On the variation, distribution and evolutionary history of Myotis armiensis (Chiroptera, Vespertilionidae) along Neotropical mountain ranges, including the description of new subspecies

On the variation, distribution and evolutionary history of Myotis armiensis (Chiroptera, Vespertilionidae) along Neotropical mountain ranges, including the description of new subspecies | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article On the variation, distribution and evolutionary history of Myotis armiensis (Chiroptera, Vespertilionidae) along Neotropical mountain ranges, including the description of new subspecies Roberto Leonan Morim Novaes, Marco Antônio Guimarães-Silva, Aldo Caccavo, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7794885/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 10 Feb, 2026 Read the published version in Mammalian Biology → Version 1 posted 4 You are reading this latest preprint version Abstract Myotis armiensis was recently described from specimens collected in the mountain ranges of Costa Rica, Panama, and Ecuador. During a systematic revision of Neotropical Myotis , we identified additional specimens from Venezuela and Colombia. Using an integrative approach that combined morphological traits, cytochrome b sequences, and ecological niche modeling, we assessed the geographic variation of M. armiensis . Our findings indicate that this species is structured into two allopatric populations: one in the Talamanca Cordillera of Central America, and another restricted to the South American Andes. These populations are reciprocally monophyletic, exhibiting genetic distances greater than 3% and notable morphometric divergence. The results suggest that M. armiensis comprises two incompletely isolated lineages, supporting the recognition of two subspecies described herein. The diversification of this taxon likely occurred during the Pleistocene, possibly driven by environmental fluctuations associated with glacial cycles. Andes Mountain range biogeography Myotinae Pleistocene speciation taxonomy. Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction Species are cornerstones of current evolutionary theory (Hull 1994 ; Ghiselin 1997 ; Claridge 2010 ) and several concepts and approaches to species delimitation have been proposed (see review in de Queiroz 2007 ). However, the boundary between intra- and interspecific variation remains one of the greatest challenges of modern taxonomy, especially in cryptic taxa, where phenotypic variation remains undetectable even after speciation events (Padial et al. 2010 ; Novaes et al. 2025a ). Neotropics harbors the richest biodiversity in the world, and several studies have shown that its species diversification is linked to complex interactions between geological and climatic history in the Americas, resulting in heterogeneous habitats (Antonelli 2021 ; Meseguer et al. 2022 ). In this context, the Mountain ranges, such as the Andes, represent an important biogeographic feature in South America, influencing rivers, precipitation, biome limits, and species distributions (Patterson et al. 2012 ; Viale and Garreaud 2015 ). The role of the Central America Cordilleras and Andes Mountain range in limiting gene flow and promoting species diversification has been widely documented for many vertebrate taxa, including amphibians, birds, and mammals (Brumfield and Capparella 1996 ; Patterson et al. 2012 ; Hutter et al. 2017 ; Solís et al. 2024 ). The complex and relatively recent Andean uplift, for example, promoted diversification in multiple taxa, but also the subdivision of relatively young species that rapidly accumulated high genetic differentiation while showing little morphological divergence (i.e., cryptic species formation), as seen in some bats of the genus Myotis Kaup, 1829 (Novaes et al. 2024a , 2025a ). Myotis is widely distributed worldwide and represents the most diverse bat genus, with approximately 140 valid species (Moratelli et al. 2019 ; Simmons and Cirranello 2025 a). Recent systematic reviews have revealed unexpectedly high diversity in Neotropical Myotis (Moratelli et al. 2011 ; Carrión-Bonilla et al. 2024 ; Novaes et al. 2024b ). Currently, 43 species of Myotis are recognized in the Neotropics, about 35% of which were described after 2010 (Carrión-Bonilla et al. 2024 ; Novaes et al. 2022a , 2025a ). Many of these recently described species still have poorly defined distributional limits, including Myotis armiensis Carrión-Bonilla & Cook, 2020 . Myotis armiensis was described based on molecular and morphological data from specimens collected in Costa Rica, Panama, and Ecuador (Carrión-Bonilla and Cook 2020 ). However, there are still gaps in its known distribution, likely due to phenotypic similarity with other species, particularly Myotis keaysi J.A. Allen, 1914 and Myotis pilosatibialis LaVal, 1973, which may complicate identification (Carrión-Bonilla and Cook 2020 ). As part of an ongoing review of Neotropical Myotis , we identified individuals of M. armiensis from Venezuela and Colombia, thereby expanding its known distribution, and obtained additional samples from Costa Rica, Panama, and Ecuador. We evaluated the intraspecific morphological and molecular variation of M. armiensis and assessed its distinction from M. pilosatibialis . In addition, we estimated the species’ potential distribution using ecological niche modeling. Because the signature of vicariance caused by the uplift of the Andes is evident in numerous mammalian taxa, particularly in many bat lineages (see Patterson et al. 2012 ), we tested the hypothesis that trans-Andean and cis-Andean populations of M. armiensis may represent different evolutionary lineages. Methods Molecular data and analyses Molecular analyses were based on 96 cytochrome b (cyt-b, ~ 1,140 bp) sequences from 29 species of Neotropical Myotis , along with two sequences from other Myotinae bats ( Myotis emarginatus and Submyotodon latirostris ) and Kerivoula papillosa (Temminck, 1840), which was used as outgroup (Appendix 1). All sequences were retrieved from NCBI GenBank, including those previously generated by us. The sequence of M. armiensis from Venezuela (USNM 370890; GenBank Accession MZ345121) was obtained from historical DNA extracted from a museum specimen, with details provided in Novaes et al. ( 2022b ). The dataset of 99 cyt-b sequences was aligned using the MUSCLE algorithm (Edgar 2004 ) with default parameters, as implemented in MEGA X (Kumar et al. 2018 ). The best-fitting model of nucleotide substitution for phylogenetic analyses was selected in JModelTest 2 (Darriba et al. 2012 ) under the Bayesian Information Criterion (BIC). The Hasegawa–Kishino–Yano model (HKY; Hasegawa et al. 1985 ), corrected for among-site rate heterogeneity with a gamma distribution and a proportion of invariant sites (HKY + Γ + I), was identified as the best fit. Phylogenetic reconstruction was performed using Bayesian inference (BI) in MrBayes 3.2 (Ronquist et al. 2012 ) with a coupled Markov chain Monte Carlo (MCMC) approach. Four simultaneous Markov chains were run for 100,000,000 generations, sampling every 10,000 generations. The first 26,000 trees were discarded as burn-in, and posterior probabilities were calculated from the consensus of the remaining trees. The robustness of Bayesian sampling was assessed by examining the effective sample size (ESS) of free parameters in Tracer v1.5 (Rambaut and Drummond 2009 ). All parameters showed ESS values greater than 300, and convergence was confirmed by plotting log-likelihood values against generation time, indicating reliable performance. Genetic distances were estimated under the Kimura two-parameter (K2P) model (Kimura 1980 ), as implemented in MEGA X. This method calculates pairwise distances by estimating the proportion of nucleotide differences between sequences. Morphological data and analyses For this study, we examined 29 specimens of Myotis armiensis and 40 specimens of M. pilosatibialis deposited in the collections of the Royal Ontario Museum (ROM, Toronto, Canada), American Museum of Natural History (AMNH, New York, USA), Carnegie Museum of Natural History (CM, Pittsburgh, USA), University of Kansas (KU, Lawrence, USA), Natural History Museum of Los Angeles County (LACM, Los Angeles, USA), Field Museum of Natural History (FMNH, Chicago, USA), Museum of Texas Tech University (TTU, Lubbock, USA), and the Smithsonian National Museum of Natural History (USNM, Washington, DC, USA) (Appendix 2). Specimens were identified based on morphological characters described by Carrión-Bonilla and Cook ( 2020 ) and by comparison with type-series material. Quantitative morphological analyses were based on five external and 16 craniodental measurements (Table 1 ). To minimize potential noise from ontogenetic variation, only adult individuals were included. Samples were organized into Operational Taxonomic Units (OTUs) following the results of phylogenetic studies on M. armiensis (e.g., Carrión-Bonilla and Cook 2020 ; Carrión-Bonilla et al. 2024 ), which identified two clades: one composed exclusively of Central American specimens and the other of South American specimens (Fig. 1 ). Samples of M. pilosatibialis were divided into OTUs corresponding to the smallest possible geographic groups that showed morphological cohesion. Table 1 External and craniodental measurements obtained from Myotis specimens. Measurement and acronym Description Forearm length (FA) From the elbow to the distal end of the forearm including carpals Third metacarpal length (3MC) From the proximal to the distal extremity of the third metacarpal Ear length (Ear) From the base to the apex of the ear Length of dorsal fur (LDF) Measured at the midpoint of the scapulae Length of ventral fur (LVF) Measured at the midpoint of the sternum Greatest length of skull (GLS) From the apex of the upper internal incisors, to the occiput Condylo-canine length (CCL) From the anterior surface of the upper canines to the occipital condyles limit Condylo-basal length (CBL) From the premaxillae to a line connecting the occipital condyles Condylo-incisive length (CIL) From the apex of upper internal incisors to the occipital condyles limit Basal length (BAL) Least distance from the apex of upper internal incisors to the ventral margin of the foramen magnum Zygomatic breadth (ZYG) Zygomatic breadth Mastoid breadth (MAB) Greatest breadth across the mastoid region Braincase breadth (BCB) Greatest breadth of the globular part of the braincase Postorbital breadth (POB) Least breadth across frontals posterior to the postorbital bulges Interorbital breadth (IOB) Least breadth between the orbits 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 M3 Molariform toothrow length (M1–3) From M1 to M3 Mandibular length (MAL) From the mandibular symphysis to the condyloid process Mandibular toothrow length (MAN) From the lower canine to m3 We used three multivariate approaches to investigate morphometric differences between the Central and South American clades of M. armiensis : (i) a multivariate analysis of variance (MANOVA), (ii) a principal component analysis (PCA) based on the variance–covariance matrix of log-transformed measurements, and (iii) a canonical variate analysis (CVA) based on the variance–covariance matrix of untransformed data. All analyses were performed in R v.4.0 using the packages stats, MASS, and lattice. Distribution patterns of measurements were visualized using violin plots and boxplots in PAST v.4.07b (Hammer et al. 2001 ). Both graphical assessments and MANOVA tests were performed exclusively on M. armiensis samples. For PCA and CVA, we also included M. pilosatibialis , a closely related and morphologically similar species, organized into three geographic groups: “Mexico,” “Guatemala,” and “Honduras + El Salvador.” Including these groups helped avoid spurious segregation patterns that might arise when comparing only two populations, especially in CVA. Slightly damaged skulls with up to two missing measurements were retained, with missing values estimated using an expectation–maximization with bootstrapping (EMB) algorithm, as implemented in the Amelia II package (Honaker et al. 2011 ) in R. The statistical significance of principal components was evaluated with the Broken Stick (BS) test (Frontier 1976 ; Peres-Neto et al. 2005 ). To reduce the risk of overfitting in CVA due to a low sample-size-to-variable ratio (Kovarovic et al. 2011 ; Mitteroecker and Bookstein 2011 ), we restricted the analysis to a subset of 10 variables (MAN, GLS, CIL, MAB, BCB, IOB, POB, BAC, BAM, and M1M3) representing overall skull dimensions, excluding measurements that showed high correlations (r > 0.8) with one or more of the selected variables. We report eigenvalues, eigenvectors, explained variance for both principal components (PCs) and canonical variates (CVs), as well as scatterplots of PC and CV scores. Qualitative morphological analyses were based on external and cranial characters traditionally used in the taxonomy of Neotropical Myotis (see Novaes et al. 2022b ). Distribution modelling To predict the current distribution of Myotis armiensis , we used 16 occurrence records based exclusively on specimens from biological collections (Table 2 ) and climatic data from WorldClim 2.1, which provides 19 variables related to precipitation, temperature, and topography for the period 1970–2000. All variables had a spatial resolution of 10 arc-minutes (≈ 18 km). Table 2 Occurrence localities of Myotis armiensis based on museum specimens. Country Federative Unit Locality Coordinates Altitude Costa Rica Puntarenas Monteverde 10°16’N, 84°49’W 931 m Costa Rica Heredia Parque Nacional Braulio Carrillo 10°09’N, 83°59’W 1,000 m Panama Chiriquí Bugaba, La Amistad International Park 08°53’N, 82°36’W 2,214 m Panama Chiriquí Tierras Altas, Cerro Punta 08°52’N, 82°35’W 1,858 m Panama Chiriquí Tierras Altas, El Volcán 08°52’N, 82°44’W 1,280 m Panama Coclé La Mesa 08°38’N, 80°07’W 850 m Venezuela Distrito Capital Pico Avila 10°33’N, 66°52’W 2,092 m Venezuela Miranda Curupao 10°31’N, 66°38’W 1,160 m Venezuela Miranda San Andrés 10°22’N, 66°50’W 1,180 m Venezuela Aragua Estación Biológica Rancho Grande 10°21’N, 67°40’W 1,081 m Venezuela Carabobo Montalbán 10°15’N, 68°21’W 1,810 m Colombia Huila San Agustín, San Antonio village 01°57’N, 76°28’W 2,185 m Ecuador Tungurahuá Azuay 01°20’S, 78°12’W 1,660 m Ecuador Napo Quijos, Cabañas del Aliso 00°34’S, 77°53’W 2,249 m Ecuador Morona Santiago Baños 02°14’S, 78°25’W 3,115 m Ecuador Zamora Chinchipe Yantzaza, Cpo. Minero Fruta del Norte 03°45’S, 78°31’W 1,200 m Ecological niche models (ENMs) were generated using the ENMTML package in R (Andrade et al. 2020 ). We applied four algorithms to obtain more robust predictions: MaxEnt, SVM, BIOCLIM, and DOMAIN. To constrain prediction limits, we used the LTP method in conjunction with MAX_TS, which defines the threshold where the sum of sensitivity and specificity is maximized. Model performance was evaluated by splitting the occurrences into two subsets: 70% for training (model calibration) and 30% for testing (model evaluation). Four environmental suitability models were generated, one for each algorithm. Model accuracy was assessed using the True Skill Statistic (TSS), with values above 0.5 considered reliable (Dudík et al. 2004 ). An ensemble model was then built by averaging the best-performing model from each algorithm, based on TSS values, resulting in a single map of environmental suitability for M. armiensis . To convert the continuous suitability model into a binary presence–absence map, we applied the Maximum Sensitivity plus Specificity (MSS) threshold. Results New records to Northern South America The museums’ samples revision allowed the identification of 13 specimens of M. armiensis from Venezuela (USNM 370890, 370893, 370901, 370929, 373920, 387714, 387715) and one specimen from Colombia (FMNH 72175), being the first records of this species for both countries (see Appendix 2). Records for Venezuela were made throughout the Cordillera de la Costa, in Rancho Grande Biological Station, Aragua (10°21'N, 67°40'W); Montalbán, Carabobo (10°15'N, 68°21'W); Pico Avila, Distrito Capital (10°32'N, 66°52'W); Curapao, Guarenas, Miranda (10°31'N, 66°38'W); and San Andrés, Miranda (10°22'N, 66°50'W), at altitudes ranging from 1,080 to 2,100 m. These specimens come from mountainous deciduous rainforests included within the Central Cordillera ecoregion. The record for Colombia was based on a specimen captured in San Agustín, Huila (01°52'N, 76°17'W), at 2,185 m above sea level. This record was made in transition between moist and dry forests in the Colombian Northwestern Andean ecoregion. The specimens from Venezuela and Colombia presents all diagnostics characters described by Carrión-Bonilla and Cook ( 2020 ). The skull is comparatively large and robust (GLS 12.9–13.7), with sagittal and lambdoidal crests present. The shape of braincase roof inclined forward; posterior region of the skull being rounded and projected beyond the limit of the occipital condyles (Fig. 2 ). All specimens have long and woolly fur; dorsal hairs are unicolored and range from medium brown to bright reddish brown; ventral hairs are bicolored and with clear contrast, with dark brown base and the tips ranging from light brown to light yellow (Fig. 3 ). All specimens have a dense fur on the dorsal surface of the uropatagium that extends beyond the level of the knees, covering most of the tibia and reaching up to 2/3 of the basal portion of the uropatagium (Fig. 4 ). Confirmation of the taxonomic identity of this sample was made from the cyt-b sequencing of a specimen from Venezuela (USNM 370929), and the results are described below. Genetic structure and variation The cyt-b dataset showed considerable variation, with 417 variable sites, 324 of which were parsimony-informative. Myotis armiensis was recovered within a clade composed exclusively of species from Mexico, Central America and northern South America. It formed a monophyletic lineage and was recovered as the sister taxon to M. pilosatibialis , a relationship strongly supported by high nodal values (Fig. 5 ). Phylogenetic inference further indicated that M. armiensis comprises two geographically structured clades: one represented by samples from Central America (Panama) and the other by samples from northern South America (Ecuador and Venezuela). Genetic distance estimates under the K2p model indicated an average divergence of 6% between M. armiensis and its sister species, M. pilosatibialis (Table 3 ). Distances between M. armiensis and other species of the ruber group ranged from 10% to 13%. In contrast, the divergence between Central and South American populations of M. armiensis was 3%, while intrapopulation distances were approximately 1% in both geographic clades. Table 3 Average genetic distances (K2p) among Myotis species of the ruber species group based on cyt-b sequences. The intraspecific variation is presented in the diagonal, boldface, and enclosed in square brackets. Taxa 1 2 3 4 5 6 7 8 9 10 11 1. M. armiensis Central [0.01] 2. M. armiensis South 0.03 [0.01] 3. M. pilosatibialis 0.06 0.06 [0.02] 4. M. extremus 0.10 0.10 0.09 [0.02] 5. M. keaysi 0.11 0.11 0.11 0.12 [0.02] 6. M. ruber 0.13 0.13 0.12 0.13 0.09 [0.00] 7. M. moratellii 0.12 0.12 0.10 0.11 0.08 0.09 [0.00] 8. M. simus 0.12 0.12 0.12 0.14 0.11 0.11 0.10 [***] 9. M. midastactus 0.12 0.12 0.12 0.13 0.09 0.09 0.09 0.07 [***] 10. M. elegans 0.11 0.12 0.11 0.11 0.08 0.10 0.07 0.11 0.09 [0.00] 11. M. riparius 0.12 0.13 0.11 0.12 0.10 0.11 0.09 0.11 0.10 0.08 [0.05] Morphological variation The MANOVA results recovered statistically significant morphometric differences between Central and South American samples of M. armiensis (Pillai F 72 = 0.680, P = 0.026; Wilks F 72 = 0.319, P = 0.026). Pairwise post hoc tests showed that only the interorbital breadth (IOB) exhibited significant differences between Central and South American samples ( p = 0.01), while MAN and GLS showed marginal values ( p = 0.052 and 0.051, respectively; Fig. 6 ). For PCA analysis, only the first PC was retained based on BS test, accounting for 57% of the total craniometric variance, while the PC2 accounted for 12% of total variation (Table 4 ). Considering the positive relationship among the original measurements and the PC1, which are all positively correlated, size seems to be the biological factor responsible for this amount of craniometric variation. All samples are highly overlap considering the projection of individual scores on the first two PCs, denoting the overall craniometric similarity among M. armiensis and M. pilosatibialis samples (Fig. 7 ). Table 4 Eigenvalues, eigenvectors, and percentage of variance for the first two principal components (PCA), and the first three canonical variates (CVA), obtained for the two samples of Myotis armiensis and three samples of Myotis pilosatibialis. Eigenv. = eigenvalues; var. = variance. Measurements PC1 PC2 CV1 CV2 CV3 MAL -0.267 -0.232 - - - MAN -0.236 -0.270 -0.540 -5.629 4.584 GLS -0.215 -0.206 -2.078 -3.094 -3.057 CIL -0.216 -0.188 4.337 1.405 -1.309 MAB -0.247 0.147 -3.676 2.499 5.795 BCB -0.241 0.187 -1.332 5.260 -4.749 IOB -0.477 0.389 -5.602 -6.678 0.693 POB -0.245 0.616 13.720 -1.289 -0.590 BAC -0.358 -0.069 -2.168 3.566 2.479 BAM -0.321 -0.048 -1.506 -1.878 -4.355 MTL -0.249 -0.285 - - - M1M3 -0.280 -0.343 0.439 10.0788 2.387 Eigenv. 