)
Myotis sp. 3 JX130539 TTU 99516 Concepción, Paraguay Larsen et al. (2012b)
Myotis sp. 3 JX130499 TTU 99802 Ñeembucu, Paraguay Larsen et al. (2012b)
Myotis sp. 3 JX130496 TTU 99743 Presidente Hayes, Paraguay Larsen et al. (2012b)
Outgroups
Myotis emarginatus MK799667 FMNH 178892 Ajlun, Jordan Patterson et al. (2019)
Submyotodon latirostris KP187906 THUMB 30036 Heping, Taiwan Ruedi et al. (2015)
Kerivoula papillosa MG194454 FMNH 205343 Luzon I, Philippine Island Sedlock et al. (2020)

Appendix 2

Specimens examined in morphological comparisons. These vouchers consist of fluid preserved specimens, stuffed skins, and skulls deposited in the American Museum of Natural History (AMNH, New York, United States); Carnegie Museum of Natural History (CM, Pittsburgh, United States); National Museum of Natural History, Smithsonian Institution (USNM, Washington, D.C., United States); Natural History Museum, University of Kansas (KU, Lawrence, United States).

Myotis attenboroughi (N = 14): Trinidad and Tobago: Tobago Island, Charlottesville, 1 km N of Pirate’s Bay, Saint John Parish (USNM 540693 [holotype], 540692 [paratype]); Tobago Island, St. Mary Parish, Hillsborough Reservoir (USNM 538064, 538065, 538066, 538067, 538068, 538069, 540619, 540620, 540621, 540694, 540695 [paratypes]). Grenada: St. David (USNM 252600).

Myotis nyctor (N = 8): Barbados: St. Thomas Parish, Cole’s cave (KU 151761, 151762, 151763, 151764, 151765, 151766, 109473 [paratype]); St. Thomas, Near Cole’s cave (AMNH 213926).

Myotis cf. nyctor (N = 1): Grenada, St. David (CM 83427).

Supplementary material

Supplementary material 1 

Supplementary data

Roberto Leonan M. Novaes, Vinícius C. Cláudio, Natasha A. Bertocchi, Edson F. Abreu, Don E. Wilson, Jesús E. Maldonado, Ricardo Moratelli

Data type: fasta

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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