Research Article |
Corresponding author: Roberto Leonan M. Novaes ( robertoleonan@gmail.com ) Academic editor: Clara Stefen
© 2022 Roberto Leonan M. Novaes, Vinícius C. Cláudio, M. Mónica Díaz, Don E. Wilson, Marcelo Weksler, Ricardo Moratelli.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Novaes RLM, Cláudio VC, Díaz MM, Wilson DE, Weksler M, Moratelli R (2022) Argentinean Myotis (Chiroptera, Vespertilionidae), including the description of a new species from the Yungas. Vertebrate Zoology 72: 1187-1216. https://doi.org/10.3897/vz.72.e90958
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Myotis is the most speciose genus of mammals in the world and recent taxonomic revisions have revealed an impressive diversity of species in South America. Even so, the phenotypic conservatism of some taxa makes taxonomic delimitation difficult. We perform a taxonomic review of Myotis from Argentina based on qualitative and quantitative morphological characters. Our results confirm the occurrence of 12 species (M. albescens, M. chiloensis, M. dinellii, M. izecksohni, M. keaysi, M. lavali, M. levis, M. nigricans, M. oxyotus, M. riparius, M. ruber, and M. cf. simus) and revealed an additional new species for the Yungas Forest. The new species is small to medium (forearm length ~ 35 mm) and can be distinguished from its congeners by a set of characters that includes forearm length, cranial measurements, discrete craniodental characters, and fur color. This review does not exhaust the need for new systematic studies with Argentinean Myotis, considering the possibility of occurrence of new species and the great morphological variation found for some complex taxa.
Argentina, morphology, Myotinae, Neotropics, systematics
Myotis Kaup, 1829 currently comprises more than 140 species distributed worldwide, and 24 are recognized from South America (
Taxonomic reviews of South American Myotis have been frequent over the last decades (i.e.,
Specifically, no reviews focusing on the species of Myotis from Argentina have been carried out so far. Nevertheless, several studies have made important contributions to our knowledge on the genus in this region (e.g.,
Based primarily on the large samples of Myotis available in scientific collections, we provide an overview of the taxonomic diversity of Myotis in Argentina, and describe a new species from Yungas Forest.
This research is part of a critical review of Neotropical Myotis, and more than 7,500 specimens from different localities in South America and Central America have been examined, covering all species currently recognized and their type specimens. Our analyses here were based on 187 specimens from Argentina, deposited in 11 scientific collections:
American Museum of Natural History (
Intra- and interspecific morphological variation were evaluated from qualitative and quantitative analyses based on adult specimens, classified from ossified epiphyses (q.v.,
Description of cranial, mandibular, and external dimensions (and their abbreviations) used in this study.
Measurements | Acronyms | Descriptions |
Greatest length of skull | GLS | From the apex of the upper internal incisors to the occiput. |
Condyle-canine length | CCL | From the anterior surface of the upper canines to a line connecting the occipital condyles. |
Condyle-incisive length | CIL | From the apex of upper internal incisors to a line connecting the occipital condyles. |
Basal length | BAL | Least distance from the apex of upper internal incisors to the ventral margin of the foramen magnum. |
Mastoid breadth | MAB | Greatest breadth across the mastoid region. |
Braincase breadth | BCB | Greatest breadth of the globular part of the braincase. |
Interorbital breadth | IOB | Least breadth between the orbits. |
Postorbital breadth | POB | Least breadth across frontals posterior to the postorbital bulges. |
Breadth across canines | BAC | Greatest breadth across outer edges of the crowns of upper canines, including cingulae. |
Breadth across molars | BAM | Greatest breadth across outer edges of the crowns of upper molars. |
Maxillary toothrow length | MTL | From the upper canine to M3. |
Molariform toothrow length | M13 | 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. |
Forearm length | FA | From the elbow to the distal end of the forearm including carpals. |
Length of the dorsal hairs | LDH | From the base to the tip of the hair in fur between scapulae. |
Length of the ventral hairs | LVH | In fur at mid thorax. |
Ear length | EL | From the base to the apex of the pinna |
To characterize and discriminate samples, Principal Component (PCA) and Discriminant Function (DFA) analyses were performed in R platform, with ‘MASS’ (
Our systematic review of Argentinean Myotis confirms the occurrence of 12 recognized species (M. albescens, M. chiloensis, M. dinellii, M. izecksohni, M. keaysi, M. lavali, M. levis, M. nigricans, M. oxyotus, M. riparius, M. ruber, M. cf. simus), and a new taxon, morphologically distinct from its congeners, representing an undescribed species.
Principal Component Analysis (Fig.
Vector correlation loadings with original variables of principal components (PC1 and PC2) and discriminant functions (DF1 and DF2) for selected samples of the Argentinean Myotis.