0.004 0.001 4.877 3.0409 2.263 % var. 57 12.3 42.4 26.4 20 In the CVA, the first three CVs account for 88% of the total variance, with the CV1 corresponding to 42% and the CV2 corresponding to 26% (Table 4 ). Despite the overall similarity observed in the PCA, the samples were recovered structured considering both the species and, in the case of M. armiensis , the geographic clades. The interpolation of individual scores on the CV1 recovered the difference between species, with most of the individuals of M. pilosatibialis with positive scores, while all but one individual of M. armiensis exhibiting negative scores. Among M. pilosatibialis , individuals from Mexico and Guatemala are mostly overlapped with scores between − 1 and 1, while individuals from Honduras and El Salvador exhibit scores mostly between 1 and 2. Most of the original measurements were negatively correlated with the CV1 except for the post-orbital breath (POB), which exhibited a positive correlation with CV1. It suggests that the difference between M. pilosatibialis and M. armiensis is mostly explained by general size, with the former being smaller than the latter, with individuals from Honduras and El Salvador being markedly small, but the post-orbital region is wider in M. pilosatibialis and narrow in M. armiensis (Fig. 7 ). The CV2 accounted for the differences between the two clades of M. armiensis , with the South American samples exhibiting almost all individuals with positive scores whereas the Central American group exhibited mostly negative scores. Recovering the observed in the MANOVA results, the differences between these groups are mostly explained by the interorbital constriction breadth (IOB), that exhibited negative correlations around − 0.2, along with mandible toothrow length (MAN) and greatest length of skull (GLS), which exhibited negative correlations slightly lower than − 0.2. On the other hand, braincase breadth (BCB) exhibited positive correlation with CV2, around 0.2. Considering this, South American individuals exhibit short and wide skulls with short mandible toothrow and narrow interorbital region, while Central American individuals exhibit long and narrow skulls with long mandible toothrow and wide interorbital region. Considering South and Central American samples, posterior classification recovered most of the individuals belonging to the given group or mistakenly assigned to one of M. pilosatibialis groups, respectively 56% and 25% for South American group and 54% and 15% for Central American group. Thus, less than 20% (3 individuals) of the South American group were suggested as member of the Central American group, while 31% (4 individuals) of the Central American group were suggested as belonging to the South American group (Fig. 7 ). Measurements these two groups are available in Table 5 . Table 5 Selected measurements (mm) of Myotis armiensis populations. Descriptive statistics include the mean, range (in parentheses), and sample size. See Table 1 for variable abbreviations. Mann-Whitney Test p-values was used to compare cranial measurements between samples and values statistically significant are bolded. Measurements with hyphen (–) were not tested. Measurements Central America South America mean (max – min) N mean (max – min) N FA 37.5 (35.8–38.7) 12 37.0 (35.9–38.4) 15 3MC 34.0 (32.8–35.2) 12 34.3 (33.4–35.8) 15 EL 12.0 (12.0–13.0) 10 12.5 (11.0–14.0) 12 LDF 6.5 (6.0–7.0) 4 7.0 (6.5–7.5) 8 LVF 5.2 (5.0–5.5) 4 5.8 (5.5–6.0) 8 GLS 13.46 (13.25–13.69) 12 13.34 (12.93–13.73) 16 CCL 12.14 (11.95–12.39) 12 11.99 (11.72–12.50) 16 CBL 12.74 (12.57–13.02) 12 12.62 (12.32–12.98) 16 CIL 12.94 (12.69–13.20) 12 12.84 (12.54–13.24) 16 BAL 11.62 (11.48–11.82) 12 11.46 (10.07–11.85) 16 ZYG 8.83 (8.67–9.10) 7 8.42 (8.06–8.83) 10 MAB 7.10 (6.83–7.45) 12 7.07 (6.82–7.42) 15 BCB 6.34 (6.21–6.56 12 6.39 (6.18–6.65) 16 POB 3.36 (3.16–3.65) 12 3.33 (3.21–3.49) 16 IOB 4.53 (4.26–4.81) 12 4.35 (4.09–4.77) 16 BAC 3.66 (3.52–3.82) 12 3.66 (3.47–3.99) 15 BAM 5.48 (5.31–5.76) 12 5.41 (5.51–5.82) 16 MTL 5.15 (5.04–5.26) 12 5.11 (4.92–5.35) 16 M1–3 2.92 (2.81–3.03) 12 2.91 (2.82–3.04) 16 MAL 9.87 (9.96–10.02) 11 9.82 (9.61–10.27) 13 MAN 5.54 (5.43–5.73) 12 5.46 (5.29–5.67) 16 Another interesting pattern recovered is that geographically close samples tended to be distinctly different on cranial dimensions, with small or none overlapping among them, while geographically distant samples were recovered overlapped. For example, individuals of M. pilosatibialis from Guatemala were partially overlapped with M. armiensis form South American group, and individuals of M. pilosatibialis from Mexico were partially overlapped with M armiensis from Central and South American groups. Individuals of M. pilosatibialis form Honduras and El Salvador were recovered as strikingly different from individuals of M. armiensis . Qualitative morphological analyses did not recover any pattern of distinction between OTUs from the characters. The sagittal crest is present in all specimens analyzed ( N = 28), ranging from low (50%) to median (50%) in samples from Central America ( N = 12); and superficial (6%), low (47%) or median (47%) in specimens from South America ( N = 16). The same pattern was repeated for lambdoidal crests, which varied among superficial (10%), low (~ 50%) or medium (~ 40%) for both population groups. The shape of braincase roof inclined forward was the predominant state (92%) in the samples from South America; while specimens from Central America had a braincase roof inclined forward (66%) or straight (34%). The shape of the posterior region of the skull, being rounded and projected beyond the limit of the occipital condyles in 92% of the South American individuals; while samples from Central America had a greater variation of this character, being 11% flattened and not projected and 89% with a rounded shape and subtly projected beyond the posterior limit of the occipital condyles. Virtually all specimens, regardless of geographical origin, have a dense fur on the dorsal surface of the uropatagium that extends beyond the level of the knees, covering most of the tibia and reaching up to 2/3 of the basal portion of the uropatagium. Only one individual from Ecuador (LACM 42951) had the fur extending to the knees. However, the condition of the skin may be biasing the observation. Specimens from the two OTUs showed the same color pattern, with unicolored dorsal fur ranging from medium brown to bright reddish brown. The ventral fur, on the other hand, is bicolored and with clear contrast, with dark brown base and the tips ranging from light brown to light yellow. Both populations of M. armiensis have long and woolly fur, but specimens from South America have slightly longer fur than those from Central America (Table 5 ). Distribution and ecological niche modelling Based on museum specimens, we found 16 occurrences localities for M. armiensis through Costa Rica, Panama, Colombia, Ecuador, and Venezuela, in altitudes ranging from 931 to 3,115 m above sea level, comprising habitats of tropical montane forests (Table 2 ). Records in Costa Rica and Panama were made in Moist Montane Forests and swampy bogs, with presence oak trees ( Quercus spp.) and bamboo ( Chusquea spp.) along Talamanca Cordillera. In Ecuador, Colombia, and Venezuela, M. armiensis was recorded in Evergreen Lower Montane Forest, Cloud Montane Forest, and Montane Dry Forest associated with Andes and Cordillera de la Costa. All algorithms showed satisfactory TSS values (> 0.8). The ensemble approach predicted that M. armiensis is highly associated with northern tropical Andes (and adjacent mountain ranges from Venezuela), occupying moist and dry montane forests from northern Venezuela to southern Peru (Fig. 8 ). In fact, 94% of the suitable areas for M. armiensis occupation is within the forested slopes of the Andes. Although there are no formal records of M. armiensis occurrence in Peru, ENM predicts high suitable areas for a large part of Peruvian Andes, from Campanquiz to Vilcabamba Cordilleras. Small areas of high suitability are also present in the mountainous areas of Costa Rica and Panama in the Talamanca Cordillera. However, the model indicates that there are no suitable areas that connect the populations of Central and South America. Areas of high suitability on the Caribbean islands, in tepuis mountain range of Venezuela and Brazil, and in the highlands forests from Northeastern Brazil can be interpreted as commission errors, as although have adequate environmental conditions for M. armiensis occurrence, are geographically isolated and, therefore, inaccessible to this species. The most important variables to predict the geographic distribution of M. armiensis were BIO10 (mean temperature of warmest quarter, 81%), BIO16 (precipitation of wettest quarter, 11%), and BIO4 (temperature seasonality, 2%). In general, the species occurs in areas that have high precipitation and low temperature variation. Taxonomy Myotis armiensis montensis , new subspecies Holotype: The specimen (USNM 370890) consists of skin, skull and mandible of an adult male collected during Smithsonian Venezuelan Project on 20 August 1965 (Figs. 2 – 4 ). The specimen is deposited in the mammal collection of the National Museum of Natural History (USNM, Washington, D.C., USA). The Cyt-b sequence is available in the National Center for Biological Information’s GenBank under accession number MZ345121. Type locality: Pico Avila, 5 km NNE Caracas, near Hotel Humboldt, Distrito Capital, Venezuela (10°32'N, 66°52'W, elevation 2,092 m). Other specimens: Another 17 specimens from Colombia, Ecuador, and Venezuela were examined and are listed in Appendix 2. Distribution. Occurs along Cordillera de la Costa and Andean humid forests from northern Venezuela to southern Ecuador, and it is likely to occur also in Peru, although have no official records. The core of its distribution is strongly associated with Andean evergreen montane forest, cloud montane forest, and montane dry forest, at altitudes ranging from 1,080 and 3,120 m a.s.l. Etymology: The subspecific epithet “ montensis ” refers to its distribution endemic to tropical forests along northern South American Mountain ranges. Diagnosis: Both populations of M. armiensis are morphologicaly quite similar, but the distinction of M. a. montensis in relation to M. a. armiensis can be made from the following set of characters: longer fur (LDF < 6.5, LVF 13.7; IOB > 4.7; MAN > 5.6); and lower sagittal and lambdoidal crests. Additionally, M. a. montensis can be readily distinguished from M. a. armiensis based on cytochrome-b gene tree, being reciprocally monophyletic. Description and comparisons: A medium to large species within Neotropical Myotis (FA 35.9–38.4). Its fur texture, cranial morphology, and phylogenetic position agree with the pattern observed in species allocated to the ruber species group. Ears are Bone Brown and medium-sized (EL 11–14 mm), reaching the portion of the rostrum between the eyes and nostrils when pushed forward. Tragus long and slender, with a broad base and narrower spear-shaped terminal half, almost straight anterior edge, and rounded tip. Membranes Bone Brown. The dorsal surface of the uropatagium is densely furred, with hairs that extends beyond the level of the knees, covering most of the tibia and reaching up to 2/3 of the basal portion of the uropatagium. There is no fringe of hairs along the trailing edge of the uropatagium. Plagiopatagium attached to the foot, at the level of the toes, by a broad band of membrane. Dorsal and ventral fur wooly and moderately long (LDH 6.5–7.5 mm, LVH 5.5–6.0 mm). Dorsal fur unicolored, ranging from medium brown to bright reddish brown. Ventral fur strongly bicolored, with dark brown base and the tips ranging from light brown to light yellow. The dental formula is I 2/3, C 1/1, PM 3/3, M 3/3 (2x) = 38, typical of most species of Myotis . Skull robust and large in length (GLS 12.9–13.7), virtually identical to M. a. armiensis in qualitative characters. Second upper premolar (P3) subtly displaced lingually, and smaller than first upper premolar (P2). First lower molar (m1) myotodont, with postcristid connecting hypoconid and entoconid (Menu 1987 ). Braincase elongated (oval shape) with the roof inclined forward in mostly specimens; sagittal and lambdoidal crests present, ranging from low to moderate in high; occipital region slightly rounded and subtle projected beyond the limit of the occipital condyles; well-developed mastoid processes. Frontal bone slightly sloping (ca. 145° angle); rostrum comparatively short (ca. 2/5 of skull total length). Myotis a. montensis is quite similar to M. a. armiensis in external and skull morphologies but vary at the frequency of characters and in size of some external and cranial measurements. Both geographic forms have reddish brown, long and woolly fur, but specimens from Andes have slightly longer fur than those from Central America (see Table 5 ). South American specimens from Andes ( M. a. montensis ) have lower sagittal and lambdoid crests than those in Central America ( M. a. armiensis ). South American individuals exhibit short and wide skulls with short mandible toothrow and narrow interorbital region, while Central American individuals exhibit long and narrow skulls with long mandible toothrow and wide interorbital region, with statistically significant differences (Table 5 ; Fig. 6 ). Myotis armiensis armiensis Carrión-Bonilla & Cook, 2020 Holotype: The specimen (MSB 262089) consists of skin, skull and skeleton of an adult male collected by J. A. Cook and collaborators on 20 March 2012. The specimen is deposited in the mammal collection of the Museum of Southwestern Biology, University of New Mexico (MSB, Albuquerque, USA), including tissues samples (NK 209314). The cyt-b sequence is available in the National Center for Biological Information’s GenBank under accession number MW025265. Type locality: La Amistad International Park Ranger Station, Bugaba District, Chiriquí Province, Panama (08°53’N, 82°36’W, elevation 2,214 m). Distribution: Occurs in premontane and montane tropical forests along Talamanca Cordillera from Costa Rica and Panama, in altitudes between 850 and 2,300 m a.s.l. Diagnosis: Can be distinguished from M. a. montensis from the following set of characters: shorter fur (LDF > 7.0, LVF > 5.5); longer and narrow skulls with longer mandible toothrow and wider interorbital region (GLS < 13.2; IOB < 4.8; MAN < 5.7); and higher sagittal and lambdoidal crests. Additionally, M. a. armiensis can be readily distinguished from M. a. montensis based on cytochrome-b gene tree, being reciprocally monophyletic. Discussion Variation and taxonomic considerations Myotis armiensis is a recently described species, and despite its unequivocal taxonomic validity, aspects of its variation and distribution remain poorly understood. The cyt-b phylogeny recovered two clades with strong biogeographic associations and an estimated genetic distance of 3.1%. Genetic distances between 2% and 11% in cyt-b may indicate either valid species or conspecific populations with high variation, warranting additional studies to clarify their specific status (Bradley and Baker 2001 ). In fact, some recognized Neotropical Myotis species show genetic distances equal to or even lower than 3% (Novaes et al. 2021 , 2022b ). Conversely, most Neotropical Myotis species display intraspecific genetic distances below 2% based on cyt-b sequences (Novaes et al. 2021 , 2022b , 2024b , 2025a ; Díaz et al. 2025 ). Thus, this comparatively high divergence between allopatric populations deserves further critical investigation. Among other factors, high divergences may result from isolation by distance or restricted gene flow (Slatkin 1987 , 1993 ), although this does not necessarily have direct implications for species delimitation and more data is required to taxonomic assessment (Johns and Avise 1998 ; Bradley and Baker 2001 ). Using a similar dataset, Carrión-Bonilla et al. ( 2024 ) applied three molecular species delimitation methods. One (mPTP) recovered the Central American (Talamanca Cordillera) and Andean populations as distinct species, while the other two (GMYC, ABGD) considered them conspecific. Despite this, those authors recognized M. armiensis as a monotypic taxon. In contrast, our morphometric analyses revealed differences in some cranial characters, though these were not corroborated by qualitative morphological traits. Ecological niche modeling suggested no suitable areas connecting the Andean and Talamanca populations, possibly reflecting strong geographic isolation and a consequent interruption in gene flow. Taken together, the phylogenetic structure, moderately high genetic divergence, and subtle size differences may reflect persistent geographic isolation and fixation of characters (Baker and Bradley 2006 ). Although these findings meet the criteria for considering the populations as divergent evolutionary lineages (De Queiroz 2007 ; Padial et al. 2010 ), the absence of morphometric discontinuities and reliable morphological or molecular synapomorphies led us to adopt a conservative approach, recognizing the trans- and cis-Andean populations as distinct allopatric subspecies. Nonetheless, we emphasize that broader taxonomic sampling and sequencing of additional specimens (and genes) could refine these results and potentially support the elevation of M. a. montensis to full species. Based on biological and phylogenetic species concepts as a framework, Avise and Ball ( 1990 ) defined subspecies as groups of populations that truly or potentially interbreed, are phylogenetically distinguishable, but remain reproductively compatible with other groups of the same species. These authors also emphasized that subspecies recognition should be based on multiple independent lines of evidence consistent with taxon distribution. However, direct evidence of interbreeding or reproductive compatibility is often difficult to obtain. Zachos ( 2016 ) argued that subspecies should be recognized when patterns of genetic or morphological variation are present, even if complete population structuring is lacking, and highlighted that the concept is inherently arbitrary. In such cases, disjunct populations with genetic, morphological, and/or ecological differentiation may be classified as subspecies, thereby organizing taxonomic and evolutionary knowledge (Zachos et al. 2014 ). More recently, De Queiroz ( 2020 ) proposed a revised concept of subspecies, defining them as incompletely separated lineages. In other words, subspecies are entities of the same fundamental kind as species (i.e., separately evolving metapopulation lineages), differing only in that they remain incompletely separated and thus belong to a more inclusive lineage, the species (De Queiroz 2020 , 2021 ). The case of M. armiensis fits well with the subspecies definitions proposed by Zachos ( 2016 ) and De Queiroz ( 2020 ), since the morphological and morphometric variation, coupled with disjunct distribution, indicates incompletely separated lineages. This supports our classification of M. a. armiensis for Talamanca populations and M. a. montensis for Andean populations. Evolutionary history A recent study estimated the divergence between M. armiensis and its sister species, M. pilosatibialis , at approximately 2.25 Ma (± 0.45 Ma) during the Early Pleistocene (Novaes et al. 2024a ). Although the split between the Talamanca and Andean lineages of M. armiensis has not been dated, it is reasonable to infer a more recent divergence, likely in the Middle Pleistocene. Myotis armiensis belongs to a clade composed exclusively of species from Mexico and Central America, being one of the most recently diverged members of this clade (Carrión-Bonilla and Cook 2020 ; Novaes et al. 2024b ). Considering its phylogenetic position, it is plausible that its origin lies in Mesoamerica, with the ancestral lineage adapted to cooler environments than currently lowlands, as supported by ecological niche models. The colonization of South America likely occurred via dispersal across the Isthmus of Panama during climatic periods favorable to population expansion. The Pleistocene was characterized by glacial (cold, dry) and interglacial (warm, wet) cycles (Ditlevsen 2022 ; Herbert 2023 ). During glaciations, glaciers advanced and high-elevation ecosystems (paramos, cloud forests, and montane forests) shifted downslope, expanding habitats now confined to highlands and forming mosaics of tropical forests and open savannas (Van Der Hammen 1974 ; Prance 1982 ; Loehle 2007; Cabanne et al. 2016 ). In contrast, interglacial periods brought glacial retreat, rising temperatures, and upslope displacement of these ecosystems, accompanied by expansion of ombrophilous and seasonal forests in lowlands (Van Der Hammen 1974 ; Prance 1982 ; Loehle 2007; Cabanne et al. 2016 ). These dynamics promoted cycles of fragmentation and reconnection of populations, generating opportunities for isolation and diversification in many taxa, including Myotis (Carnaval and Moritz 2008 ; Goin et al. 2012 ; Cabanne et al. 2016 ; Novaes et al. 2024a , 2025b ). Thus, during glacial phases, M. armiensis likely expanded into lower altitudes, increasing population size, colonizing South America, and occupying a wide range of suitable forests. Conversely, during interglacial phases, as cloud and montane forests contracted upslope, M. armiensis populations retreated to cooler and more stable highland habitats, where they remain