a new species of myotis (chiroptera, vespertilionidae) from bolivia

Journal of Mammalogy, 95(4):E17–E25, 2014

A new species of Myotis (Chiroptera, Vespertilionidae) from Bolivia

RICARDO MORATELLI* AND DON E. WILSON

Campus Fiocruz da Mata Atlantica, Fundacao Oswaldo Cruz, CEP 22713-375, Rio de Janeiro, RJ, Brazil (RM)Division of Mammals, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC20013-7012, USA (DEW, RM)

* Correspondent: [email protected]

We describe Myotis midastactus sp. nov. from the Bolivian savanna on the basis of differences in fur color, and

cranial and external features that unquestionably distinguish it from all other Neotropical Myotis. This new

species is morphologically allied to M. simus Thomas, 1901 and other species in the M. ruber group. Myotismidastactus is endemic to Bolivia, where it occurs with 6 congeners—albescens, dinellii, keaysi, nigricans,

riparius, and oxyotus. Previously identified as M. simus, M. midastactus is in syntopy with M. nigricans and M.riparius in the department of Santa Cruz, and there is no evidence that true M. simus occurs in Bolivia.

RESUMEN

Aquı describimos Myotis midastactus sp. nov., una nueva especie de murcielago de las sabanas de Bolivia.

Diferencias en el color de la piel y en caracterısticas craneanas y externas sin duda le distinguen de todas las

demas especies del genero Myotis presentes en el Neotropico. Esta nueva especie se alıa morfologicamente a M.simus Thomas, 1901 y otras especies en el grupo Myotis ruber. Myotis midastactus es endemica de Bolivia,

donde tambien se encuentran 6 congeneres—albescens, dinellii, keaysi, nigricans, riparius, y oxyotus. Myotismidastactus se encuentra en sintopıa con M. nigricans y M. riparius en el departamento de Santa Cruz. Con la

descripcion de esta nueva especie, no hay pruebas de la presencia de M. simus en Bolivia.

RESUMO

Descrevemos Myotis midastactus sp. nov. do Cerrado Boliviano. Diferencas na coloracao da pelagem e em

caracteres externos e cranianos inquestionavelmente distinguem-na de todas as especies de Myotis neotropicais.

Myotis midastactus e morfologicamente similar a Myotis simus e outras especies do grupo Myotis ruber. Myotismidastactus e endemica da Bolıvia, onde outras 6 especies do genero tambem ocorrem—albescens, dinellii,keaysi, nigricans, riparius e oxyotus. Myotis midastactus ocorre em sintopia com M. nigricans e M. riparius no

departamento de Santa Cruz. Apos a descricao dessa nova especie, nao ha evidencia da ocorrencia de M. simusna Bolıvia.

Key words: Myotinae, Myotis simus, Myotis ruber group, Neotropics, South America

� 2014 American Society of Mammalogists

DOI: 10.1644/14-MAMM-149

The genus Myotis Kaup, 1829 is unique among mammals in

its high diversity and near-global distribution. The genus

includes over 100 species (Simmons 2005), with representa-

tives in nearly all biogeographical regions. The biogeographic

reconstruction by Ruedi et al. (2013) retrieved East Asia as the

place of origin of the genus, with the divergence between New

and Old World lineages dating from the mid- to late Miocene.

Ruedi et al. (2013) suggested the Bering Strait as the gateway

for New World colonization, with an ancestral form reaching

the Neotropics by the Late Miocene. Early New World Myotis

diverged into 3 main branches. The South American and

Caribbean assemblage recognized by Simmons (2005) and

Wilson (2008) includes representatives of 2: the keaysi–riparius assemblage (a monophyletic lineage that includes

keaysi, ruber, simus, elegans, riparius) and the martiniquen-sis–levis assemblage (martiniquensis, dominicensis, nyctor,

w w w . m a m m a l o g y . o r g

E17

http://www.mammalogy.org

atacamensis, nesopolus larensis, chiloensis, albescens, oxy-otus, nigricans, levis—see Ruedi et al. 2013:7, figure 3).

Moratelli et al. (2013) assigned the oldest available names to

each lineage; thus the branch including keaysi and riparius is

the ruber group, and the branch including martiniquensis and

levis became the albescens group. Species in the ruber group

generally are characterized by woolly fur, a flattened occipital

region, and a moderate to high sagittal crest.