Measurements | PC1 | PC2 | DF1 | DF2 |
GLS | 0.573 | –0.538 | 0.367 | –0.847 |
CIL | 0.555 | –0.194 | 0.732 | –0.839 |
MAB | 0.260 | 0.512 | 0.589 | –0.137 |
BCB | 0.240 | 0.540 | 0.469 | –0.030 |
POB | 0.093 | 0.491 | 0.468 | 0.017 |
IOB | 0.187 | 0.254 | 0.336 | 0.018 |
BAM | 0.248 | 0.085 | 0.352 | –0.463 |
MTL | 0.199 | –0.380 | 0.237 | –0.563 |
M1-3 | 0.124 | –0.222 | 0.173 | –0.461 |
MAN | 0.176 | –0.495 | 0.142 | –0.665 |
MAL | 0.384 | 0.095 | 0.729 | –0.591 |
Skull and external linear morphometric measurements indicated that Myotis sp. nov. is consistently smaller than most species found in Argentina; but there is partial overlap for forearm length and some cranial measurements with M. lavali, M. nigricans, and M. riparius (Tables
Family Vespertilionidae Gray, 1821
Subfamily Myotinae Tate, 1942
Genus Myotis Kaup, 1829
Dried skin, skull, and mandible of an adult female (CML 7623; Figs
Finca Alto Verde, ca. 20 km SW San Ramón de La Nueva Orán, Orán Department, Salta Province, Argentina (23°13′S, 64°32″W; 670 m elevation).
Dried skin, skull and mandible of an adult female (CML 7622), collected by M. D. Miotti (field number 539) on September 14, 2006 at the type locality.
Myotis barquezi is known only from the type locality, in Salta Province, northern Argentina, where it inhabits lowland tropical forests inside the Southern Andean Yungas ecoregion (Fig.
The climate presents hot, rainy summers and cold, dry winters, and the annual mean temperature exceeds 21°C (
Myotis barquezi is named in honor of Dr. Rubén M. Barquez, in recognition of his outstanding contributions to Neotropical mammalogy, especially on the bat fauna from Argentina. This species name is a noun in the genitive case formed by adding −i to the stem of the name (ICZN, 1999; 31.1.2).
Barquez’s Myotis [English]; Myotis de Barquez [Spanish].
Small to medium sized species (FA 35.1–35.2 mm; GLS 13.1–13.5 mm); sagittal crest present but low; robust and broad skull; braincase inflated and remarkably high in lateral profile; braincase roof formed by the parietal bone strongly inclined forward; frontal bone with a slight slope towards the rostrum; posterior region of the braincase flattened and non-projected beyond the limit of the occipital condyles; well-developed mastoid processes; dorsal fur moderately long (LDF 5–6 mm), woolly and clearly bicolored, with tips ranging from Dresden Brown to Snuff Brown, and darker bases (Natal Brown); legs and dorsal surface of the uropatagium covered by fur that extend up to the knees or just beyond; plagiopatagium inserted into the foot by a broad band of membrane.
Myotis barquezi is a small to medium species of Myotis (Table
Selected measurements (mm) and body mass (g) of the holotype (CML 7623) and paratype of Myotis barquezi (CML 7622) from Salta, Argentina.
Measurements | Holotype CML 7623 | Paratype CML 7622 |
FA | 35.1 | 35.2 |
EL | 15.0 | 14.0 |
Body mass | 5.0 | 6.0 |
LDF | 6.0 | 5.0 |
LVF | 5.3 | 4.5 |
GLS | 12.9 | 13.4 |
CCL | 11.8 | 11.7 |
CBL | 12.3 | 12.4 |
CIL | 12.6 | 12.7 |
BAL | 11.2 | 11.6 |
ZYG | — | — |
MAB | 6.7 | 6.7 |
BCB | 6.6 | 6.3 |
POB | 3.4 | 3.4 |
IOB | 4.5 | 4.6 |
BAC | 3.4 | 3.4 |
BAM | 5.4 | 5.3 |
MTL | 5.1 | 5.0 |
M1–3 | 3.0 | 3.0 |
MAN | 5.3 | — |
MAL | — | — |
Dorsal and ventral fur woolly and medium-sized (LDH 5–6 mm, LVH ~4.5 mm). Dorsal pelage clearly bicolored with medium-brown bases (near Natal Brown) and reddish tips, ranging from Dresden Brown to Snuff Brown. Darker bases comprise 2/3 of total hair length and changes abruptly from darker to lighter, with lighter tips comprising about 1/3 of hair length. Ventral pelage is also strongly bicolored, with Natal Brown bases (2/3 of hair length) and bright orangish tips (1/3 of hair length [near Clay Color]). The orangish venter contrasts with the reddish-brown dorsum.