today. Moreover, major mountain ranges may have acted as geographic barriers even during climatically stable intervals, promoting speciation and shaping evolutionary trajectories (Vuilleumier 1969 ; Cadle 1991 ; Patterson et al. 1992 ; Torres-Carvajal and Mafla-Endara 2013 ). The genetic structure revealed by our analyses, along with shifts in cranial character frequencies, may reflect recent population bottlenecks driven by Pleistocene environmental fluctuations, as documented for other Myotis species (Furman et al. 2013 ; Lu et al. 2013 ; Novaes et al. 2024a , 2025b ). These events may have caused population declines, reduced gene flow, and increased the frequency of rare haplotypes within populations (Willis and Wiese 1993 ; Hartl 2008). Although the evolutionary history of M. armiensis requires further study with broader genetic sampling, current spatial segregation patterns are sufficient to support recognition of two subspecies, consistent with the arguments of Zachos ( 2016 ) and De Queiroz ( 2020 ). Future phylogeographic research based on larger datasets and additional genetic markers will be essential to clarify demographic dynamics and diversification processes underlying intraspecific variation. Declarations Acknowledgements We are grateful for staff and curators of the institutions that received us and Edson Fiedler de Abreu for helping in generating molecular data. RLMN has received 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 has received financial support from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil; 313963/2018-5) and FAPERJ (E-26/200.967/2021). Author contributions RLMN, study conception and design, project management, data analysis, writing and review; MAGS, data analysis, writing and review; AC, data analysis, writing and review; DEW, advising and review; MW, advising and review; RM, project management, advising and review. All authors read and approved the final manuscript. Funding 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) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil; 313963/2018-5). Data availability DNA sequences generated in this study have been deposited in GenBank (Appendix I). Data from distribution is available in Table 2. Morphological data matrices are available upon request from the corresponding author. Conflict of interest The authors declare no conflict of interest. Ethics declaration Not applicable. References Andrade AFA, Velazco SJE, De Marco PJ (2020) ENMTML: An R package for a straightforward construction of complex ecological niche models. Environ Model Soft 125:104615. https://doi.org/10.1016/j.envsoft.2019.104615 Antonelli A (2021) The rise and fall of Neotropical biodiversity. Bot J Lin Soc 199:8–24. https://doi.org/10.1093/botlinnean/boab061 Avise JC, Ball RM (1990) Principles of genealogical concordance in species concepts and biological taxonomy. In: Futuyma D, Antonovics J (eds) Oxford surveys in evolutionary biology. Oxford University Press, Oxford, pp 45–67 Baker RJ, Bradley RD (2006) Speciation in mammals and the genetic species concept. J Mammal 87:643–662. https://doi.org/10.1644/06-mamm-f-038r2.1 Bradley RD, Baker RJ (2001) A test of the genetic species concept: cytochrome-b sequences and mammals. J Mammal 82: 960–973. https://doi.org/10.1644/1545-1542 (2001)082%3C0960:ATOTGS%3E2.0.CO;2 Brumfield RT, Capparella AP (1996) Historical diversification of birds in northwestern South America: a molecular perspective on the role of vicariant events. Evolution 50:1607–1624. https://doi.org/10.1111/j.1558-5646.1996.tb03933.x Cabanne GS, Calderón L, Trujillo-Arias N, Flores P, Pessoa R, d’Horta FM, Miyaki CY (2016) Effects of Pleistocene climate changes on species ranges and evolutionary processes in the Neotropical Atlantic Forest. Biol J Lin Soc 119:856–872. https://doi.org/10.1111/bij.12844 Cadle JE (1991) Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northern Perú: new species and comments on relationships and distribution patterns. Proc Acad Nat Sci Phil 143:1–96 Caraballo DA, Montani ME, Martınez LM, Antoniazzi LR, Sambrana TC, Fernandez 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:e0244750. https://doi.org/10.1371/journal. pone.0244750 Carnaval AC, Moritz C (2008) Historical climate modelling predicts patterns of current biodiversity in the Brazilian Atlantic Forest. J Biogeogr 35:1187–1201. https://doi.org/10.1111/j.1365-2699.2007.01870.x 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. J Mammal 105:241–258. https://doi.org/10.1093/jmammal/gyad124 Claridge MF (2010) Species are real biological entities. In: Ayala FK, Arp R (eds) Contemporary debates in philosophy of biology. Wiley-Blackwell, Malden, pp 91–109 Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nat Met 9:a772. https://doi.org/10.1038/nmeth.2109 De Queiroz K (2007) Species concepts and species delimitation. Syst Biol 56:879–886. https://doi.org/10.1080/10635150701701083 De Queiroz K (2020) An updated concept of subspecies resolves a dispute about the taxonomy of incompletely separated lineages. Herpetol Rev 51(2):459–461 De Queiroz K (2021) Response to critiscism of na updated subspecies concept. Herpetol Rev 52(4):773–776 Díaz DR, Medina CR, Arias S, López E, Carrión-Bonilla CA (2025) A new species of Myotis (Chiroptera: Vespertilionidae) from the coastal desert of southern Peru. J Mammal gyaf028. https://doi.org/10.1093/jmammal/gyaf028 Ditlevsen P (2022) The Pleistocene glacial cycles and millennial-scale climate variability. Atmos Ocean 60:233–244. https://doi.org/10.1080/07055900.2022.2077172 Dudík M, Phillips SJ, Schapire RE (2004) Performance guarantees for regularized maximum entropy density estimation. In: Shawe-Taylor J, Singer Y (eds) Conference on Computational Learning Theory (COLT). Lecture Notes in Computer Science, vol. 3120. Springer, Berlin. https://doi.org/10.1007/978-3-540-27819-1_33 Edgar RC (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucl Acid Res 32(5):1792–1797. https://doi.org/10.1093/nar/gkh340 Frontier S (1976) Étude de la decroissance des valeurs propres dans une analyze en composantes principales: comparison avec le modèle de baton brisé. J Exp Mar Biol Ecol 25(1):67–75. https://doi.org/10.1016/0022-0981(76)90076-9 Furman A, Çoraman E, Nagy ZL, Postawa T, Bilgin R, Gajewska M, Bogdanowicz W (2013) Phylogeography of the large Myotis bats (Chiroptera: Vespertilionidae) in Europe, Asia Minor, and Transcaucasia. Biol J Lin Soc 108:189–209. https://doi.org/10.1111/j.1095-8312.2012.01994.x Ghiselin MT (1997) Metaphysics and the origin of species. State University of New York, New York, p 377 Goin FJ, Gelfo JN, Chornogubsky L, Woodburne MO, Martin T (2012) Origins, radiations, and distribution of South American mammals: From greenhouse to icehouse worlds. In: Patterson BD, Costa LP (eds) Bones, clones, and biomes: the history and geography of recent Neotropical mammals. University of Chicago Press, Chicago, pp 20–50. https://doi.org/10.7208/9780226649214-003 Hammer Ø, Harper DAT, Ryan PD (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontol Elect 4(1):1–9 Hartl DL (2006) Principles of population genetics, 4th edn. OUP, New York, p 545 Hasegawa M, Kishino H, Yano T (1985) Dating of human-ape splitting by a molecular clock of mitochondrial DNA. J Mol Evol 22:160–174. https://doi.org/10.1007/bf02101694 Herbert TD (2023) The Mid-Pleistocene climate transition. Rev Earth Plan Sci 51:389–418. https://doi.org/10.1146/annurev-earth-032320-104209 Honaker J, King G, Blackwell M (2011) Amelia II: A program for missing data. J Stat Softw 45(1):1–47. https://doi.org/10.18637/jss.v045.i07 Hull D (1994) Contemporary systematic philosophies. In: Sober E (ed) Conceptual issues in evolutionary biology, 2nd edn. MIT Press, Cambridge, pp 295–330 Hutter CR, Lambert SM, Wiens JJ (2017) Rapid diversification and time explain amphibian richness at different scales in the tropical Andes, Earth’s most biodiverse hotspot. Ame Nat 190:828–843. https://doi.org/10.1086/694319 Johns GC, Avise JC (1998) A comparative summary of genetic distances in the vertebrates from the mitochondrial cytochrome b gene. Mol Biol Evol 15(11):1481–1490. https://doi.org/10.1093/oxfordjournals.molbev.a025875 Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. J Mol Evol 16:111–120. https://doi.org/10.1007/bf01731581 Kovarovic K, Aiello LC, Cardini A, Lockwood CA (2011) Discriminant function analysis in archaeology: are classification rates too good to be true? J Archaeol Sci 38:3006–3018. https://doi.org/10.1016/j.jas.2011.06.028 Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular evolutionary genetics analysis across computing platforms. Mol Biol Evol 35:1547–1549. https://doi.org/10.1093/molbev/msy096 Larsen RJ, Knapp MC, Genoways HH, Khan FAA, Larsen PA, Wilson DE, Baker RJ (2012a) 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 Larsen RJ, Larsen PA, Genoways HH, Catzeflis FM, Geluso K, Kwiecinski KK, Pedersen SC, Simal F, Baker RJ (2012b) Evolutionary history of Caribbean species of Myotis , with evidence of a third Lesser Antillean endemic. Mammal Biol 77:124–134. https://doi.org/10.1016/j.mambio.2011.11.003 Loehle C (2006) Predicting Pleistocene climate from vegetation in North America. Clim Past 3:109–118. https://doi.org/10.5194/cp-3-109-2007 Lu G, Lin A, Luo J, Blondel DV, Meiklejohn KA, Sun K, Feng J (2013) Phylogeography of the Rickett’s big-footed bat, Myotis pilosus (Chiroptera: Vespertilionidae): a novel pattern of genetic structure of bats in China. BMC Evol Biol 13:e241. https://doi.org/10.1186/1471-2148-13-241 Menu H (1987) Morphotypes dentaires actuels et fossiles des chiropteres vespertilionines, 2 partie: implications systematiques et phylogeniques. Paleovertebrata 17(1):77–150 Meseguer AS, Michel A, Fabre PH, Pérez-Escobar AO, Chomicki G, Riina R, Antonelli A, Antoine PO, Delsuc F, Condamine FL (2022) Diversification dynamics in the Neotropics through time, clades, and biogeographic regions. eLife 11:e74503. https://doi.org/10.7554/eLife.74503 Mitteroecker P, Bookstein F (2011) Linear discrimination, ordination, and the visualization of selection gradients in modern morphometrics. Evol Biol 38:100–114. https://doi.org/10.1007/s11692-011-9109-8 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. Mammal Biol 76(5):592–607. https://doi.org/10.1016/j.mambio.2011.01.003 Moratelli R, Burgin C, Cláudio VC, Novaes RLM, López-Baucells A, Haslauer R (2019) Family Vespertilionidae (Vesper Bats). In: Wilson DE, Mittermeier AR (eds) Handbook of the mammals of the world, vol 9. Bats. Lynx Editions, Barcelona, pp 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 1015:145–167. https://doi.org/10.3897/zookeys.1015.59248 Novaes RLM, Wilson DE, Moratelli R (2022a) Catalogue of primary types of Neotropical Myotis (Chiroptera, Vespertilionidae). ZooKeys 1105:127–164. https://doi.org/10.3897/zookeys.1105.85055 Novaes RLM, Cláudio VC, Carrión-Bonilla C, Abreu EF, Wilson DE, Maldonado JE, Weksler M (2022b) Variation in the Myotis keaysi complex (Chiroptera, Vespertilionidae), with description of a new species from Ecuador. J Mammal 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 bioacoustics data. Zootaxa 5188(5):430–452. https://doi.org/10.11646/zootaxa.5188.5.2 Novaes RLM, Cláudio VC, Wilson DE, Weber MM, Weksler M, Moratelli R (2024a) Integrative taxonomy and evolutionary history reveal cryptic diversity in Myotis riparius (Chiroptera: Vespertilionidae). Biol J Lin Soc 142:91–112. https://doi.org/10.1093/biolinnean/blad123 Novaes RLM, Cláudio VC, Wilson DE, Weksler M, Moratelli R (2024b) Taxonomic status of Myotis extremus (Chiroptera, Vespertilionidae) from Mesoamerica, with comments on the distribution and systematics of Myotis nigricans . Zool Anz 308:99–110. https://doi.org/10.1016/j.jcz.2023.11.009 Novaes RLM, Cláudio VC, Bertocchi NA, Oliveira K, Semedo TBF, Saldanha J, Wilson DE, Moratelli R (2025a) Unveiling the shelf life: a new cryptic species of Myotis (Chiroptera, Vespertilionidae) from South America revealed by an integrative taxonomy approach. J Mammal 106(4):878–897. https://doi.org/10.1093/jmammal/gyaf016 Novaes RLM, Cláudio VC, Bertocchi NA, Abreu EF, Wilsom DE, Maldonado JE, Moratelli R (2025b) Islands in the stream: Distribution of Myotis attenboroughi (Chiroptera, Vespertilionidae) in Grenada and mainland South America illuminates the evolutionary history of Caribbean Myotis . Evol Syst 9:107–122. https://doi.org/10.3897/evolsyst.9.146107 Padial JM, Miralles A, La Riva ID, Vences M (2010) The integrative future of taxonomy. Front Zool 7:e16. https://doi.org/10.1186/1742-9994-7-16 Patterson BD, Pacheco V, Ashley MV (1992) On the origins of the western slope region of endemism: systematics of fig-eating bats, genus Artibeus . Mem Mus Hist Nat Lima 21:189–205 Patterson BD, Solari S, Velazco PM (2012) The role of the Andes in the diversification and biogeography of Neotropical mammals. In: Patterson BD, Costa LP (eds) Bones, clones, and biomes: the history and geography of recent Neotropical mammals. University of Chicago Press, Chicago, pp 351–378. https://doi.org/10.7208/9780226649214-015 Patterson BD, Webala PW, Peterhans JCK, Goodman SM, Bartonjo M, Demos TC (2019) Genetic variation and relationships among Afrotropical species of Myotis (Chiroptera: Vespertilionidae). J Mammal 100:1130–1143. https://doi.org/10.1093/jmammal/gyz087 Peres-Neto PR, Jackson DA, Somers KM (2005) How many principal components? Stopping rules for determining the number of non-trivial axes revisited. Comp Stat Data Anal 49:974–997. https://doi.org/10.1016/j.csda.2004.06.015 Platt RN, 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. Syst Biol 67(2):236–249. https://doi.org/10.1093/sysbio/syx070 Prance GT (1982) A review of the phytogeographic evidences for Pleistocene climate changes in the Neotropics. Miss Bot Gard 69(3):594–624. https://doi.org/10.2307/2399085 Rambaut A, Drummond AJ (2009) Tracer v.1.5: MCMC Trace Analysis Packpage. Available at http://tree.bio.ed.ac.uk/software/tracer/ Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 32: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst Biol 61:539–542. https://doi.org/10.1093/sysbio/sys029 Ruedi M, Mayer F (2001) Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggest deterministic ecomorphological convergences. Mol Phylogen Evol 21:463–448. https://doi.org/10.1006/mpev.2001.1017 Ruedi M, Csorba G, Lin LK, Chou CH (2015) Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920:301–342. http://dx.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 (2025) Bat species of the world: A taxonomic and geographic database. Version 1.7. Available at https://batnames.org/home.html , Accessed on April 09, 2025 Slatkin M (1987) Gene flow and the geographic structure of natural populations. Science 236:787–792. https://doi.org/10.1126/science.3576198 Slatkin M (1993) Isolation by distance in equilibrium and non-equilibrium populations. Evolution 47:264–279. https://doi.org/10.1111/j.1558-5646.1993.tb01215.x Solís A, Kohlmann B, Alvarado GE (2024) A Costa Rican biological puzzle—The geological, landscape, and vicariance history as a conditioning of the past and present biogeography. In: Quesada-Román A (ed) Landscapes and landforms of Costa Rica. Springer, Cham. https://doi.org/10.1007/978-3-031-64940-0_10 Stadelmann B, Lin LK, Kunz TH, Ruedi M (2007) Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes. Mol Phylogen Evol 43:32–48. https://doi.org/10.1016/j.ympev.2006.06.019 Torres-Carvajal O, Mafla-Endara P (2013) Evolutionary history of Andean Pholidobolus and Macropholidus (Squamata: Gymnophthalmidae) lizards. Mol Phylogenet Evol 68(2):212–217. https://doi.org/10.1016/j.ympev.2013.03.013 Van Der Hammen T (1974) The Pleistocene changes of vegetation and climate in tropical South America. J Biogeog 1:3–26. https://doi.org/10.2307/3038066 Viale M, Garreaud R (2015) Orographic effects of the subtropical and extratropical Andes on upwind precipitating clouds. JGR Atmos 120:4962–4974. https://doi.org/10.1002/2014JD023014 Vuilleumier F (1969) Pleistocene speciation in birds living in the high Andes. Nature 223:1179–1180. https://doi.org/10.1038/2231179a0 Willis K, Wiese RJ (1993) Effect of new founders on retention of gene diversity in captive populations: A formalization of the nucleus population concept. ZooBiology 12(6):535–548. https://doi.org/10.1002/zoo.1430120605 Zachos FE (2016) Species concepts in biology: historical development, theoretical foundations and practical relevance. Springer International Publishing, Geneve, p 220. https://doi.org/10.1007/978-3-319-44966-1 Zachos FE, Mattioli S, Ferretti F, Lorenzini R (2014) The unique Mesola red deer of Italy: taxonomic recognition ( Cervus elaphus italicus nova ssp., Cervidae) would endorse conservation. Ital J Zool 81:136–143. https://doi.org/10.1080/11250003.2014.895060 Supplementary Files Appendix.docx Cite Share Download PDF Status: Published Journal Publication published 10 Feb, 2026 Read the published version in Mammalian Biology → Version 1 posted Reviewers agreed at journal 15 Oct, 2025 Reviewers invited by journal 15 Oct, 2025 Editor assigned by journal 10 Oct, 2025 First submitted to journal 08 Oct, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7794885","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":529852958,"identity":"9aace6c9-39a1-494d-a6fb-7f9fef848e98","order_by":0,"name":"Roberto Leonan Morim Novaes","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6UlEQVRIiWNgGAWjYFACHjApA+FUADEzcwMRWhKgGhnOgLQwkqKFsQ1M4tdi3t578HHlDxsefvbeg58L59VG87cDtfyo2IZTi8yZc8mGZxLSeCR7ziVLz9x2PHfGYcYGxp4zt3FqkZDIMZNsSDjMY3Ajx0Cad9ux3AagFmbGNrxazH+CtNjfyDH+zTvnWO58IrSYMYJtATKkeRtqcjcQ1MJzLlmyIS2NR+LMGTNrnmMHcjcCtRzE6xdgQH1ssLGR42/vMb7NU1OXO+/84YMPflTg1oIODoPJA0SrB4I6UhSPglEwCkbBCAEADeZVHxPtcsoAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0003-1657-2807","institution":"Fundação Oswaldo Cruz: Fundacao Oswaldo Cruz","correspondingAuthor":true,"prefix":"","firstName":"Roberto","middleName":"Leonan Morim","lastName":"Novaes","suffix":""},{"id":529852959,"identity":"ea7dc683-96e5-475e-99f2-0013c77ec5ee","order_by":1,"name":"Marco Antônio Guimarães-Silva","email":"","orcid":"","institution":"Fundação Oswaldo Cruz: Fundacao Oswaldo Cruz","correspondingAuthor":false,"prefix":"","firstName":"Marco","middleName":"Antônio","lastName":"Guimarães-Silva","suffix":""},{"id":529852960,"identity":"251a1d20-f9d5-4bce-833f-dd1475f541ba","order_by":2,"name":"Aldo Caccavo","email":"","orcid":"","institution":"Universidade Federal de Juiz de Fora","correspondingAuthor":false,"prefix":"","firstName":"Aldo","middleName":"","lastName":"Caccavo","suffix":""},{"id":529852961,"identity":"253fd452-36ae-4857-b6c7-ab113a382ca4","order_by":3,"name":"Don E. 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(blue).\u003c/p\u003e","description":"","filename":"Figure1OccurrencepointsofMyotisarmiensis.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/e555ab3edb3b986e4c660c0b.jpg"},{"id":94673383,"identity":"f5a40bca-3cac-4a5c-a304-86d6656b4184","added_by":"auto","created_at":"2025-10-29 13:41:21","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":343665,"visible":true,"origin":"","legend":"\u003cp\u003eDorsal, ventral, and lateral views of the skull and lateral view of the mandible of the holotype of \u003cem\u003eMyotis armiensis montensis\u003c/em\u003efrom Pico Avila, Distrito Capital, Venezuela (USNM 370890).\u003c/p\u003e","description":"","filename":"Figure2Skull.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/c706cbd4f013197db8a1c273.jpg"},{"id":94673495,"identity":"cb55a33f-be06-445f-ab0f-0035a3cfb972","added_by":"auto","created_at":"2025-10-29 13:41:26","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":2072324,"visible":true,"origin":"","legend":"\u003cp\u003eDorsal (left) and ventral (right) views of the skin of the holotype of \u003cem\u003eMyotis armiensis montensis\u003c/em\u003e from Pico Avila, Distrito Capital, Venezuela (USNM 370890).\u003c/p\u003e","description":"","filename":"Figure3Skin.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/b0c001abd5d4d3b023a2676b.jpg"},{"id":94666197,"identity":"3ca2b96d-8650-4ca9-a35f-329b80622f42","added_by":"auto","created_at":"2025-10-29 12:33:28","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1413714,"visible":true,"origin":"","legend":"\u003cp\u003eDorsal surface of uropatagium densely furred in the holotype of \u003cem\u003eMyotis armiensis montensis\u003c/em\u003e from Pico Avila, Distrito Capital, Venezuela (USNM 370890).\u003c/p\u003e","description":"","filename":"Figure4Uropatagiumfur.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/014b87e49c98089e1af58a2f.jpg"},{"id":94666241,"identity":"78906cf5-a798-4bf9-9366-4614f7081834","added_by":"auto","created_at":"2025-10-29 12:33:32","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":1786236,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree resulting from the Bayesian inference of cytochrome-b sequences (1,140 bp) of Neotropical \u003cem\u003eMyotis\u003c/em\u003e. Nodal support was calculated by posterior probabilities for BI.\u003c/p\u003e","description":"","filename":"Figure5Phylogenetictree.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/a91fad97793e04e0347b67f6.jpg"},{"id":94666192,"identity":"c28d59b9-df96-4c92-884b-557bde4c3765","added_by":"auto","created_at":"2025-10-29 12:33:28","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":1131307,"visible":true,"origin":"","legend":"\u003cp\u003eViolin and box plot showing the pattern of distribution of three selected cranial measurements (GLS, MAN and IOB) for the two geographic clades recovered for \u003cem\u003eMyotis armiensis\u003c/em\u003e, South (blue) and Central (red).