Myotis simus Thomas, 1901 (sensu lato), is the most

distinctive species in the ruber group and among Neotropical

Myotis generally. Its uniqueness led Cabrera (1958) to erect the

subgenus Hesperomyotis to set M. simus apart from the other

South American species. Myotis simus occurs only on the

eastern side of the Andes and currently is known from the

Amazon and Parana basins, including northern Argentina,

Paraguay, Bolivia, northern Brazil, Peru, Ecuador, and

Colombia (Moratelli et al. 2011a, 2013; Moratelli 2012).

Moratelli et al. (2011b) suggested that population samples

currently assigned to M. simus from Bolivia are morphomet-

rically and morphologically distinct from Amazon basin

samples, and probably represented a separate species. Speci-

mens from Bolivia have brighter dorsal and ventral fur colors,

larger skulls, and more laterally projecting mastoids (Moratelli

et al. 2011b). After further analyses of M. simus (sensu lato)

from Bolivia and the Amazon Basin, including topotypes from

Peru, we describe the Myotis from Bolivia as a new species,

which we compare with other taxa in the ruber group.

MATERIALS AND METHODS

The material we analyzed includes 106 M. simus (sensu

Wilson 2008) from Bolivia and the Amazon basin in

Colombia, Ecuador, northern Brazil, and Peru. Collections

are in the American Museum of Natural History, New York

FIG. 1.—Dorsal, ventral, and lateral views of the skull (bar¼ 5 mm) of the holotype of Myotis midastactus (AMNH 211156). See Table 1 for

measurements.

E18 Vol. 95, No. 4JOURNAL OF MAMMALOGY

(AMNH); Museum of Texas Tech University, Lubbock (TTU);

National Museum of Natural History, Smithsonian Institution,

Washington, DC (USNM); Universidade Federal Rural do Rio

de Janeiro, Seropedica, Brazil (ALP); and the Museu de

Zoologia da Universidade de Sao Paulo, Sao Paulo, Brazil

(MZUSP). A complete list of specimens examined is in

Appendix I. Most of this material was used by Moratelli et al.

(2011b) in their analyses of geographic variation in M. simus.

These samples cover most of the distribution of M. simus (see

Moratelli 2012), including the type locality (Sarayacu, Rıo

Ucayali, Loreto, Peru), but do not include southernmost

localities in Paraguay and northern Argentina. The samples

we examined from Bolivia (27 adults) are reassigned to the

species described here; the 57 adults and 22 subadults or

juveniles from the Amazon basin constitute our samples of M.simus (sensu stricto) on the basis of their similarity with the

holotype and topotypes (see Moratelli et al. 2011b). This

research is part of a broader systematic review of Neotropical

Myotis, and more than 2,000 vouchers have been examined and

include representatives of most New World species. Within the

ruber group, where this new species belongs, 339 adults and

juveniles representing M. elegans Hall, 1962, M. keaysi J. A.

Allen, 1914, M. pilosatibialis LaVal, 1973, M. ripariusHandley, 1960, and M. ruber (E. Geoffroy, 1806) have been

examined.

All measurements in this report are either in millimeters or

grams (body mass) and are from adults. The lengths of head

and body (HB), tail, hind foot, ear, and the body weight (mass)

were recorded from skin labels. Other dimensions were

measured using digital calipers accurate to 0.02 mm. They

include the forearm length (FA), 3rd metacarpal length (3ML),

length of the dorsal hair (LDH), length of the ventral hair

(LVH), greatest length of skull (GLS), condylocanine length

(CCL), condylobasal length (CBL), condylo-incisive length

(CIL), basal length (BAL), zygomatic breadth (ZB), mastoid

breadth (MAB), braincase breadth (BCB), interorbital breadth

(IOB), postorbital breadth (POB), breadth across canines

(BAC), breadth across molars (BAM), maxillary tooth-row

length (MTL), molariform tooth-row length (M13), mandibular

length (MAL), and mandibular tooth-row length (MAN).

These measurements are defined in Moratelli et al. (2013:3).

Skull and external measurements (except those from skin

labels) were rounded to tenths. Descriptive statistics (mean and

range) were calculated for all dimensions. Capitalized color

nomenclature is based on Ridgway (1912).