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 medium-sized in length, resembling Myotis species of the ruber-group (GLS 13.1–13.5 mm). The second upper premolar (P3) aligned with toothrow, not displaced to the lingual side, and barely smaller than first upper premolar (P2). First lower molar (m1) myotodont, with postcristid connecting hypoconid and entoconid (
Myotis barquezi can be distinguished from all the species in the ruber-group (armiensis, elegans, keaysi, midastactus, moratellii, pampa, pilosatibialis, riparius, ruber, and simus), and species from the albescens-group that confirmedly or potentially occur in the South American southern cone (albescens, chiloensis, dinellii, izecksohni, lavali, levis, nigricans, and oxyotus), by the set of diagnostic traits reported above.
Among species from the ruber-group, M. barquezi most resembles M. keaysi from the Central Andes (see
Myotis barquezi differs from M. ruber by its smaller size (both external and cranial; FA > 37.5 mm and GLS > 14.0 mm in M. ruber), shorter and clearly bicolored dorsal fur, dense fur along the leg and dorsal surface of the uropatagium, narrower interorbital constriction, and more developed mastoid processes. It differs from M. armiensis by its smaller size (FA > 36.0 mm in M. armiensis), clearly bicolored dorsal fur, lower sagittal crest, posterior region of the braincase flattened and non-projected beyond the limit of the occipital condyles. It differs from M. pilosatibialis by its smaller size (both external and cranial; FA > 36.0 mm in M. pilosatibialis), clearly bicolored dorsal fur, globose braincase (elongated in pilosatibialis), parietal bone strongly inclined forward, and shorter and broader rostrum. It differs from M. moratellii by its general smaller size (FA > 35.0 and GLS > 13.8 in moratellii), clearly bicolored dorsal fur, braincase lower in profile, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles, and sagittal crest lower.
Myotis barquezi can be distinguished from M. riparius by its clearly bicolored and reddish dorsal fur, presence of dense fur along the leg and dorsal surface of the uropatagium, parietal bone strongly inclined forward, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles, narrower interorbital constriction, and more developed mastoid processes. It differs from M. elegans by its larger size (both external and cranial; FA < 34.5 mm and GLS < 13.0 mm in M. elegans), more robust skull, higher sagittal crest, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles, narrower interorbital constriction, and more developed mastoid processes. It differs from M. pampa by its shorter fur (LDF < 6 in M. barquezi, LDF > 7 in M. pampa), ventral fur bicolored (being tricolored in M. pampa), skull more robust, sagittal crest present, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles, and more developed mastoid processes.
Myotis barquezi can be easily distinguished from M. simus and M. midastactus by its smaller size (FA > 36.0 mm and GLS > 13.5 mm in M. simus and M. midastactus), longer and clearly bicolored dorsal fur (being shorter [LDF < 4] and unicolored in M. simus and M. midastactus), legs and dorsal surface of the uropatagium covered by fur that extend up to the knees, plagiopatagium inserted into the foot by a broad band of membrane (attached at ankles in M. simus and M. midastactus); and narrower skull. In addition, tympanic bullae are comparatively larger in M. barquezi than in any other species from ruber-group, except M. elegans.
In comparison to the species from the albescens-group, M. barquezi can be easily distinguished from M. albescens by the absence of a fringe of hairs on the posterior margin of the uropatagium, reddish dorsal fur clearly bicolored (brownish with frosted appearance in albescens), yellowish ventral fur (whitish in albescens), frontal bone slightly sloping, sagittal crest present, and posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles. It differs from M. dinellii and M. levis by its smaller size, absence of a fringe of hairs on the posterior margin of the uropatagium, comparatively shorter ears, narrower skull, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles. It differs from M. izecksohni and M. nigricans by the reddish and clearly bicolored dorsal fur, parietal bone strongly inclined forward, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles, and well-developed mastoid processes.
Myotis barquezi can be distinguished from M. chiloensis from its general smaller size (FA < 35.2 in M. barquezi; FA > 37 mm in M. chiloensis), ventral fur strongly bicolored with bright orange tips, shorter and broader skull, less inflated braincase, parietal bone strongly inclined forward, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles. It differs from M. lavali by its shorter and reddish fur, broader skull, parietal bone strongly inclined forward, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles, and well-developed mastoid processes. Differs from M. oxyotus by its smaller size (FA > 37 mm), shorter dorsal fur, ventral fur strongly bicolored with bright orange tips, shorter and broader skull, broader skull, parietal bone strongly inclined forward, posterior region of the braincase flattened and not projected beyond the limit of the occipital condyles. In addition, M. barquezi can be distinguished from all species of the albescens-group by its woolly fur (silky fur in albescens-group species, except M. chiloensis), dense fur on dorsal surface of the uropatagium (absent in all species of albescens-group), and tympanic bullae comparatively larger.