\u003c/p\u003e","description":"","filename":"Figure6Plotvariationincranialcharacters.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/533208cc7f19999fe055c729.jpg"},{"id":94666195,"identity":"4b9a2c81-a65c-4712-ae4d-42975b5c685f","added_by":"auto","created_at":"2025-10-29 12:33:28","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":2215415,"visible":true,"origin":"","legend":"\u003cp\u003eInterpolation of individual scores (left) and vector correlation (right) for the first two principal components (PCA, above) and the first two canonical variates (CVA, below) for the two geographic samples of \u003cem\u003eMyotis armiensis\u003c/em\u003e, Central America (red) and South America (blue), along with three samples of \u003cem\u003eMyotis pilosatibialis\u003c/em\u003e (grey). Shaded polygons represent convex hull for each of the five samples (above) and for the two samples of \u003cem\u003eM. armiensis\u003c/em\u003e and one encompassing all individuals of \u003cem\u003eM. pilosatibialis\u003c/em\u003e(below). Above: vectorial correlations marked in grey represent cranial measurements only used in PCA. Below: \u003cem\u003eMyotis armiensis\u003c/em\u003e: Central America = red diamonds; South America = blue dots; \u003cem\u003eMyotis pilosatibialis\u003c/em\u003e: Mexico = downward triangle, Guatemala = square, and Honduras + El Salvador = upward triangle. Note that the first CV account for the difference between species while the second CV account for the difference between the two clades found within \u003cem\u003eM. armiensis\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"Figure7PlotPCAandCVA.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/58544d884a3338ddb44e4d24.jpg"},{"id":94666198,"identity":"8b89771e-153e-4dda-95fe-cae3231e516a","added_by":"auto","created_at":"2025-10-29 12:33:28","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":11046293,"visible":true,"origin":"","legend":"\u003cp\u003eCurrent species distribution model for \u003cem\u003eMyotis armiensis\u003c/em\u003e: A) environmental suitability map; and B) binary distribution maps obtained with the Maximum Sensitivity and Specificity threshold.\u003c/p\u003e","description":"","filename":"Figure8ENM.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/6d48fbcd51484329ad38fea0.jpg"},{"id":102785494,"identity":"8f900a35-a6b3-4604-8c84-83bc2f272db2","added_by":"auto","created_at":"2026-02-16 16:07:34","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":23503432,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/558e0dcf-443a-4761-b66e-94343a4f83db.pdf"},{"id":94666225,"identity":"aeb026b5-69d0-42af-ac24-87efab092434","added_by":"auto","created_at":"2025-10-29 12:33:30","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":28586,"visible":true,"origin":"","legend":"","description":"","filename":"Appendix.docx","url":"https://assets-eu.researchsquare.com/files/rs-7794885/v1/b6bc653829447dff152e6b83.docx"}],"financialInterests":"","formattedTitle":"On the variation, distribution and evolutionary history of Myotis armiensis (Chiroptera, Vespertilionidae) along Neotropical mountain ranges, including the description of new subspecies","fulltext":[{"header":"Introduction","content":"\u003cp\u003eSpecies are cornerstones of current evolutionary theory (Hull \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Ghiselin \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Claridge \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and several concepts and approaches to species delimitation have been proposed (see review in de Queiroz \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). However, the boundary between intra- and interspecific variation remains one of the greatest challenges of modern taxonomy, especially in cryptic taxa, where phenotypic variation remains undetectable even after speciation events (Padial et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Novaes et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eNeotropics harbors the richest biodiversity in the world, and several studies have shown that its species diversification is linked to complex interactions between geological and climatic history in the Americas, resulting in heterogeneous habitats (Antonelli \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Meseguer et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In this context, the Mountain ranges, such as the Andes, represent an important biogeographic feature in South America, influencing rivers, precipitation, biome limits, and species distributions (Patterson et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Viale and Garreaud \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The role of the Central America Cordilleras and Andes Mountain range in limiting gene flow and promoting species diversification has been widely documented for many vertebrate taxa, including amphibians, birds, and mammals (Brumfield and Capparella \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Patterson et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Hutter et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Sol\u0026iacute;s et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The complex and relatively recent Andean uplift, for example, promoted diversification in multiple taxa, but also the subdivision of relatively young species that rapidly accumulated high genetic differentiation while showing little morphological divergence (i.e., cryptic species formation), as seen in some bats of the genus \u003cem\u003eMyotis\u003c/em\u003e Kaup, 1829 (Novaes et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cem\u003eMyotis\u003c/em\u003e is widely distributed worldwide and represents the most diverse bat genus, with approximately 140 valid species (Moratelli et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Simmons and Cirranello \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2025\u003c/span\u003ea). Recent systematic reviews have revealed unexpectedly high diversity in Neotropical \u003cem\u003eMyotis\u003c/em\u003e (Moratelli et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Carri\u0026oacute;n-Bonilla et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Novaes et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e). Currently, 43 species of \u003cem\u003eMyotis\u003c/em\u003e are recognized in the Neotropics, about 35% of which were described after 2010 (Carri\u0026oacute;n-Bonilla et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Novaes et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e). Many of these recently described species still have poorly defined distributional limits, including \u003cem\u003eMyotis armiensis\u003c/em\u003e Carri\u0026oacute;n-Bonilla \u0026amp; Cook, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e. \u003cem\u003eMyotis armiensis\u003c/em\u003e was described based on molecular and morphological data from specimens collected in Costa Rica, Panama, and Ecuador (Carri\u0026oacute;n-Bonilla and Cook \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, there are still gaps in its known distribution, likely due to phenotypic similarity with other species, particularly \u003cem\u003eMyotis keaysi\u003c/em\u003e J.A. Allen, 1914 and \u003cem\u003eMyotis pilosatibialis\u003c/em\u003e LaVal, 1973, which may complicate identification (Carri\u0026oacute;n-Bonilla and Cook \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eAs part of an ongoing review of Neotropical \u003cem\u003eMyotis\u003c/em\u003e, we identified individuals of \u003cem\u003eM. armiensis\u003c/em\u003e from Venezuela and Colombia, thereby expanding its known distribution, and obtained additional samples from Costa Rica, Panama, and Ecuador. We evaluated the intraspecific morphological and molecular variation of \u003cem\u003eM. armiensis\u003c/em\u003e and assessed its distinction from \u003cem\u003eM. pilosatibialis\u003c/em\u003e. In addition, we estimated the species\u0026rsquo; potential distribution using ecological niche modeling. Because the signature of vicariance caused by the uplift of the Andes is evident in numerous mammalian taxa, particularly in many bat lineages (see Patterson et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), we tested the hypothesis that trans-Andean and cis-Andean populations of \u003cem\u003eM. armiensis\u003c/em\u003e may represent different evolutionary lineages.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eMolecular data and analyses\u003c/h2\u003e\u003cp\u003eMolecular analyses were based on 96 cytochrome b (cyt-b, ~\u0026thinsp;1,140 bp) sequences from 29 species of Neotropical \u003cem\u003eMyotis\u003c/em\u003e, along with two sequences from other Myotinae bats (\u003cem\u003eMyotis emarginatus\u003c/em\u003e and \u003cem\u003eSubmyotodon latirostris\u003c/em\u003e) and \u003cem\u003eKerivoula papillosa\u003c/em\u003e (Temminck, 1840), which was used as outgroup (Appendix 1). All sequences were retrieved from NCBI GenBank, including those previously generated by us. The sequence of \u003cem\u003eM. armiensis\u003c/em\u003e from Venezuela (USNM 370890; GenBank Accession MZ345121) was obtained from historical DNA extracted from a museum specimen, with details provided in Novaes et al. (\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe dataset of 99 cyt-b sequences was aligned using the MUSCLE algorithm (Edgar \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) with default parameters, as implemented in MEGA X (Kumar et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The best-fitting model of nucleotide substitution for phylogenetic analyses was selected in JModelTest 2 (Darriba et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) under the Bayesian Information Criterion (BIC). The Hasegawa\u0026ndash;Kishino\u0026ndash;Yano model (HKY; Hasegawa et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1985\u003c/span\u003e), corrected for among-site rate heterogeneity with a gamma distribution and a proportion of invariant sites (HKY\u0026thinsp;+\u0026thinsp;Γ\u0026thinsp;+\u0026thinsp;I), was identified as the best fit.\u003c/p\u003e\u003cp\u003ePhylogenetic reconstruction was performed using Bayesian inference (BI) in MrBayes 3.2 (Ronquist et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) with a coupled Markov chain Monte Carlo (MCMC) approach. Four simultaneous Markov chains were run for 100,000,000 generations, sampling every 10,000 generations. The first 26,000 trees were discarded as burn-in, and posterior probabilities were calculated from the consensus of the remaining trees. The robustness of Bayesian sampling was assessed by examining the effective sample size (ESS) of free parameters in Tracer v1.5 (Rambaut and Drummond \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). All parameters showed ESS values greater than 300, and convergence was confirmed by plotting log-likelihood values against generation time, indicating reliable performance.\u003c/p\u003e\u003cp\u003eGenetic distances were estimated under the Kimura two-parameter (K2P) model (Kimura \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e1980\u003c/span\u003e), as implemented in MEGA X. This method calculates pairwise distances by estimating the proportion of nucleotide differences between sequences.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eMorphological data and analyses\u003c/h3\u003e\n\u003cp\u003eFor this study, we examined 29 specimens of \u003cem\u003eMyotis armiensis\u003c/em\u003e and 40 specimens of \u003cem\u003eM. pilosatibialis\u003c/em\u003e deposited in the collections of the Royal Ontario Museum (ROM, Toronto, Canada), American Museum of Natural History (AMNH, New York, USA), Carnegie Museum of Natural History (CM, Pittsburgh, USA), University of Kansas (KU, Lawrence, USA), Natural History Museum of Los Angeles County (LACM, Los Angeles, USA), Field Museum of Natural History (FMNH, Chicago, USA), Museum of Texas Tech University (TTU, Lubbock, USA), and the Smithsonian National Museum of Natural History (USNM, Washington, DC, USA) (Appendix 2). Specimens were identified based on morphological characters described by Carri\u0026oacute;n-Bonilla and Cook (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and by comparison with type-series material.\u003c/p\u003e\u003cp\u003eQuantitative morphological analyses were based on five external and 16 craniodental measurements (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). To minimize potential noise from ontogenetic variation, only adult individuals were included. Samples were organized into Operational Taxonomic Units (OTUs) following the results of phylogenetic studies on \u003cem\u003eM. armiensis\u003c/em\u003e (e.g., Carri\u0026oacute;n-Bonilla and Cook \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Carri\u0026oacute;n-Bonilla et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), which identified two clades: one composed exclusively of Central American specimens and the other of South American specimens (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Samples of \u003cem\u003eM. pilosatibialis\u003c/em\u003e were divided into OTUs corresponding to the smallest possible geographic groups that showed morphological cohesion.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eExternal and craniodental measurements obtained from \u003cem\u003eMyotis\u003c/em\u003e specimens.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"2\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMeasurement and acronym\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDescription\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eForearm length (FA)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the elbow to the distal end of the forearm including carpals\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eThird metacarpal length (3MC)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the proximal to the distal extremity of the third metacarpal\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEar length (Ear)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the base to the apex of the ear\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLength of dorsal fur (LDF)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMeasured at the midpoint of the scapulae\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLength of ventral fur (LVF)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMeasured at the midpoint of the sternum\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGreatest length of skull (GLS)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the apex of the upper internal incisors, to the occiput\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCondylo-canine length (CCL)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the anterior surface of the upper canines to the occipital condyles limit\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCondylo-basal length (CBL)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the premaxillae to a line connecting the occipital condyles\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCondylo-incisive length (CIL)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the apex of upper internal incisors to the occipital condyles limit\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBasal length (BAL)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLeast distance from the apex of upper internal incisors to the ventral margin of the foramen magnum\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eZygomatic breadth (ZYG)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eZygomatic breadth\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMastoid breadth (MAB)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGreatest breadth across the mastoid region\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBraincase breadth (BCB)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGreatest breadth of the globular part of the braincase\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePostorbital breadth (POB)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLeast breadth across frontals posterior to the postorbital bulges\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eInterorbital breadth (IOB)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLeast breadth between the orbits\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBreadth across canines (BAC)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGreatest breadth across outer edges of the crowns of upper canines, including cingulae\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBreadth across molars (BAM)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGreatest breadth across outer edges of the crowns of upper molars\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMaxillary toothrow length (MTL)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the upper canine to M3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMolariform toothrow length (M1\u0026ndash;3)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom M1 to M3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMandibular length (MAL)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the mandibular symphysis to the condyloid process\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMandibular toothrow length (MAN)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFrom the lower canine to m3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eWe used three multivariate approaches to investigate morphometric differences between the Central and South American clades of \u003cem\u003eM. armiensis\u003c/em\u003e: (i) a multivariate analysis of variance (MANOVA), (ii) a principal component analysis (PCA) based on the variance\u0026ndash;covariance matrix of log-transformed measurements, and (iii) a canonical variate analysis (CVA) based on the variance\u0026ndash;covariance matrix of untransformed data. All analyses were performed in R v.4.0 using the packages stats, MASS, and lattice.\u003c/p\u003e\u003cp\u003eDistribution patterns of measurements were visualized using violin plots and boxplots in PAST v.4.07b (Hammer et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Both graphical assessments and MANOVA tests were performed exclusively on \u003cem\u003eM. armiensis\u003c/em\u003e samples. For PCA and CVA, we also included \u003cem\u003eM. pilosatibialis\u003c/em\u003e, a closely related and morphologically similar species, organized into three geographic groups: \u0026ldquo;Mexico,\u0026rdquo; \u0026ldquo;Guatemala,\u0026rdquo; and \u0026ldquo;Honduras\u0026thinsp;+\u0026thinsp;El Salvador.\u0026rdquo; Including these groups helped avoid spurious segregation patterns that might arise when comparing only two populations, especially in CVA. Slightly damaged skulls with up to two missing measurements were retained, with missing values estimated using an expectation\u0026ndash;maximization with bootstrapping (EMB) algorithm, as implemented in the Amelia II package (Honaker et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) in R.\u003c/p\u003e\u003cp\u003eThe statistical significance of principal components was evaluated with the Broken Stick (BS) test (Frontier \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Peres-Neto et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). To reduce the risk of overfitting in CVA due to a low sample-size-to-variable ratio (Kovarovic et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Mitteroecker and Bookstein \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), we restricted the analysis to a subset of 10 variables (MAN, GLS, CIL, MAB, BCB, IOB, POB, BAC, BAM, and M1M3) representing overall skull dimensions, excluding measurements that showed high correlations (r\u0026thinsp;\u0026gt;\u0026thinsp;0.8) with one or more of the selected variables. We report eigenvalues, eigenvectors, explained variance for both principal components (PCs) and canonical variates (CVs), as well as scatterplots of PC and CV scores. Qualitative morphological analyses were based on external and cranial characters traditionally used in the taxonomy of Neotropical \u003cem\u003eMyotis\u003c/em\u003e (see Novaes et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eDistribution modelling\u003c/h3\u003e\n\u003cp\u003eTo predict the current distribution of \u003cem\u003eMyotis armiensis\u003c/em\u003e, we used 16 occurrence records based exclusively on specimens from biological collections (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) and climatic data from WorldClim 2.1, which provides 19 variables related to precipitation, temperature, and topography for the period 1970\u0026ndash;2000. All variables had a spatial resolution of 10 arc-minutes (\u0026asymp;\u0026thinsp;18 km).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eOccurrence localities of \u003cem\u003eMyotis armiensis\u003c/em\u003e based on museum specimens.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCountry\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFederative Unit\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLocality\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCoordinates\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eAltitude\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCosta Rica\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePuntarenas\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMonteverde\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;16\u0026rsquo;N, 84\u0026deg;49\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e931 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCosta Rica\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHeredia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eParque Nacional Braulio Carrillo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;09\u0026rsquo;N, 83\u0026deg;59\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,000 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePanama\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChiriqu\u0026iacute;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBugaba, La Amistad International Park\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e08\u0026deg;53\u0026rsquo;N, 82\u0026deg;36\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2,214 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePanama\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChiriqu\u0026iacute;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTierras Altas, Cerro Punta\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e08\u0026deg;52\u0026rsquo;N, 82\u0026deg;35\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,858 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePanama\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChiriqu\u0026iacute;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTierras Altas, El Volc\u0026aacute;n\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e08\u0026deg;52\u0026rsquo;N, 82\u0026deg;44\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,280 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePanama\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCocl\u0026eacute;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLa Mesa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e08\u0026deg;38\u0026rsquo;N, 80\u0026deg;07\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e850 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVenezuela\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDistrito Capital\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePico Avila\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;33\u0026rsquo;N, 66\u0026deg;52\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2,092 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVenezuela\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMiranda\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCurupao\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;31\u0026rsquo;N, 66\u0026deg;38\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,160 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVenezuela\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMiranda\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSan Andr\u0026eacute;s\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;22\u0026rsquo;N, 66\u0026deg;50\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,180 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVenezuela\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAragua\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eEstaci\u0026oacute;n Biol\u0026oacute;gica Rancho Grande\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;21\u0026rsquo;N, 67\u0026deg;40\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,081 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVenezuela\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCarabobo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMontalb\u0026aacute;n\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10\u0026deg;15\u0026rsquo;N, 68\u0026deg;21\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,810 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eColombia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHuila\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSan Agust\u0026iacute;n, San Antonio village\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e01\u0026deg;57\u0026rsquo;N, 76\u0026deg;28\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2,185 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEcuador\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTungurahu\u0026aacute;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAzuay\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e01\u0026deg;20\u0026rsquo;S, 78\u0026deg;12\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,660 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEcuador\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNapo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eQuijos, Caba\u0026ntilde;as del Aliso\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e00\u0026deg;34\u0026rsquo;S, 77\u0026deg;53\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2,249 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEcuador\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMorona Santiago\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBa\u0026ntilde;os\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e02\u0026deg;14\u0026rsquo;S, 78\u0026deg;25\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3,115 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEcuador\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eZamora Chinchipe\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eYantzaza, Cpo. Minero Fruta del Norte\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e03\u0026deg;45\u0026rsquo;S, 78\u0026deg;31\u0026rsquo;W\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1,200 m\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eEcological niche models (ENMs) were generated using the ENMTML package in R (Andrade et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). We applied four algorithms to obtain more robust predictions: MaxEnt, SVM, BIOCLIM, and DOMAIN. To constrain prediction limits, we used the LTP method in conjunction with MAX_TS, which defines the threshold where the sum of sensitivity and specificity is maximized. Model performance was evaluated by splitting the occurrences into two subsets: 70% for training (model calibration) and 30% for testing (model evaluation). Four environmental suitability models were generated, one for each algorithm.\u003c/p\u003e\u003cp\u003eModel accuracy was assessed using the True Skill Statistic (TSS), with values above 0.5 considered reliable (Dud\u0026iacute;k et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). An ensemble model was then built by averaging the best-performing model from each algorithm, based on TSS values, resulting in a single map of environmental suitability for \u003cem\u003eM. armiensis\u003c/em\u003e. To convert the continuous suitability model into a binary presence\u0026ndash;absence map, we applied the Maximum Sensitivity plus Specificity (MSS) threshold.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e\u003ch2\u003eNew records to Northern South America\u003c/h2\u003e\u003cp\u003eThe museums\u0026rsquo; samples revision allowed the identification of 13 specimens of \u003cem\u003eM. armiensis\u003c/em\u003e from Venezuela (USNM 370890, 370893, 370901, 370929, 373920, 387714, 387715) and one specimen from Colombia (FMNH 72175), being the first records of this species for both countries (see Appendix 2). Records for Venezuela were made throughout the Cordillera de la Costa, in Rancho Grande Biological Station, Aragua (10\u0026deg;21'N, 67\u0026deg;40'W); Montalb\u0026aacute;n, Carabobo (10\u0026deg;15'N, 68\u0026deg;21'W); Pico Avila, Distrito Capital (10\u0026deg;32'N, 66\u0026deg;52'W); Curapao, Guarenas, Miranda (10\u0026deg;31'N, 66\u0026deg;38'W); and San Andr\u0026eacute;s, Miranda (10\u0026deg;22'N, 66\u0026deg;50'W), at altitudes ranging from 1,080 to 2,100 m. These specimens come from mountainous deciduous rainforests included within the Central Cordillera ecoregion. The record for Colombia was based on a specimen captured in San Agust\u0026iacute;n, Huila (01\u0026deg;52'N, 76\u0026deg;17'W), at 2,185 m above sea level. This record was made in transition between moist and dry forests in the Colombian Northwestern Andean ecoregion.\u003c/p\u003e\u003cp\u003eThe specimens from Venezuela and Colombia presents all diagnostics characters described by Carri\u0026oacute;n-Bonilla and Cook (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The skull is comparatively large and robust (GLS 12.9\u0026ndash;13.7), with sagittal and lambdoidal crests present. The shape of braincase roof inclined forward; posterior region of the skull being rounded and projected beyond the limit of the occipital condyles (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). All specimens have long and woolly fur; dorsal hairs are unicolored and range from medium brown to bright reddish brown; ventral hairs are bicolored and with clear contrast, with dark brown base and the tips ranging from light brown to light yellow (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). All specimens have a dense fur on the dorsal surface of the uropatagium that extends beyond the level of the knees, covering most of the tibia and reaching up to 2/3 of the basal portion of the uropatagium (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Confirmation of the taxonomic identity of this sample was made from the cyt-b sequencing of a specimen from Venezuela (USNM 370929), and the results are described below.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eGenetic structure and variation\u003c/h2\u003e\u003cp\u003eThe cyt-b dataset showed considerable variation, with 417 variable sites, 324 of which were parsimony-informative. \u003cem\u003eMyotis armiensis\u003c/em\u003e was recovered within a clade composed exclusively of species from Mexico, Central America and northern South America. It formed a monophyletic lineage and was recovered as the sister taxon to \u003cem\u003eM. pilosatibialis\u003c/em\u003e, a relationship strongly supported by high nodal values (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Phylogenetic inference further indicated that \u003cem\u003eM. armiensis\u003c/em\u003e comprises two geographically structured clades: one represented by samples from Central America (Panama) and the other by samples from northern South America (Ecuador and Venezuela).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eGenetic distance estimates under the K2p model indicated an average divergence of 6% between \u003cem\u003eM. armiensis\u003c/em\u003e and its sister species, \u003cem\u003eM. pilosatibialis\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Distances between \u003cem\u003eM. armiensis\u003c/em\u003e and other species of the \u003cem\u003eruber\u003c/em\u003e group ranged from 10% to 13%. In contrast, the divergence between Central and South American populations of \u003cem\u003eM. armiensis\u003c/em\u003e was 3%, while intrapopulation distances were approximately 1% in both geographic clades.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eAverage genetic distances (K2p) among \u003cem\u003eMyotis\u003c/em\u003e species of the \u003cem\u003eruber\u003c/em\u003e species group based on cyt-b sequences. The intraspecific variation is presented in the diagonal, boldface, and enclosed in square brackets.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"12\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTaxa\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c12\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1. \u003cem\u003eM. armiensis\u003c/em\u003e Central\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e[0.01]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e2. \u003cem\u003eM. armiensis\u003c/em\u003e South\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e[0.01]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3. \u003cem\u003eM. pilosatibialis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e[0.02]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e4. \u003cem\u003eM. extremus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e[0.02]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e5. \u003cem\u003eM. keaysi\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e[0.02]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e6. \u003cem\u003eM. ruber\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003e[0.00]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e7. \u003cem\u003eM. moratellii\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003e[0.00]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e8. \u003cem\u003eM. simus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e\u003cb\u003e[***]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e9. \u003cem\u003eM. midastactus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e\u003cb\u003e[***]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e10. \u003cem\u003eM. elegans\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e\u003cb\u003e[0.00]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e11. \u003cem\u003eM. riparius\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e\u003cp\u003e\u003cb\u003e[0.05]\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eMorphological variation\u003c/h3\u003e\n\u003cp\u003eThe MANOVA results recovered statistically significant morphometric differences between Central and South American samples of \u003cem\u003eM. armiensis\u003c/em\u003e (Pillai \u003cem\u003eF\u003c/em\u003e\u003csub\u003e\u003cem\u003e72\u003c/em\u003e\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.680, P\u0026thinsp;=\u0026thinsp;0.026; Wilks \u003cem\u003eF\u003c/em\u003e\u003csub\u003e\u003cem\u003e72\u003c/em\u003e\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.319, P\u0026thinsp;=\u0026thinsp;0.026). Pairwise post hoc tests showed that only the interorbital breadth (IOB) exhibited significant differences between Central and South American samples (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.01), while MAN and GLS showed marginal values (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.052 and 0.051, respectively; Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eFor PCA analysis, only the first PC was retained based on BS test, accounting for 57% of the total craniometric variance, while the PC2 accounted for 12% of total variation (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Considering the positive relationship among the original measurements and the PC1, which are all positively correlated, size seems to be the biological factor responsible for this amount of craniometric variation. All samples are highly overlap considering the projection of individual scores on the first two PCs, denoting the overall craniometric similarity among \u003cem\u003eM. armiensis\u003c/em\u003e and \u003cem\u003eM. pilosatibialis\u003c/em\u003e samples (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eEigenvalues, eigenvectors, and percentage of variance for the first two principal components (PCA), and the first three canonical variates (CVA), obtained for the two samples of Myotis armiensis and three samples of Myotis pilosatibialis. Eigenv. = eigenvalues; var. = variance.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMeasurements\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePC1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePC2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCV1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCV2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eCV3\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMAL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.267\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.232\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMAN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.236\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.270\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-0.540\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-5.629\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.584\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGLS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.215\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.206\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-2.078\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-3.094\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-3.057\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCIL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.216\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.188\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.337\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.405\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-1.309\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMAB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.247\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.147\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-3.676\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.499\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5.795\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBCB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.241\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.187\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-1.332\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.260\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-4.749\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eIOB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.477\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.389\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-5.602\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-6.678\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.693\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePOB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.245\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.616\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e13.720\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-1.289\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.590\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBAC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.358\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.069\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-2.168\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.566\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.479\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBAM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.321\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.048\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-1.506\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-1.878\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-4.355\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMTL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.249\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.285\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM1M3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.280\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e-0.343\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10.0788\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.387\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEigenv.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.004\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.877\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.0409\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2.263\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e% var.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e57\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e12.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e42.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e26.