The specimens used here to describe M. barquezi (CML 7622 and 7623) were originally misidentified as Myotis lavali (
Medium-sized species (FA 33.5–37.5 mm, body mass 4–8 g; Table
External and craniodental measurements of Argentinean populations of Myotis albescens, M. ruber, and M. nigricans, including morphometric variation and number of samples (N). The measurements are in millimeters. Acronyms and descriptions are available in Table
Measurements | Myotis albescens | Myotis ruber | Myotis nigricans |
mean (min–max) N | mean (min–max) N | mean (min–max) N | |
FA | 35.0 (33.7–36.8) 22 | 38.5 (37.5–40.5) 5 | 33.8 (33.0–35.0) 7 |
EL | 14.0 (12.0–15.5) 22 | 14.2 (13.0–15.0) 4 | 13.0 (11.0–15.0) 7 |
Body mass | 6.0 (5.5–7.0) 18 | 7.0 (4.0–8.0) 4 | 4.5 (3.5–6.0) 7 |
LDF | 6.5 (5.5–7.5) 17 | 7.0 (6.5–8.0) 4 | 7.0 (6.0–7.5) 7 |
LVF | 5.0 (4.5–6.0) 17 | 5.5 (5.0–6.0) 4 | 5.5 (5.0–6.5) 7 |
GLS | 14.3 (13.9–15.0) 20 | 15.3 (15.1–15.6) 5 | 13.5 (12.4–14.3) 7 |
CCL | 12.7 (12.1–14.2) 20 | 13.5 (13.5–13.7) 5 | 12.0 (11.6–12.7) 7 |
CBL | 13.1 (12.6–13.8) 20 | 14.2 (14.0–14.4) 5 | 12.7 (12.3–13.5) 7 |
CIL | 13.4 (12.9–14.0) 20 | 14.5 (14.4–14.8) 5 | 12.8 (12.4–13.5) 7 |
BAL | 12.1 (11.5–13.4) 20 | 13.0 (12.8–13.2) 5 | 11.5 (11.0–12.3) 7 |
ZYG | 8.8 (8.6–9.0) 11 | 9.5 (9.4–9.6) 3 | 8.3 (8.1–8.4) 4 |
MAB | 7.4 (7.0–7.7) 20 | 7.8 (7.7–8.2) 5 | 6.9 (6.6–7.1) 7 |
BCB | 7.1 (6.8–7.4) 20 | 7.0 (6.7–7.2) 5 | 6.4 (6.2–6.6) 7 |
POB | 4.1 (3.9–4.2) 20 | 3.7 (3.6–3.9) 5 | 3.5 (3.3–3.8) 7 |
IOB | 4.8 (4.5–5.0) 20 | 4.9 (4.7–5.2) 5 | 4.5 (4.1–4.7) 7 |
BAC | 3.8 (3.5–4.0) 20 | 4.1 (4.0–4.3) 5 | 3.4 (3.2–3.5) 5 |
BAM | 5.5 (5.2–5.7) 20 | 6.0 (5.8–6.2) 5 | 5.2 (4.9–5.6) 7 |
MTL | 5.1 (4.9–5.5) 20 | 5.9 (5.9–5.9) 5 | 5.0 (4.8–5.2) 7 |
M1–3 | 2.9 (2.8–3.1) 20 | 3.3 (3.2–3.3) 5 | 2.8 (2.7–3.0) 7 |
MAN | 5.8 (5.1–6.1) 20 | 6.3 (6.2–6.4) 4 | 5.3 (5.1–5.5) 7 |
MAL | 10.0 (9.5–10.4) 20 | 11.1 (10.9–11.4) 4 | 9.4 (9.0–9.7) 7 |
Occurs from southern Veracruz, Mexico, southward through Central America into Uruguay, northern Argentina and eastern Brazil, from humid tropical forests to savannas and semi-arid environments (
Medium to large-sized species (FA 35.8–40.5 mm; body mass 4–8 g; Table
The distribution is associated with ombrophilous and seasonal tropical forests from Northeastern Brazil to northern Argentina and Paraguay, where it appears to be strongly associated to humid and dense forested environments (
Recent studies have indicated that M. nigricans is a complex of allopatric species (
In Argentina, they occur in ombrophilous tropical forests from Humid Chaco (Provinces of Chaco and Corrientes) and moist Atlantic Forest (Misiones Province), in an altitudinal range from 40 to 400 m. The distribution of this species still needs to be reviewed considering the recent taxonomic changes in populations originally identified as M. nigricans. Thus, records for other Argentinean provinces (e.g.,
Medium-sized species (FA 36.5–40.1 mm, body mass 6–8 g; Table
External and craniodental measurements of Argentinean populations of Myotis levis, M. chiloensis, and M. oxyotus, including morphometric variation and number of samples (N). The measurements are in millimeters. Acronyms and descriptions are available in Table
Measurements | Myotis levis | Myotis chiloensis | Myotis oxyotus |
mean (min–max) N | mean (min–max) N | CML 10860 | |
FA | 38.6 (36.6–40.1) 5 | 38.5 (37.0–41.2) 30 | 40.9 |
EL | 17.0 (16.0–19.0) 5 | 15.0 (14.0–16.0) 22 | 19.0 |
Body mass | 7.0 (6.0–8.0) 4 | 7.5 (5.5–9.5) 20 | 4.0 |
LDF | 7.0 (6.0–8.0) 5 | 7.0 (6.0–8.5) 20 | 9.5 |
LVF | 6.0 (4.5–7.0) 5 | 6.0 (5.0–7.5) 20 | 7.5 |
GLS | 15.1 (14.4–15.5) 5 | 14.7 (13.8–15.3) 30 | 14.7 |
CCL | 13.2 (12.7–13.5) 5 | 13.0 (12.4–13.7) 30 | 12.8 |
CBL | 13.9 (13.5–14.2) 5 | 13.7 (13.3–14.4) 30 | 13.5 |
CIL | 14.1 (13.6–14.5) 5 | 14.0 (13.6–14.6) 30 | 13.9 |
BAL | 12.7 (12.1–13.2) 5 | 12.6 (12.1–13.1) 30 | 12.5 |
ZYG | 9.0 (8.5–9.3) 4 | - | 8.7 |
MAB | 7.6 (7.2–7.8) 5 | 7.6 (7.3–7.8) 30 | 7.1 |
BCB | 7.1 (6.8–7.5) 5 | 7.0 (6.4–7.5) 30 | 6.9 |
POB | 3.7 (3.5–3.8) 5 | 3.8 (3.5–4.0) 30 | 3.8 |
IOB | 4.7 (4.3–4.5) 5 | 4.7 (4.6–5.0) 30 | 4.9 |
BAC | 3.7 (3.5–3.9) 5 | 3.8 (3.6–4.0) 30 | 3.5 |
BAM | 5.8 (5.6–6.0) 5 | 5.8 (5.4–6.1) 30 | 5.5 |
MTL | 5.5 (5.4–5.6) 5 | 5.7 (5.5–5.8) 30 | 5.6 |
M1–3 | 3.2 (3.1–3.2) 5 | 3.2 (3.1–3.3) 30 | 3.6 |
MAN | 6.0 (5.8–6.1) 4 | 6.0 (5.6–6.2) 30 | 5.8 |
MAL | 10.6 (10.0–10.8) 4 | 10.6 (10.1–11.1) 30 | 11.0 |
This species occurs from Southeastern Brazil southward to Uruguay, Paraguay, and eastern Argentina, from ombrophilous Atlantic Forest to Pampa grasslands (
Medium to large-sized species (FA 37.0–41.2 mm, body mass 5.5–9.5 g; Table
Myotis chiloensis occurs from Southern Chile, eastward into western Argentina and southward to Tierra del Fuego, in evergreen-deciduous forests, montane temperate forests, and Patagonian scrublands (
Among all Myotis specimens from Argentina analyzed, only one voucher was identified as M. oxyotus (CML 10860). It is a medium-sized bat (FA 40.9 mm, body mass 4.0 g; Table
Myotis oxyotus is divided in two allopatric subspecies: Myotis o. oxyotus from South America and M. o. gardneri from Central America. Myotis o. oxyotus occurs along the Andes, from Venezuela southward to northern Argentina (
Medium to large-sized species (FA 34.7–39.4 mm, body mass 5–6 g; Table
External and craniodental measurements of Argentinean populations of Myotis cf. simus, M. dinellii, and M. keaysi, including morphometric variation and number of samples (N). The measurements are in millimeters. Acronyms and descriptions are available in Table
Measurements | Myotis cf. simus | Myotis dinellii | Myotis keaysi |
mean (min–max) N | mean (min–max) N | mean (min–max) N | |
FA | 39.1 (38.0–41.3) 4 | 36.8 (34.7–39.4) 51 | 39.4 (38.7–40.0) 8 |
EL | 10.5 (10.0–12.0) 4 | 15.5 (14.0–17.0) 33 | 13.5 (13.0–14.0) 4 |
Body mass | 9.0 (8.0–9.5) 3 | 5.0 (5.0–6.0) 33 | 7.0 (6.0–8.0) 4 |
LDF | 3.1 (2.7–3.4) 4 | 8.0 (7.0–9.1) 33 | 7.5 (6.5–8.0) 3 |
LVF | 2.8 (2.5–2.9) 4 | 6.5 (6.0–8.8) 33 | 5.5 (5.0–6.0) 3 |
GLS | 14.5 (14.0–15.0) 4 | 14.5 (13.9–15.3) 51 | 13.7 (13.5–13.9) 8 |
CCL | 12.6 (11.8–13.0) 4 | 12.8 (12.3–13.8) 51 | 12.4 (12.3–12.6) 8 |
CBL | 13.3 (12.9–13.8) 4 | 13.5 (12.9–14.5) 51 | 13.0 (12.8–13.1) 8 |
CIL | 13.7 (13.2–14.2) 4 | 13.7 (13.0–14.7) 51 | 13.3 (13.1–13.4) 8 |
BAL | 12.3 (11.8–13.0) 4 | 12.3 (11.7–13.2) 51 | 11.9 (11.7–12.0) 8 |
ZYG | 9.4 (9.3–9.5) 3 | 8.7 (8.4–9.4) 39 | 8.9 (8.8–9.0) 5 |
MAB | 7.7 (7.5–7.9) 4 | 7.2 (6.8–7.7) 51 | 7.3 (7.0–7.6) 8 |
BCB | 7.1 (6.8–7.5) 4 | 6.8 (6.5–7.1) 51 | 6.8 (6.7–7.0) 8 |
POB | 3.8 (3.6–4.1) 4 | 3.6 (3.3–3.8) 51 | 3.7 (3.6–3.8) 8 |
IOB | 4.9 (4.7–5.1) 4 | 4.6 (4.0–5.0) 51 | 4.5 (4.5–4.5) 8 |
BAC | 4.0 (3.7–4.1) 3 | 3.6 (3.1–3.9) 51 | 3.6 (3.6–3.6) 8 |
BAM | 5.7 (5.2–6.0) 4 | 5.6 (5.2–6.0) 51 | 5.5 (5.3–5.7) 8 |