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eIn the CVA, the first three CVs account for 88% of the total variance, with the CV1 corresponding to 42% and the CV2 corresponding to 26% (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Despite the overall similarity observed in the PCA, the samples were recovered structured considering both the species and, in the case of \u003cem\u003eM. armiensis\u003c/em\u003e, the geographic clades. The interpolation of individual scores on the CV1 recovered the difference between species, with most of the individuals of \u003cem\u003eM. pilosatibialis\u003c/em\u003e with positive scores, while all but one individual of \u003cem\u003eM. armiensis\u003c/em\u003e exhibiting negative scores. Among \u003cem\u003eM. pilosatibialis\u003c/em\u003e, individuals from Mexico and Guatemala are mostly overlapped with scores between \u0026minus;\u0026thinsp;1 and 1, while individuals from Honduras and El Salvador exhibit scores mostly between 1 and 2. Most of the original measurements were negatively correlated with the CV1 except for the post-orbital breath (POB), which exhibited a positive correlation with CV1. It suggests that the difference between \u003cem\u003eM. pilosatibialis\u003c/em\u003e and \u003cem\u003eM. armiensis\u003c/em\u003e is mostly explained by general size, with the former being smaller than the latter, with individuals from Honduras and El Salvador being markedly small, but the post-orbital region is wider in \u003cem\u003eM. pilosatibialis\u003c/em\u003e and narrow in \u003cem\u003eM. armiensis\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe CV2 accounted for the differences between the two clades of \u003cem\u003eM. armiensis\u003c/em\u003e, with the South American samples exhibiting almost all individuals with positive scores whereas the Central American group exhibited mostly negative scores. Recovering the observed in the MANOVA results, the differences between these groups are mostly explained by the interorbital constriction breadth (IOB), that exhibited negative correlations around \u0026minus;\u0026thinsp;0.2, along with mandible toothrow length (MAN) and greatest length of skull (GLS), which exhibited negative correlations slightly lower than \u0026minus;\u0026thinsp;0.2. On the other hand, braincase breadth (BCB) exhibited positive correlation with CV2, around 0.2. Considering this, South American individuals exhibit short and wide skulls with short mandible toothrow and narrow interorbital region, while Central American individuals exhibit long and narrow skulls with long mandible toothrow and wide interorbital region. Considering South and Central American samples, posterior classification recovered most of the individuals belonging to the given group or mistakenly assigned to one of \u003cem\u003eM. pilosatibialis\u003c/em\u003e groups, respectively 56% and 25% for South American group and 54% and 15% for Central American group. Thus, less than 20% (3 individuals) of the South American group were suggested as member of the Central American group, while 31% (4 individuals) of the Central American group were suggested as belonging to the South American group (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e). Measurements these two groups are available in Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSelected measurements (mm) of \u003cem\u003eMyotis armiensis\u003c/em\u003e populations. Descriptive statistics include the mean, range (in parentheses), and sample size. See Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e for variable abbreviations. Mann-Whitney Test p-values was used to compare cranial measurements between samples and values statistically significant are bolded. Measurements with hyphen (\u0026ndash;) were not tested.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"3\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eMeasurements\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCentral America\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSouth America\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003emean (max\u003c/b\u003e\u0026ndash;\u003cb\u003emin)\u003c/b\u003e \u003cb\u003eN\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003emean (max\u003c/b\u003e\u0026ndash;\u003cb\u003emin)\u003c/b\u003e \u003cb\u003eN\u003c/b\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eFA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e37.5 (35.8\u0026ndash;38.7) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e37.0 (35.9\u0026ndash;38.4) 15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3MC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e34.0 (32.8\u0026ndash;35.2) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e34.3 (33.4\u0026ndash;35.8) 15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e12.0 (12.0\u0026ndash;13.0) 10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e12.5 (11.0\u0026ndash;14.0) 12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLDF\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6.5 (6.0\u0026ndash;7.0) 4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7.0 (6.5\u0026ndash;7.5) 8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLVF\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5.2 (5.0\u0026ndash;5.5) 4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5.8 (5.5\u0026ndash;6.0) 8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGLS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e13.46 (13.25\u0026ndash;13.69) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e13.34 (12.93\u0026ndash;13.73) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCCL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e12.14 (11.95\u0026ndash;12.39) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e11.99 (11.72\u0026ndash;12.50) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCBL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e12.74 (12.57\u0026ndash;13.02) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e12.62 (12.32\u0026ndash;12.98) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCIL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e12.94 (12.69\u0026ndash;13.20) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e12.84 (12.54\u0026ndash;13.24) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBAL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e11.62 (11.48\u0026ndash;11.82) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e11.46 (10.07\u0026ndash;11.85) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eZYG\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8.83 (8.67\u0026ndash;9.10) 7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e8.42 (8.06\u0026ndash;8.83) 10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMAB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e7.10 (6.83\u0026ndash;7.45) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7.07 (6.82\u0026ndash;7.42) 15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBCB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6.34 (6.21\u0026ndash;6.56 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6.39 (6.18\u0026ndash;6.65) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePOB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3.36 (3.16\u0026ndash;3.65) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3.33 (3.21\u0026ndash;3.49) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eIOB\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e4.53 (4.26\u0026ndash;4.81) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4.35 (4.09\u0026ndash;4.77) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBAC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3.66 (3.52\u0026ndash;3.82) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3.66 (3.47\u0026ndash;3.99) 15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBAM\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5.48 (5.31\u0026ndash;5.76) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5.41 (5.51\u0026ndash;5.82) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMTL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5.15 (5.04\u0026ndash;5.26) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5.11 (4.92\u0026ndash;5.35) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM1\u0026ndash;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2.92 (2.81\u0026ndash;3.03) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2.91 (2.82\u0026ndash;3.04) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMAL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e9.87 (9.96\u0026ndash;10.02) 11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9.82 (9.61\u0026ndash;10.27) 13\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMAN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5.54 (5.43\u0026ndash;5.73) 12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e5.46 (5.29\u0026ndash;5.67) 16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eAnother interesting pattern recovered is that geographically close samples tended to be distinctly different on cranial dimensions, with small or none overlapping among them, while geographically distant samples were recovered overlapped. For example, individuals of \u003cem\u003eM. pilosatibialis\u003c/em\u003e from Guatemala were partially overlapped with \u003cem\u003eM. armiensis\u003c/em\u003e form South American group, and individuals of \u003cem\u003eM. pilosatibialis\u003c/em\u003e from Mexico were partially overlapped with \u003cem\u003eM armiensis\u003c/em\u003e from Central and South American groups. Individuals of \u003cem\u003eM. pilosatibialis\u003c/em\u003e form Honduras and El Salvador were recovered as strikingly different from individuals of \u003cem\u003eM. armiensis\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eQualitative morphological analyses did not recover any pattern of distinction between OTUs from the characters. The sagittal crest is present in all specimens analyzed (\u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;28), ranging from low (50%) to median (50%) in samples from Central America (\u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;12); and superficial (6%), low (47%) or median (47%) in specimens from South America (\u003cem\u003eN\u003c/em\u003e\u0026thinsp;=\u0026thinsp;16). The same pattern was repeated for lambdoidal crests, which varied among superficial (10%), low (~\u0026thinsp;50%) or medium (~\u0026thinsp;40%) for both population groups. The shape of braincase roof inclined forward was the predominant state (92%) in the samples from South America; while specimens from Central America had a braincase roof inclined forward (66%) or straight (34%). The shape of the posterior region of the skull, being rounded and projected beyond the limit of the occipital condyles in 92% of the South American individuals; while samples from Central America had a greater variation of this character, being 11% flattened and not projected and 89% with a rounded shape and subtly projected beyond the posterior limit of the occipital condyles.\u003c/p\u003e\u003cp\u003eVirtually all specimens, regardless of geographical origin, have a dense fur on the dorsal surface of the uropatagium that extends beyond the level of the knees, covering most of the tibia and reaching up to 2/3 of the basal portion of the uropatagium. Only one individual from Ecuador (LACM 42951) had the fur extending to the knees. However, the condition of the skin may be biasing the observation. Specimens from the two OTUs showed the same color pattern, with unicolored dorsal fur ranging from medium brown to bright reddish brown. The ventral fur, on the other hand, is bicolored and with clear contrast, with dark brown base and the tips ranging from light brown to light yellow. Both populations of \u003cem\u003eM. armiensis\u003c/em\u003e have long and woolly fur, but specimens from South America have slightly longer fur than those from Central America (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eDistribution and ecological niche modelling\u003c/h3\u003e\n\u003cp\u003eBased on museum specimens, we found 16 occurrences localities for \u003cem\u003eM. armiensis\u003c/em\u003e through Costa Rica, Panama, Colombia, Ecuador, and Venezuela, in altitudes ranging from 931 to 3,115 m above sea level, comprising habitats of tropical montane forests (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Records in Costa Rica and Panama were made in Moist Montane Forests and swampy bogs, with presence oak trees (\u003cem\u003eQuercus\u003c/em\u003e spp.) and bamboo (\u003cem\u003eChusquea\u003c/em\u003e spp.) along Talamanca Cordillera. In Ecuador, Colombia, and Venezuela, \u003cem\u003eM. armiensis\u003c/em\u003e was recorded in Evergreen Lower Montane Forest, Cloud Montane Forest, and Montane Dry Forest associated with Andes and Cordillera de la Costa.\u003c/p\u003e\u003cp\u003eAll algorithms showed satisfactory TSS values (\u0026gt;\u0026thinsp;0.8). The ensemble approach predicted that \u003cem\u003eM. armiensis\u003c/em\u003e is highly associated with northern tropical Andes (and adjacent mountain ranges from Venezuela), occupying moist and dry montane forests from northern Venezuela to southern Peru (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). In fact, 94% of the suitable areas for \u003cem\u003eM. armiensis\u003c/em\u003e occupation is within the forested slopes of the Andes. Although there are no formal records of \u003cem\u003eM. armiensis\u003c/em\u003e occurrence in Peru, ENM predicts high suitable areas for a large part of Peruvian Andes, from Campanquiz to Vilcabamba Cordilleras. Small areas of high suitability are also present in the mountainous areas of Costa Rica and Panama in the Talamanca Cordillera. However, the model indicates that there are no suitable areas that connect the populations of Central and South America. Areas of high suitability on the Caribbean islands, in tepuis mountain range of Venezuela and Brazil, and in the highlands forests from Northeastern Brazil can be interpreted as commission errors, as although have adequate environmental conditions for \u003cem\u003eM. armiensis\u003c/em\u003e occurrence, are geographically isolated and, therefore, inaccessible to this species.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe most important variables to predict the geographic distribution of \u003cem\u003eM. armiensis\u003c/em\u003e were BIO10 (mean temperature of warmest quarter, 81%), BIO16 (precipitation of wettest quarter, 11%), and BIO4 (temperature seasonality, 2%). In general, the species occurs in areas that have high precipitation and low temperature variation.\u003c/p\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eTaxonomy\u003c/h2\u003e\u003cp\u003e\u003cb\u003eMyotis armiensis montensis\u003c/b\u003e, new subspecies\u003c/p\u003e\u003cp\u003eHolotype: The specimen (USNM 370890) consists of skin, skull and mandible of an adult male collected during Smithsonian Venezuelan Project on 20 August 1965 (Figs.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The specimen is deposited in the mammal collection of the National Museum of Natural History (USNM, Washington, D.C., USA). The Cyt-b sequence is available in the National Center for Biological Information\u0026rsquo;s GenBank under accession number MZ345121.\u003c/p\u003e\u003cp\u003eType locality: Pico Avila, 5 km NNE Caracas, near Hotel Humboldt, Distrito Capital, Venezuela (10\u0026deg;32'N, 66\u0026deg;52'W, elevation 2,092 m).\u003c/p\u003e\u003cp\u003eOther specimens: Another 17 specimens from Colombia, Ecuador, and Venezuela were examined and are listed in Appendix 2.\u003c/p\u003e\u003cp\u003eDistribution. Occurs along Cordillera de la Costa and Andean humid forests from northern Venezuela to southern Ecuador, and it is likely to occur also in Peru, although have no official records. The core of its distribution is strongly associated with Andean evergreen montane forest, cloud montane forest, and montane dry forest, at altitudes ranging from 1,080 and 3,120 m a.s.l.\u003c/p\u003e\u003cp\u003eEtymology: The subspecific epithet \u0026ldquo;\u003cem\u003emontensis\u003c/em\u003e\u0026rdquo; refers to its distribution endemic to tropical forests along northern South American Mountain ranges.\u003c/p\u003e\u003cp\u003eDiagnosis: Both populations of \u003cem\u003eM. armiensis\u003c/em\u003e are morphologicaly quite similar, but the distinction of \u003cem\u003eM. a. montensis\u003c/em\u003e in relation to \u003cem\u003eM. a. armiensis\u003c/em\u003e can be made from the following set of characters: longer fur (LDF\u0026thinsp;\u0026lt;\u0026thinsp;6.5, LVF\u0026thinsp;\u0026lt;\u0026thinsp;5.5); short and wide skulls with short mandible toothrow and narrow interorbital region (GLS\u0026thinsp;\u0026gt;\u0026thinsp;13.7; IOB\u0026thinsp;\u0026gt;\u0026thinsp;4.7; MAN\u0026thinsp;\u0026gt;\u0026thinsp;5.6); and lower sagittal and lambdoidal crests. Additionally, \u003cem\u003eM. a. montensis\u003c/em\u003e can be readily distinguished from \u003cem\u003eM. a. armiensis\u003c/em\u003e based on cytochrome-b gene tree, being reciprocally monophyletic.\u003c/p\u003e\u003cp\u003eDescription and comparisons: A medium to large species within Neotropical \u003cem\u003eMyotis\u003c/em\u003e (FA 35.9\u0026ndash;38.4). Its fur texture, cranial morphology, and phylogenetic position agree with the pattern observed in species allocated to the \u003cem\u003eruber\u003c/em\u003e species group. Ears are Bone Brown and medium-sized (EL 11\u0026ndash;14 mm), reaching the portion of the rostrum between the eyes and nostrils when pushed forward. Tragus long and slender, with a broad base and narrower spear-shaped terminal half, almost straight anterior edge, and rounded tip. Membranes Bone Brown. The dorsal surface of the uropatagium is densely furred, with hairs that extends beyond the level of the knees, covering most of the tibia and reaching up to 2/3 of the basal portion of the uropatagium. There is no fringe of hairs along the trailing edge of the uropatagium. Plagiopatagium attached to the foot, at the level of the toes, by a broad band of membrane. Dorsal and ventral fur wooly and moderately long (LDH 6.5\u0026ndash;7.5 mm, LVH 5.5\u0026ndash;6.0 mm). Dorsal fur unicolored, ranging from medium brown to bright reddish brown. Ventral fur strongly bicolored, with dark brown base and the tips ranging from light brown to light yellow.\u003c/p\u003e\u003cp\u003eThe dental formula is I 2/3, C 1/1, PM 3/3, M 3/3 (2x)\u0026thinsp;=\u0026thinsp;38, typical of most species of \u003cem\u003eMyotis\u003c/em\u003e. Skull robust and large in length (GLS 12.9\u0026ndash;13.7), virtually identical to \u003cem\u003eM. a. armiensis\u003c/em\u003e in qualitative characters. Second upper premolar (P3) subtly displaced lingually, and smaller than first upper premolar (P2). First lower molar (m1) myotodont, with postcristid connecting hypoconid and entoconid (Menu \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1987\u003c/span\u003e). Braincase elongated (oval shape) with the roof inclined forward in mostly specimens; sagittal and lambdoidal crests present, ranging from low to moderate in high; occipital region slightly rounded and subtle projected beyond the limit of the occipital condyles; well-developed mastoid processes. Frontal bone slightly sloping (ca. 145\u0026deg; angle); rostrum comparatively short (ca. 2/5 of skull total length).\u003c/p\u003e\u003cp\u003e\u003cem\u003eMyotis a. montensis\u003c/em\u003e is quite similar to \u003cem\u003eM. a. armiensis\u003c/em\u003e in external and skull morphologies but vary at the frequency of characters and in size of some external and cranial measurements. Both geographic forms have reddish brown, long and woolly fur, but specimens from Andes have slightly longer fur than those from Central America (see Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). South American specimens from Andes (\u003cem\u003eM. a. montensis\u003c/em\u003e) have lower sagittal and lambdoid crests than those in Central America (\u003cem\u003eM. a. armiensis\u003c/em\u003e). South American individuals exhibit short and wide skulls with short mandible toothrow and narrow interorbital region, while Central American individuals exhibit long and narrow skulls with long mandible toothrow and wide interorbital region, with statistically significant differences (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cb\u003eMyotis armiensis armiensis\u003c/b\u003e Carri\u0026oacute;n-Bonilla \u0026amp; Cook, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e\u003c/p\u003e\u003cp\u003eHolotype: The specimen (MSB 262089) consists of skin, skull and skeleton of an adult male collected by J. A. Cook and collaborators on 20 March 2012. The specimen is deposited in the mammal collection of the Museum of Southwestern Biology, University of New Mexico (MSB, Albuquerque, USA), including tissues samples (NK 209314). The cyt-b sequence is available in the National Center for Biological Information\u0026rsquo;s GenBank under accession number MW025265.\u003c/p\u003e\u003cp\u003eType locality: La Amistad International Park Ranger Station, Bugaba District, Chiriqu\u0026iacute; Province, Panama (08\u0026deg;53\u0026rsquo;N, 82\u0026deg;36\u0026rsquo;W, elevation 2,214 m).\u003c/p\u003e\u003cp\u003eDistribution: Occurs in premontane and montane tropical forests along Talamanca Cordillera from Costa Rica and Panama, in altitudes between 850 and 2,300 m a.s.l.\u003c/p\u003e\u003cp\u003eDiagnosis: Can be distinguished from \u003cem\u003eM. a. montensis\u003c/em\u003e from the following set of characters: shorter fur (LDF\u0026thinsp;\u0026gt;\u0026thinsp;7.0, LVF\u0026thinsp;\u0026gt;\u0026thinsp;5.5); longer and narrow skulls with longer mandible toothrow and wider interorbital region (GLS\u0026thinsp;\u0026lt;\u0026thinsp;13.2; IOB\u0026thinsp;\u0026lt;\u0026thinsp;4.8; MAN\u0026thinsp;\u0026lt;\u0026thinsp;5.7); and higher sagittal and lambdoidal crests. Additionally, \u003cem\u003eM. a. armiensis\u003c/em\u003e can be readily distinguished from \u003cem\u003eM. a. montensis\u003c/em\u003e based on cytochrome-b gene tree, being reciprocally monophyletic.\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eVariation and taxonomic considerations\u003c/h2\u003e\u003cp\u003e\u003cem\u003eMyotis armiensis\u003c/em\u003e is a recently described species, and despite its unequivocal taxonomic validity, aspects of its variation and distribution remain poorly understood. The cyt-b phylogeny recovered two clades with strong biogeographic associations and an estimated genetic distance of 3.1%. Genetic distances between 2% and 11% in cyt-b may indicate either valid species or conspecific populations with high variation, warranting additional studies to clarify their specific status (Bradley and Baker \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). In fact, some recognized Neotropical \u003cem\u003eMyotis\u003c/em\u003e species show genetic distances equal to or even lower than 3% (Novaes et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e). Conversely, most Neotropical \u003cem\u003eMyotis\u003c/em\u003e species display intraspecific genetic distances below 2% based on cyt-b sequences (Novaes et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e; D\u0026iacute;az et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Thus, this comparatively high divergence between allopatric populations deserves further critical investigation.