MTL | 5.4 (5.1–5.8) 3 | 5.5 (5.2–5.8) 51 | 5.3 (5.1–5.4) 8 |
M1–3 | 3.1 (2.8–3.3) 4 | 3.2 (3.0–3.3) 50 | 2.9 (2.9–3.1) 8 |
MAN | 5.7 (5.4–6.2) 3 | 5.9 (5.6–6.2) 51 | 5.7 (5.6–5.8) 8 |
MAL | 10.8 (10.5–11.3) 4 | 10.3 (9.8–11.0) 51 | 9.9 (9.8–10.1) 8 |
This species occurs from southern Bolivia southward through Argentina, occupying deciduous forests, savannas, and semiarid open environments (
Medium to large-sized species (FA 38.7–42.0 mm, body mass 6–8 g; Table
Endemic to South America and strongly associated with highlands in the Andes, from Venezuela to northern Argentina, where it is present in habitats formed by Dense Ombrophylous Forest, Montana Nebular Forest and, in the areas of higher altitude, in Seasonal Forests dominated by shrubby vegetation and with sparse patches of arboreal vegetation (
Small to medium-sized species (FA 33.4–36.3 mm, body mass 4–5 g; Table
External and craniodental measurements of Argentinean populations of Myotis riparius, M. izecksohni, and M. lavali, including morphometric variation and number of samples (N). The measurements are in millimeters. Acronyms and descriptions are available in Table
Measurements | Myotis riparius | Myote izecksohni | Myotis lavali |
Mean (min–max) N | CML 10200 | Mean (min–max) N | |
FA | 34.3 (32.3–35.6) 10 | 37.8 | 34.0 (33.2–35.1) 9 |
EL | 13.0 (12.0–15.0) 7 | 12.1 | 14.0 (13.0–15.0) 5 |
Body mass | 5.0 (4.0–5.0) 7 | 4.0 | 6.5 (6.0–7.0) 5 |
LDF | 7.1 (6.5–7.5) 7 | 6.5 | 7.0 (6.5–7.5) 5 |
LVF | 6.0 (5.5–7.0) 7 | 5.3 | 6.0 (5.5–6.5) 5 |
GLS | 13.6 (13.4–13.8) 10 | 14.6 | 13.9 (13.6–14.3) 9 |
CCL | 12.1 (11.8–12.3) 10 | 12.6 | 12.3 (12.1–12.7) 9 |
CBL | 12.7 (12.4–12.8) 10 | 13.4 | 13.0 (12.7–13.3) 9 |
CIL | 12.9 (12.6–13.1) 10 | 13.8 | 13.2 (12.9–13.5) 9 |
BAL | 11.5 (11.3–11.7) 10 | 12.1 | 11.7 (11.4–12.1) 9 |
ZYG | 8.7 (8.6–8.7) 6 | 8.7 | 8.3 (8.3–8.3) 4 |
MAB | 7.0 (6.9–7.0) 10 | 7.1 | 7.0 (6.7–7.4) 9 |
BCB | 6.4 (6.3–6.4) 10 | 6.8 | 6.6 (6.1–6.9) 9 |
POB | 3.5 (3.4–3.6) 10 | 3.5 | 3.5 (3.4–3.6) 9 |
IOB | 4.4 (4.3–4.7) 10 | 4.5 | 4.5 (4.2–4.7) 9 |
BAC | 3.6 (3.5–3.6) 10 | 3.7 | 3.4 (3.3–3.6) 9 |
BAM | 5.4 (5.2–5.6) 10 | 5.7 | 5.4 (5.2–5.5) 9 |
MTL | 5.1 (4.9–5.2) 10 | 5.5 | 5.0 (4.9–5.2) 9 |
M1–3 | 2.9 (2.8–3.0) 10 | 3.1 | 2.9 (2.8–2.9) 9 |
MAN | 5.5 (5.2–5.7) 10 | 5.7 | 5.4 (5.2–5.5) 8 |
MAL | 9.8 (9.5–10.1) 10 | 10.5 | 9.7 (9.3–10.1) 8 |
Myotis riparius occurs from Honduras, southward through South America into northern Argentina and eastern Brazil, occupying humid tropical forests to savanna environments (
In Argentina, M. riparius occurs in the northern portion, from Southern Andean Yungas (Jujuy, Salta, and Tucumán Provinces) throughout ombrophilous tropical forests in Humid Chaco (Province of Chaco and Formosa), Dry Chaco (Santiago del Estero), and moist Atlantic Forest (Misiones Province), in an altitudinal range from 70 to 2,000 m (
Among all Myotis specimens from Argentina analyzed, only one voucher (CML 10200) was identified as M. izecksohni. It is a medium-sized species (FA 37.8 mm, body mass 4 g; Table
This species occurs from southeastern Brazil to northern Argentina, in dense ombrophilous forests (
Small to medium-sized species (FA 33.8–39.6 mm, body mass 5–6 g; Table
Myotis lavali occurs from Northeastern Brazil, throughout South American diagonal of open formations, to Paraguay and northern Argentina, in savannas, semi-arid fields, and seasonal forests (
The M. simus complex was taxonomically revised resulting in the recognition of the recently described species Myotis midastactus Moratelli & Wilson, 2014, based on individuals originally identified as M. simus from Bolivia (
In Argentina, this taxon is medium to large-sized (FA 38.0–41.3 mm, body mass 8.0–9.5 g; Table
Argentinean populations of M. cf. simus are present in Humid Chaco and Mesopotamian Savanna from Formosa and Corrientes Provinces, and in the Delta e Islas del Paraná in Santa Fe Province, occupying an altitudinal range from 46 to 200 m (
Myotis barquezi occurs in humid seasonal forests from Southern Andean Yungas, an ecoregion with a high richness of mammals, including endemic species (
Despite recent taxonomic revisions of Neotropical Myotis, we consider that a large part of Myotis species from Argentina have insufficient taxonomic knowledge. Of the 13 species listed for Argentina, three (M. keaysi, M. riparius, and M. simus) show strong morphological variation in relation to the type specimens and topotype series (
Until recently, M. dinellii was considered a subspecies of M. levis (
The present study indicates the occurrence of 13 species of Myotis in Argentina but does not end the need for further revisions. On the contrary, the pattern of variation found in several taxa suggests that the diversity of Argentinian Myotis may be underestimated and the taxonomic status of some populations need to be revised, especially in light of new morphological and, mainly, molecular evidence. We suggest that further studies of Myotis from Argentina should focus on investigating the morphological and genetic variation of the forms currently named M. riparius, M. simus, and M. keaysi.
With the description of M. barquezi, we recognize 25 species of South American Myotis, including M. albescens (É. Geoffroy, 1806), M. ruber (É. Geoffroy, 1806), M. nigricans (Schinz, 1821), M. levis (I. Geoffroy, 1824), M. chiloensis (Waterhouse, 1840), M. oxyotus (Peters, 1866), M. atacamensis (Lataste, 1892), M. simus Thomas, 1901, M. dinellii Thomas, 1902, M. caucensis Allen, 1914, M. keaysi J.A. Allen, 1914, M. riparius Handley, 1960, M. larensis LaVal, 1973, M. diminutus Moratelli & Wilson, 2011, M. lavali Moratelli, Peracchi, Dias & Oliveira, 2011, M. izecksohni Moratelli, Peracchi, Dias & Oliveira, 2011, M. handleyi Moratelli, Gardner, Oliveira & Wilson, 2013, M. midastactus Moratelli & Wilson, 2014, M. clydejonesi Moratelli, Wilson, Gardner, Fisher & Gutiérrez, 2016, M. attenboroughi Moratelli, Wilson, Novaes, Helgen & Gutiérrez, 2017, M. bakeri Moratelli, Novaes, Carrión-Bonilla & Wilson, 2019, M. armiensis Carrión-Bonilla & Cook, 2020, M. pampa Novaes, Wilson & Moratelli 2021, and M. moratellii Novaes, Cláudio, Carrión-Bonilla, Abreu, Wilson, Maldonado & Weksler, 2021. In line with other taxonomic revisions (i.e.,
The following curators and collection staff provided access to specimens under their care: P. Teta (
Listed below are localities with specimens examined from following collections: American Museum of Natural History (
Myotis albescens (N = 64): Argentina: Corrientes, San Martín, La Higuera Cué (
Myotis armiensis (N = 14): Ecuador: Tungurahua, Baños (
Myotis barquezi (N = 2): Argentina: Salta, Orán, ca. 20 km SW San Ramón de La Nueva Orán, Finca Alto Verde (CML 7622 [paratype], CML 7623 [holotype]).