\u003c/p\u003e\u003cp\u003eAmong other factors, high divergences may result from isolation by distance or restricted gene flow (Slatkin \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e1987\u003c/span\u003e, \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1993\u003c/span\u003e), although this does not necessarily have direct implications for species delimitation and more data is required to taxonomic assessment (Johns and Avise \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Bradley and Baker \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Using a similar dataset, Carri\u0026oacute;n-Bonilla et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) applied three molecular species delimitation methods. One (mPTP) recovered the Central American (Talamanca Cordillera) and Andean populations as distinct species, while the other two (GMYC, ABGD) considered them conspecific. Despite this, those authors recognized \u003cem\u003eM. armiensis\u003c/em\u003e as a monotypic taxon. In contrast, our morphometric analyses revealed differences in some cranial characters, though these were not corroborated by qualitative morphological traits. Ecological niche modeling suggested no suitable areas connecting the Andean and Talamanca populations, possibly reflecting strong geographic isolation and a consequent interruption in gene flow. Taken together, the phylogenetic structure, moderately high genetic divergence, and subtle size differences may reflect persistent geographic isolation and fixation of characters (Baker and Bradley \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Although these findings meet the criteria for considering the populations as divergent evolutionary lineages (De Queiroz \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Padial et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), the absence of morphometric discontinuities and reliable morphological or molecular synapomorphies led us to adopt a conservative approach, recognizing the trans- and cis-Andean populations as distinct allopatric subspecies. Nonetheless, we emphasize that broader taxonomic sampling and sequencing of additional specimens (and genes) could refine these results and potentially support the elevation of \u003cem\u003eM. a. montensis\u003c/em\u003e to full species.\u003c/p\u003e\u003cp\u003eBased on biological and phylogenetic species concepts as a framework, Avise and Ball (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1990\u003c/span\u003e) defined subspecies as groups of populations that truly or potentially interbreed, are phylogenetically distinguishable, but remain reproductively compatible with other groups of the same species. These authors also emphasized that subspecies recognition should be based on multiple independent lines of evidence consistent with taxon distribution. However, direct evidence of interbreeding or reproductive compatibility is often difficult to obtain. Zachos (\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) argued that subspecies should be recognized when patterns of genetic or morphological variation are present, even if complete population structuring is lacking, and highlighted that the concept is inherently arbitrary. In such cases, disjunct populations with genetic, morphological, and/or ecological differentiation may be classified as subspecies, thereby organizing taxonomic and evolutionary knowledge (Zachos et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). More recently, De Queiroz (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) proposed a revised concept of subspecies, defining them as incompletely separated lineages. In other words, subspecies are entities of the same fundamental kind as species (i.e., separately evolving metapopulation lineages), differing only in that they remain incompletely separated and thus belong to a more inclusive lineage, the species (De Queiroz \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe case of \u003cem\u003eM. armiensis\u003c/em\u003e fits well with the subspecies definitions proposed by Zachos (\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and De Queiroz (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), since the morphological and morphometric variation, coupled with disjunct distribution, indicates incompletely separated lineages. This supports our classification of \u003cem\u003eM. a. armiensis\u003c/em\u003e for Talamanca populations and \u003cem\u003eM. a. montensis\u003c/em\u003e for Andean populations.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eEvolutionary history\u003c/h2\u003e\u003cp\u003eA recent study estimated the divergence between \u003cem\u003eM. armiensis\u003c/em\u003e and its sister species, \u003cem\u003eM. pilosatibialis\u003c/em\u003e, at approximately 2.25 Ma (\u0026plusmn;\u0026thinsp;0.45 Ma) during the Early Pleistocene (Novaes et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e). Although the split between the Talamanca and Andean lineages of \u003cem\u003eM. armiensis\u003c/em\u003e has not been dated, it is reasonable to infer a more recent divergence, likely in the Middle Pleistocene. \u003cem\u003eMyotis armiensis\u003c/em\u003e belongs to a clade composed exclusively of species from Mexico and Central America, being one of the most recently diverged members of this clade (Carri\u0026oacute;n-Bonilla and Cook \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Novaes et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e). Considering its phylogenetic position, it is plausible that its origin lies in Mesoamerica, with the ancestral lineage adapted to cooler environments than currently lowlands, as supported by ecological niche models. The colonization of South America likely occurred via dispersal across the Isthmus of Panama during climatic periods favorable to population expansion.\u003c/p\u003e\u003cp\u003eThe Pleistocene was characterized by glacial (cold, dry) and interglacial (warm, wet) cycles (Ditlevsen \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Herbert \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). During glaciations, glaciers advanced and high-elevation ecosystems (paramos, cloud forests, and montane forests) shifted downslope, expanding habitats now confined to highlands and forming mosaics of tropical forests and open savannas (Van Der Hammen \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1974\u003c/span\u003e; Prance \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Loehle 2007; Cabanne et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). In contrast, interglacial periods brought glacial retreat, rising temperatures, and upslope displacement of these ecosystems, accompanied by expansion of ombrophilous and seasonal forests in lowlands (Van Der Hammen \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1974\u003c/span\u003e; Prance \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Loehle 2007; Cabanne et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). These dynamics promoted cycles of fragmentation and reconnection of populations, generating opportunities for isolation and diversification in many taxa, including \u003cem\u003eMyotis\u003c/em\u003e (Carnaval and Moritz \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Goin et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Cabanne et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Novaes et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). Thus, during glacial phases, \u003cem\u003eM. armiensis\u003c/em\u003e likely expanded into lower altitudes, increasing population size, colonizing South America, and occupying a wide range of suitable forests. Conversely, during interglacial phases, as cloud and montane forests contracted upslope, \u003cem\u003eM. armiensis\u003c/em\u003e populations retreated to cooler and more stable highland habitats, where they remain today. Moreover, major mountain ranges may have acted as geographic barriers even during climatically stable intervals, promoting speciation and shaping evolutionary trajectories (Vuilleumier \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e1969\u003c/span\u003e; Cadle \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Patterson et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Torres-Carvajal and Mafla-Endara \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe genetic structure revealed by our analyses, along with shifts in cranial character frequencies, may reflect recent population bottlenecks driven by Pleistocene environmental fluctuations, as documented for other \u003cem\u003eMyotis\u003c/em\u003e species (Furman et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Lu et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Novaes et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). These events may have caused population declines, reduced gene flow, and increased the frequency of rare haplotypes within populations (Willis and Wiese \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Hartl 2008). Although the evolutionary history of \u003cem\u003eM. armiensis\u003c/em\u003e requires further study with broader genetic sampling, current spatial segregation patterns are sufficient to support recognition of two subspecies, consistent with the arguments of Zachos (\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and De Queiroz (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Future phylogeographic research based on larger datasets and additional genetic markers will be essential to clarify demographic dynamics and diversification processes underlying intraspecific variation.\u003c/p\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are grateful for staff and curators of the institutions that received us and Edson Fiedler de Abreu for helping in generating molecular data. RLMN has received 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 has received financial support from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil; 313963/2018-5) and FAPERJ (E-26/200.967/2021).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eRLMN, study conception and design, project management, data analysis, writing and review; MAGS, data analysis, writing and review; AC, data analysis, writing and review; DEW, advising and review; MW, advising and review; RM, project management, advising and review. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFundação 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) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil; 313963/2018-5).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eDNA sequences generated in this study have been deposited in GenBank (Appendix I). Data from distribution is available in Table 2. Morphological data matrices are available upon request from the corresponding author.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflict of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics declaration\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAndrade AFA, Velazco SJE, De Marco PJ (2020) ENMTML: An R package for a straightforward construction of complex ecological niche models. Environ Model Soft 125:104615. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.envsoft.2019.104615\u003c/span\u003e\u003cspan address=\"10.1016/j.envsoft.2019.104615\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eAntonelli A (2021) The rise and fall of Neotropical biodiversity. Bot J Lin Soc 199:8\u0026ndash;24. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/botlinnean/boab061\u003c/span\u003e\u003cspan address=\"10.1093/botlinnean/boab061\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eAvise JC, Ball RM (1990) Principles of genealogical concordance in species concepts and biological taxonomy. In: Futuyma D, Antonovics J (eds) Oxford surveys in evolutionary biology. Oxford University Press, Oxford, pp 45\u0026ndash;67\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBaker RJ, Bradley RD (2006) Speciation in mammals and the genetic species concept. J Mammal 87:643\u0026ndash;662. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1644/06-mamm-f-038r2.1\u003c/span\u003e\u003cspan address=\"10.1644/06-mamm-f-038r2.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBradley RD, Baker RJ (2001) A test of the genetic species concept: cytochrome-b sequences and mammals. J Mammal 82: 960\u0026ndash;973. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1644/1545-1542\u003c/span\u003e\u003cspan address=\"10.1644/1545-1542\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e(2001)082%3C0960:ATOTGS%3E2.0.CO;2\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eBrumfield RT, Capparella AP (1996) Historical diversification of birds in northwestern South America: a molecular perspective on the role of vicariant events. Evolution 50:1607\u0026ndash;1624. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1558-5646.1996.tb03933.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1558-5646.1996.tb03933.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCabanne GS, Calder\u0026oacute;n L, Trujillo-Arias N, Flores P, Pessoa R, d\u0026rsquo;Horta FM, Miyaki CY (2016) Effects of Pleistocene climate changes on species ranges and evolutionary processes in the Neotropical Atlantic Forest. Biol J Lin Soc 119:856\u0026ndash;872. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/bij.12844\u003c/span\u003e\u003cspan address=\"10.1111/bij.12844\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCadle JE (1991) Systematics of lizards of the genus \u003cem\u003eStenocercus\u003c/em\u003e (Iguania: Tropiduridae) from northern Per\u0026uacute;: new species and comments on relationships and distribution patterns. Proc Acad Nat Sci Phil 143:1\u0026ndash;96\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCaraballo DA, Montani ME, Martınez LM, Antoniazzi LR, Sambrana TC, Fernandez C, Cisterna DM, Beltr\u0026aacute;n FJ, Colombo VC (2020) Heterogeneous taxonomic resolution of cytochrome b gene identification of bats from Argentina: implications for field studies. PLoS ONE 15:e0244750. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal. pone.0244750\u003c/span\u003e\u003cspan address=\"10.1371/journal. pone.0244750\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCarnaval AC, Moritz C (2008) Historical climate modelling predicts patterns of current biodiversity in the Brazilian Atlantic Forest. J Biogeogr 35:1187\u0026ndash;1201. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-2699.2007.01870.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-2699.2007.01870.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCarri\u0026oacute;n-Bonilla CA, Cook JA (2020) A new bat species of the genus \u003cem\u003eMyotis\u003c/em\u003e with comments on the phylogenetic placement of \u003cem\u003eM. keaysi\u003c/em\u003e and \u003cem\u003eM. pilosatibialis\u003c/em\u003e. Therya 11:508\u0026ndash;532. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.12933/therya-20-999\u003c/span\u003e\u003cspan address=\"10.12933/therya-20-999\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eCarri\u0026oacute;n-Bonilla CA, Ron S, Cook JA (2024) Species richness in \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera: Vespertilionidae): species delimitation and expanded geographic sampling reveal high Neotropical diversity. J Mammal 105:241\u0026ndash;258. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jmammal/gyad124\u003c/span\u003e\u003cspan address=\"10.1093/jmammal/gyad124\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eClaridge MF (2010) Species are real biological entities. In: Ayala FK, Arp R (eds) Contemporary debates in philosophy of biology. Wiley-Blackwell, Malden, pp 91\u0026ndash;109\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDarriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nat Met 9:a772. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/nmeth.2109\u003c/span\u003e\u003cspan address=\"10.1038/nmeth.2109\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDe Queiroz K (2007) Species concepts and species delimitation. Syst Biol 56:879\u0026ndash;886. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/10635150701701083\u003c/span\u003e\u003cspan address=\"10.1080/10635150701701083\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDe Queiroz K (2020) An updated concept of subspecies resolves a dispute about the taxonomy of incompletely separated lineages. Herpetol Rev 51(2):459\u0026ndash;461\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDe Queiroz K (2021) Response to critiscism of na updated subspecies concept. Herpetol Rev 52(4):773\u0026ndash;776\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eD\u0026iacute;az DR, Medina CR, Arias S, L\u0026oacute;pez E, Carri\u0026oacute;n-Bonilla CA (2025) A new species of \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera: Vespertilionidae) from the coastal desert of southern Peru. J Mammal gyaf028. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jmammal/gyaf028\u003c/span\u003e\u003cspan address=\"10.1093/jmammal/gyaf028\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDitlevsen P (2022) The Pleistocene glacial cycles and millennial-scale climate variability. Atmos Ocean 60:233\u0026ndash;244. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/07055900.2022.2077172\u003c/span\u003e\u003cspan address=\"10.1080/07055900.2022.2077172\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eDud\u0026iacute;k M, Phillips SJ, Schapire RE (2004) Performance guarantees for regularized maximum entropy density estimation. In: Shawe-Taylor J, Singer Y (eds) Conference on Computational Learning Theory (COLT). Lecture Notes in Computer Science, vol. 3120. Springer, Berlin. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/978-3-540-27819-1_33\u003c/span\u003e\u003cspan address=\"10.1007/978-3-540-27819-1_33\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eEdgar RC (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucl Acid Res 32(5):1792\u0026ndash;1797. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/nar/gkh340\u003c/span\u003e\u003cspan address=\"10.1093/nar/gkh340\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFrontier S (1976) \u0026Eacute;tude de la decroissance des valeurs propres dans une analyze en composantes principales: comparison avec le mod\u0026egrave;le de baton bris\u0026eacute;. J Exp Mar Biol Ecol 25(1):67\u0026ndash;75. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/0022-0981(76)90076-9\u003c/span\u003e\u003cspan address=\"10.1016/0022-0981(76)90076-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eFurman A, \u0026Ccedil;oraman E, Nagy ZL, Postawa T, Bilgin R, Gajewska M, Bogdanowicz W (2013) Phylogeography of the large \u003cem\u003eMyotis\u003c/em\u003e bats (Chiroptera: Vespertilionidae) in Europe, Asia Minor, and Transcaucasia. Biol J Lin Soc 108:189\u0026ndash;209. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1095-8312.2012.01994.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1095-8312.2012.01994.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGhiselin MT (1997) Metaphysics and the origin of species. State University of New York, New York, p 377\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eGoin FJ, Gelfo JN, Chornogubsky L, Woodburne MO, Martin T (2012) Origins, radiations, and distribution of South American mammals: From greenhouse to icehouse worlds. In: Patterson BD, Costa LP (eds) Bones, clones, and biomes: the history and geography of recent Neotropical mammals. University of Chicago Press, Chicago, pp 20\u0026ndash;50. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7208/9780226649214-003\u003c/span\u003e\u003cspan address=\"10.7208/9780226649214-003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHammer \u0026Oslash;, Harper DAT, Ryan PD (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontol Elect 4(1):1\u0026ndash;9\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHartl DL (2006) Principles of population genetics, 4th edn. OUP, New York, p 545\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHasegawa M, Kishino H, Yano T (1985) Dating of human-ape splitting by a molecular clock of mitochondrial DNA. J Mol Evol 22:160\u0026ndash;174. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/bf02101694\u003c/span\u003e\u003cspan address=\"10.1007/bf02101694\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHerbert TD (2023) The Mid-Pleistocene climate transition. Rev Earth Plan Sci 51:389\u0026ndash;418. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1146/annurev-earth-032320-104209\u003c/span\u003e\u003cspan address=\"10.1146/annurev-earth-032320-104209\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHonaker J, King G, Blackwell M (2011) Amelia II: A program for missing data. J Stat Softw 45(1):1\u0026ndash;47. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.18637/jss.v045.i07\u003c/span\u003e\u003cspan address=\"10.18637/jss.v045.i07\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHull D (1994) Contemporary systematic philosophies. In: Sober E (ed) Conceptual issues in evolutionary biology, 2nd edn. MIT Press, Cambridge, pp 295\u0026ndash;330\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eHutter CR, Lambert SM, Wiens JJ (2017) Rapid diversification and time explain amphibian richness at different scales in the tropical Andes, Earth\u0026rsquo;s most biodiverse hotspot. Ame Nat 190:828\u0026ndash;843. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1086/694319\u003c/span\u003e\u003cspan address=\"10.1086/694319\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eJohns GC, Avise JC (1998) A comparative summary of genetic distances in the vertebrates from the mitochondrial cytochrome b gene. Mol Biol Evol 15(11):1481\u0026ndash;1490. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/oxfordjournals.molbev.a025875\u003c/span\u003e\u003cspan address=\"10.1093/oxfordjournals.molbev.a025875\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. J Mol Evol 16:111\u0026ndash;120. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/bf01731581\u003c/span\u003e\u003cspan address=\"10.1007/bf01731581\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKovarovic K, Aiello LC, Cardini A, Lockwood CA (2011) Discriminant function analysis in archaeology: are classification rates too good to be true? J Archaeol Sci 38:3006\u0026ndash;3018. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jas.2011.06.028\u003c/span\u003e\u003cspan address=\"10.1016/j.jas.2011.06.028\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eKumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular evolutionary genetics analysis across computing platforms. Mol Biol Evol 35:1547\u0026ndash;1549. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/molbev/msy096\u003c/span\u003e\u003cspan address=\"10.1093/molbev/msy096\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLarsen RJ, Knapp MC, Genoways HH, Khan FAA, Larsen PA, Wilson DE, Baker RJ (2012a) Genetic diversity of Neotropical \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera: Vespertilionidae) with an emphasis on South American species. PLoS ONE 7:e46578. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0046578\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0046578\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLarsen RJ, Larsen PA, Genoways HH, Catzeflis FM, Geluso K, Kwiecinski KK, Pedersen SC, Simal F, Baker RJ (2012b) Evolutionary history of Caribbean species of \u003cem\u003eMyotis\u003c/em\u003e, with evidence of a third Lesser Antillean endemic. Mammal Biol 77:124\u0026ndash;134. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mambio.2011.11.003\u003c/span\u003e\u003cspan address=\"10.1016/j.mambio.2011.11.003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLoehle C (2006) Predicting Pleistocene climate from vegetation in North America. Clim Past 3:109\u0026ndash;118. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.5194/cp-3-109-2007\u003c/span\u003e\u003cspan address=\"10.5194/cp-3-109-2007\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eLu G, Lin A, Luo J, Blondel DV, Meiklejohn KA, Sun K, Feng J (2013) Phylogeography of the Rickett\u0026rsquo;s big-footed bat, \u003cem\u003eMyotis pilosus\u003c/em\u003e (Chiroptera: Vespertilionidae): a novel pattern of genetic structure of bats in China. BMC Evol Biol 13:e241. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/1471-2148-13-241\u003c/span\u003e\u003cspan address=\"10.1186/1471-2148-13-241\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMenu H (1987) Morphotypes dentaires actuels et fossiles des chiropteres vespertilionines, 2 partie: implications systematiques et phylogeniques. Paleovertebrata 17(1):77\u0026ndash;150\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMeseguer AS, Michel A, Fabre PH, P\u0026eacute;rez-Escobar AO, Chomicki G, Riina R, Antonelli A, Antoine PO, Delsuc F, Condamine FL (2022) Diversification dynamics in the Neotropics through time, clades, and biogeographic regions. eLife 11:e74503. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7554/eLife.74503\u003c/span\u003e\u003cspan address=\"10.7554/eLife.74503\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMitteroecker P, Bookstein F (2011) Linear discrimination, ordination, and the visualization of selection gradients in modern morphometrics. Evol Biol 38:100\u0026ndash;114. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11692-011-9109-8\u003c/span\u003e\u003cspan address=\"10.1007/s11692-011-9109-8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMoratelli R, Peracchi AL, Dias D, Oliveira JA (2011) Geographic variation in South American populations of \u003cem\u003eMyotis nigricans\u003c/em\u003e (Schinz, 1821) (Chiroptera, Vespertilionidae), with the description of two new species. Mammal Biol 76(5):592\u0026ndash;607. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mambio.2011.01.003\u003c/span\u003e\u003cspan address=\"10.1016/j.mambio.2011.01.003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eMoratelli R, Burgin C, Cl\u0026aacute;udio VC, Novaes RLM, L\u0026oacute;pez-Baucells A, Haslauer R (2019) Family Vespertilionidae (Vesper Bats). In: Wilson DE, Mittermeier AR (eds) Handbook of the mammals of the world, vol 9. Bats. Lynx Editions, Barcelona, pp 716\u0026ndash;981\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Cl\u0026aacute;udio 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 1015:145\u0026ndash;167. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3897/zookeys.1015.59248\u003c/span\u003e\u003cspan address=\"10.3897/zookeys.1015.59248\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Wilson DE, Moratelli R (2022a) Catalogue of primary types of Neotropical \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera, Vespertilionidae). ZooKeys 1105:127\u0026ndash;164. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3897/zookeys.1105.85055\u003c/span\u003e\u003cspan address=\"10.3897/zookeys.1105.85055\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Cl\u0026aacute;udio VC, Carri\u0026oacute;n-Bonilla C, Abreu EF, Wilson DE, Maldonado JE, Weksler M (2022b) Variation in the \u003cem\u003eMyotis keaysi\u003c/em\u003e complex (Chiroptera, Vespertilionidae), with description of a new species from Ecuador. J Mammal 103:540\u0026ndash;559. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jmammal/gyab139\u003c/span\u003e\u003cspan address=\"10.1093/jmammal/gyab139\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Rodr\u0026iacute;guez-San Pedro A, Saldarriaga-C\u0026oacute;rdoba MM, Aguilera-Acu\u0026ntilde;a O, Wilson DE, Moratelli R (2022c) Systematic review of \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera, Vespertilionidae) from Chile based on molecular, morphological, and bioacoustics data. Zootaxa 5188(5):430\u0026ndash;452. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.11646/zootaxa.5188.5.2\u003c/span\u003e\u003cspan address=\"10.11646/zootaxa.5188.5.2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Cl\u0026aacute;udio VC, Wilson DE, Weber MM, Weksler M, Moratelli R (2024a) Integrative taxonomy and evolutionary history reveal cryptic diversity in \u003cem\u003eMyotis riparius\u003c/em\u003e (Chiroptera: Vespertilionidae). Biol J Lin Soc 142:91\u0026ndash;112. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/biolinnean/blad123\u003c/span\u003e\u003cspan address=\"10.1093/biolinnean/blad123\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Cl\u0026aacute;udio VC, Wilson DE, Weksler M, Moratelli R (2024b) Taxonomic status of \u003cem\u003eMyotis extremus\u003c/em\u003e (Chiroptera, Vespertilionidae) from Mesoamerica, with comments on the distribution and systematics of \u003cem\u003eMyotis nigricans\u003c/em\u003e. Zool Anz 308:99\u0026ndash;110. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jcz.2023.11.009\u003c/span\u003e\u003cspan address=\"10.1016/j.jcz.2023.11.009\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Cl\u0026aacute;udio VC, Bertocchi NA, Oliveira K, Semedo TBF, Saldanha J, Wilson DE, Moratelli R (2025a) Unveiling the shelf life: a new cryptic species of \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera, Vespertilionidae) from South America revealed by an integrative taxonomy approach. J Mammal 106(4):878\u0026ndash;897. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jmammal/gyaf016\u003c/span\u003e\u003cspan address=\"10.1093/jmammal/gyaf016\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eNovaes RLM, Cl\u0026aacute;udio VC, Bertocchi NA, Abreu EF, Wilsom DE, Maldonado JE, Moratelli R (2025b) Islands in the stream: Distribution of \u003cem\u003eMyotis attenboroughi\u003c/em\u003e (Chiroptera, Vespertilionidae) in Grenada and mainland South America illuminates the evolutionary history of Caribbean \u003cem\u003eMyotis\u003c/em\u003e. Evol Syst 9:107\u0026ndash;122. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3897/evolsyst.9.146107\u003c/span\u003e\u003cspan address=\"10.3897/evolsyst.9.146107\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePadial JM, Miralles A, La Riva ID, Vences M (2010) The integrative future of taxonomy. Front Zool 7:e16. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/1742-9994-7-16\u003c/span\u003e\u003cspan address=\"10.1186/1742-9994-7-16\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePatterson BD, Pacheco V, Ashley MV (1992) On the origins of the western slope region of endemism: systematics of fig-eating bats, genus \u003cem\u003eArtibeus\u003c/em\u003e. Mem Mus Hist Nat Lima 21:189\u0026ndash;205\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePatterson BD, Solari S, Velazco PM (2012) The role of the Andes in the diversification and biogeography of Neotropical mammals. In: Patterson BD, Costa LP (eds) Bones, clones, and biomes: the history and geography of recent Neotropical mammals. University of Chicago Press, Chicago, pp 351\u0026ndash;378. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7208/9780226649214-015\u003c/span\u003e\u003cspan address=\"10.7208/9780226649214-015\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePatterson BD, Webala PW, Peterhans JCK, Goodman SM, Bartonjo M, Demos TC (2019) Genetic variation and relationships among Afrotropical species of \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera: Vespertilionidae). J Mammal 100:1130\u0026ndash;1143. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jmammal/gyz087\u003c/span\u003e\u003cspan address=\"10.1093/jmammal/gyz087\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePeres-Neto PR, Jackson DA, Somers KM (2005) How many principal components? Stopping rules for determining the number of non-trivial axes revisited. Comp Stat Data Anal 49:974\u0026ndash;997. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.csda.2004.06.015\u003c/span\u003e\u003cspan address=\"10.1016/j.csda.2004.06.015\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePlatt RN, 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 \u003cem\u003eMyotis\u003c/em\u003e bats. Syst Biol 67(2):236\u0026ndash;249. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/sysbio/syx070\u003c/span\u003e\u003cspan address=\"10.1093/sysbio/syx070\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003ePrance GT (1982) A review of the phytogeographic evidences for Pleistocene climate changes in the Neotropics. Miss Bot Gard 69(3):594\u0026ndash;624. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/2399085\u003c/span\u003e\u003cspan address=\"10.2307/2399085\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eRambaut A, Drummond AJ (2009) Tracer v.1.5: MCMC Trace Analysis Packpage. Available at \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://tree.bio.ed.ac.uk/software/tracer/\u003c/span\u003e\u003cspan address=\"http://tree.bio.ed.ac.uk/software/tracer/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eRonquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, H\u0026ouml;hna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 32: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst Biol 61:539\u0026ndash;542. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/sysbio/sys029\u003c/span\u003e\u003cspan address=\"10.1093/sysbio/sys029\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eRuedi M, Mayer F (2001) Molecular systematics of bats of the genus \u003cem\u003eMyotis\u003c/em\u003e (Vespertilionidae) suggest deterministic ecomorphological convergences. Mol Phylogen Evol 21:463\u0026ndash;448. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1006/mpev.2001.1017\u003c/span\u003e\u003cspan address=\"10.1006/mpev.2001.1017\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eRuedi M, Csorba G, Lin LK, Chou CH (2015) Molecular phylogeny and morphological revision of \u003cem\u003eMyotis\u003c/em\u003e bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920:301\u0026ndash;342. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.11646/zootaxa.3920.2.6\u003c/span\u003e\u003cspan address=\"10.11646/zootaxa.3920.2.6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSedlock JL, Heaney LR, Balete DS, Ruedi M (2020) Philippine bats of the genus \u003cem\u003eKerivoula\u003c/em\u003e (Chiroptera: Vespertilionidae): overview and assessment of variation in \u003cem\u003eK. pellucida\u003c/em\u003e and \u003cem\u003eK. whiteheadi\u003c/em\u003e. Zootaxa 4755: 454\u0026ndash;490. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.11646/zootaxa.4755.3.2\u003c/span\u003e\u003cspan address=\"10.11646/zootaxa.4755.3.2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSimmons NB, Cirranello AL (2025) Bat species of the world: A taxonomic and geographic database. Version 1.7. Available at \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://batnames.org/home.html\u003c/span\u003e\u003cspan address=\"https://batnames.org/home.html\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, Accessed on April 09, 2025\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSlatkin M (1987) Gene flow and the geographic structure of natural populations. Science 236:787\u0026ndash;792. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1126/science.3576198\u003c/span\u003e\u003cspan address=\"10.1126/science.3576198\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSlatkin M (1993) Isolation by distance in equilibrium and non-equilibrium populations. Evolution 47:264\u0026ndash;279. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1558-5646.1993.tb01215.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1558-5646.1993.tb01215.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eSol\u0026iacute;s A, Kohlmann B, Alvarado GE (2024) A Costa Rican biological puzzle\u0026mdash;The geological, landscape, and vicariance history as a conditioning of the past and present biogeography. In: Quesada-Rom\u0026aacute;n A (ed) Landscapes and landforms of Costa Rica. Springer, Cham. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/978-3-031-64940-0_10\u003c/span\u003e\u003cspan address=\"10.1007/978-3-031-64940-0_10\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eStadelmann B, Lin LK, Kunz TH, Ruedi M (2007) Molecular phylogeny of New World \u003cem\u003eMyotis\u003c/em\u003e (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes. Mol Phylogen Evol 43:32\u0026ndash;48. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ympev.2006.06.019\u003c/span\u003e\u003cspan address=\"10.1016/j.ympev.2006.06.019\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eTorres-Carvajal O, Mafla-Endara P (2013) Evolutionary history of Andean \u003cem\u003ePholidobolus\u003c/em\u003e and \u003cem\u003eMacropholidus\u003c/em\u003e (Squamata: Gymnophthalmidae) lizards. Mol Phylogenet Evol 68(2):212\u0026ndash;217. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ympev.2013.03.013\u003c/span\u003e\u003cspan address=\"10.1016/j.ympev.2013.03.013\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eVan Der Hammen T (1974) The Pleistocene changes of vegetation and climate in tropical South America. J Biogeog 1:3\u0026ndash;26. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/3038066\u003c/span\u003e\u003cspan address=\"10.2307/3038066\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eViale M, Garreaud R (2015) Orographic effects of the subtropical and extratropical Andes on upwind precipitating clouds. JGR Atmos 120:4962\u0026ndash;4974. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/2014JD023014\u003c/span\u003e\u003cspan address=\"10.1002/2014JD023014\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eVuilleumier F (1969) Pleistocene speciation in birds living in the high Andes. Nature 223:1179\u0026ndash;1180. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/2231179a0\u003c/span\u003e\u003cspan address=\"10.1038/2231179a0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eWillis K, Wiese RJ (1993) Effect of new founders on retention of gene diversity in captive populations: A formalization of the nucleus population concept. ZooBiology 12(6):535\u0026ndash;548. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/zoo.1430120605\u003c/span\u003e\u003cspan address=\"10.1002/zoo.1430120605\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eZachos FE (2016) Species concepts in biology: historical development, theoretical foundations and practical relevance. Springer International Publishing, Geneve, p 220. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/978-3-319-44966-1\u003c/span\u003e\u003cspan address=\"10.1007/978-3-319-44966-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eZachos FE, Mattioli S, Ferretti F, Lorenzini R (2014) The unique Mesola red deer of Italy: taxonomic recognition (\u003cem\u003eCervus elaphus italicus\u003c/em\u003e nova ssp., Cervidae) would endorse conservation. Ital J Zool 81:136\u0026ndash;143. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/11250003.2014.895060\u003c/span\u003e\u003cspan address=\"10.1080/11250003.2014.895060\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"mammalian-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mamb","sideBox":"Learn more about [Mammalian Biology](https://link.springer.com/journal/42991)","snPcode":"42991","submissionUrl":"https://www.editorialmanager.com/mamb/default2.aspx","title":"Mammalian Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Andes Mountain range, biogeography, Myotinae, Pleistocene, speciation, taxonomy.","lastPublishedDoi":"10.21203/rs.3.rs-7794885/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7794885/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cem\u003eMyotis armiensis\u003c/em\u003e was recently described from specimens collected in the mountain ranges of Costa Rica, Panama, and Ecuador. During a systematic revision of Neotropical \u003cem\u003eMyotis\u003c/em\u003e, we identified additional specimens from Venezuela and Colombia. Using an integrative approach that combined morphological traits, cytochrome b sequences, and ecological niche modeling, we assessed the geographic variation of \u003cem\u003eM. armiensis\u003c/em\u003e. Our findings indicate that this species is structured into two allopatric populations: one in the Talamanca Cordillera of Central America, and another restricted to the South American Andes. These populations are reciprocally monophyletic, exhibiting genetic distances greater than 3% and notable morphometric divergence. The results suggest that \u003cem\u003eM. armiensis\u003c/em\u003e comprises two incompletely isolated lineages, supporting the recognition of two subspecies described herein. The diversification of this taxon likely occurred during the Pleistocene, possibly driven by environmental fluctuations associated with glacial cycles.\u003c/p\u003e","manuscriptTitle":"On the variation, distribution and evolutionary history of Myotis armiensis (Chiroptera, Vespertilionidae) along Neotropical mountain ranges, including the description of new subspecies","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-29 12:33:24","doi":"10.21203/rs.3.rs-7794885/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-10-15T07:40:32+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-10-15T06:46:53+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-10-10T05:19:30+00:00","index":"","fulltext":""},{"type":"submitted","content":"Mammalian Biology","date":"2025-10-08T08:30:47+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"mammalian-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mamb","sideBox":"Learn more about [Mammalian Biology](https://link.springer.com/journal/42991)","snPcode":"42991","submissionUrl":"https://www.editorialmanager.com/mamb/default2.aspx","title":"Mammalian Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"78eb0011-8161-407d-9c80-22d11fdb3d99","owner":[],"postedDate":"October 29th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-16T16:04:23+00:00","versionOfRecord":{"articleIdentity":"rs-7794885","link":"https://doi.org/10.1007/s42991-026-00563-w","journal":{"identity":"mammalian-biology","isVorOnly":false,"title":"Mammalian Biology"},"publishedOn":"2026-02-10 15:59:27","publishedOnDateReadable":"February 10th, 2026"},"versionCreatedAt":"2025-10-29 12:33:24","video":"","vorDoi":"10.1007/s42991-026-00563-w","vorDoiUrl":"https://doi.org/10.1007/s42991-026-00563-w","workflowStages":[]},"version":"v1","identity":"rs-7794885","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7794885","identity":"rs-7794885","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}