Myotis chiloensis (N = 31): Argentina: Chubut, El Hoyo [de Epuyén] (CML 5218,
Myotis dinellii (N = 73): Argentina: Buenos Aires, General Guido, Canal 2 y Ruta 2 (
Myotis elegans (N = 25): Costa Rica (KU 158651). El Salvador (ROM 101293, 101319). Honduras: Olancho (
Myotis izecksohni (N = 48): Argentina: Misiones, San José, Apóstoles, Parque Provincial de la Sierra, Colonia Taranco (CML 10200). Brazil: Rio de Janeiro, Tinguá, Reserva Biológica do Tinguá (ALP 6618, 6626, 6675, 6676, MN 74357, 74358); Rio de Janeiro, Teresópolis, Parque Nacional da Serra dos Órgãos (ALP 6450, 6498, 6501, 6513, 6524); Paraná, Balsa Nova, São Luis do Purunã (CCMZ-DZUP 196, 197, 198, 199, 200, 410, 419, 420, 421, 422); Paraná, Campinhos (CCMZ-DZUP 56, 57, 58, 59, 61, 62, 63, 64, 65, 66, 67, 85, 86, 87, 88, 92, 93, 96, 97, 99, 105, 107, 108, 109, 110, 112).
Myotis keaysi (N = 36): Argentina: Tucumán, Burruyacú, Piedra Tendida, 8 km W Dique El Cajón (CML 6177, 7600, 9839); Tucumán, Burruyacú, Reserva Provincial Aguas Chiquitas, Arroyo Aguas Chiquitas (CML 8938); Tucumán, Burruyacú (
Myotis lavali (N = 58): Argentina: Salta, 1 km al E de Tonono sobre río Itiyuro (CML 5324); Santiago del Estero, Pellegrini, Santo Domingo (
Myotis levis (N = 47): Argentina: Buenos Aires, La Valle (
Myotis midastactus (N = 30): Bolivia: Beni, Cercado (
Myotis moratellii (N = 9): Ecuador: Los Ríos, Abras de Mantequilla, Hacienda Santa Teresita (
Myotis nigricans (N = 72): Argentina: Chaco, Sargento Cabral, La Forestal, Capitan Solari (
Myotis oxyotus (N = 26): Argentina: Jujuy, Yavi, el Mirador (CML 10860). Colombia: Nariño: El Guabo (
Myotis pampa (N = 15): Uruguay: Artigas, ca. 6 km NW from Belén (
Myotis pilosatibialis (N = 32): El Salvador: Ahuachapan, El Imposible, San Francisco Menendez (ROM 101273), Santa Ana, Parque Nacional Montecristo (ROM 101352, 101353, 101356, 101357, 101430, 101433, 101465–101467, 101524). Honduras: Francisco Morazan, 1 km W Talanga (LACM 36879 [holotype]), Cortes (
Myotis riparius (N = 63): Argentina: Corrientes, Capital, Laguna Paivas, Barrio Los Lomas (CML 2994); Formosa, Rio Porteño, km 64, a 5 km al sur de Estancia Sta. Catalina (OMHN 18889); Formosa, Río Bermejo, 10 millas al S de Colonia km 503 (CML 5412); Formosa, Parque Nacional Pilcomayo (
Myotis ruber (N = 60): Argentina: Formosa, Laguna Blanca, Parque Nacional Pilcomayo (CML 4666, 4673, 4676, 4686); Misiones, Guarani, jat. Hwy 21, Arroyo Oveja Negra, ~2 km W Parque Provincial Moconá (CML 3877); Misiones, Parque Nacional Iguazú, Ayui (
Myotis simus (N = 68): Brazil: Amazonas, Manaus (
Myotis cf. simus (N = 4): Argentina: Corrientes, Isla Apipé (