RESULTS

MYOTIS MIDASTACTUS, NEW SPECIES

GOLDEN MYOTIS, MURCIELAGO VESPERTINO DE ORO

FIGS. 1–3

Myotis simus: Anderson et al., 1982:10; not M. simus Thomas,

1901.

Myotis simus: Wilson and Salazar, 1990:52; not M. simusThomas, 1901.

Myotis simus: Anderson, 1997:276; not M. simus Thomas,

1901.

Myotis simus: Aguirre et al., 2002:1272; not M. simus Thomas,

1901.


1901.

Myotis simus: Emmons et al., 2006:56; not M. simus Thomas,

1901.


1901.

Myotis simus: Espinoza, 2007:326; not M. simus Thomas,

1901.

Myotis simus: Tarifa and Aguirre, 2009; not M. simus Thomas,

1901.

FIG. 3.—Adult female of Myotis midastactus (USNM 584502)

captured at Noel Kempff Mercado National Park, department of Santa

Cruz, Bolivia. Photograph courtesy of Marco Tschapka.

FIG. 2.—A) Dorsal and B) ventral views of the skin of the holotype

of Myotis midastactus (AMNH 211156). Forearm length 38 mm.

August 2014 E19MORATELLI AND WILSON—NEW MYOTIS FROM BOLIVIA


1901.

Myotis simus: Moratelli et al., 2011a; part, not M. simusThomas, 1901.

Myotis simus: Moratelli et al., 2011b; part, not M. simusThomas, 1901.

Myotis simus: Moratelli 2012; part, not M. simus Thomas,

1901.

Holotype.—Adult male, AMNH 211156, with skin,

complete skull, and postcranial skeleton (Figs. 1 and 2),

collected by S. Anderson (field number SA 5871), on 9

September 1965. External and craniodental measurements for

the type series are in Table 1.

Type locality.—Cercado, Rıo Mamore, Beni, Bolivia, about

23 km W of San Javier (148340S, 648550W).

Paratypes.—Includes 27 adults (26 females, 1 male), 26 of

which are from the type locality (AMNH 211155, 211156,

211167–211176, 211178–211184, 211190, 211192–211198),

and one is from El Refugio Huanchaca, Parque Nacional Noel

Kempff Mercado (PNNKM), Santa Cruz, Bolivia (USNM

584502; Fig. 3). Paratypes from Beni were collected by S.

Anderson and D. E. Anez on 9 September 1965, and the 1 from

Santa Cruz was collected by L. Emmons on 12 September

2000. Among paratypes deposited at AMNH, only skulls and

skins were examined.

Other specimens.—Seventeen additional specimens (16

females, 1 sex undetermined; AMNH 211157–211166,

211177, 211185–211189, 211191), previously identified as

M. simus, and collected by S. Anderson and D. E. Anez at the

type locality, are not included in the paratype series because we

did not examine this material.

Distribution.—Myotis midastactus is known only from

Bolivia, with all records from the departments of El Beni and

Santa Cruz (Fig. 4). Material from Beni examined here is from

23 km W of San Javier (148340S, 648550W). Anderson (1997)

also reported material from near Espıritu (» 148130S, 668400W)

and Curiraba River (148500S, 668230W; see Anderson [1997]

for localities, and Gardner’s [2008] Gazetteer for coordinates).

The specimen from Santa Cruz is from El Refugio Huanchaca

(14846 002 00S, 61802 005 00W [L. Emmons, pers. comm.]),

PNNKM, elevation 150 m. All localities are in the Cerrado

Biogeographical Province (see Anderson 1997:75, figure 470).

The PNNKM locality is in semideciduous forest (Fig. 5—

Emmons et al. 2006).

Diagnosis.—Myotis midastactus can be distinguished from

all New World congeners by its golden-yellow fur. M.

midastactus can be distinguished from all other Neotropical

Myotis by the following set of traits: plagiopatagium attached

to the foot by a narrow band of membrane (� 1.5 mm—see

Lopez-Gonzalez et al. 2001:141, figure 1b); woolly, extremely

TABLE 1.—Selected measurements (mm) and body weight (g) of the holotype (AMNH 211156) and the paratype series of Myotis midastactus,

and of a pooled sample of Myotis simus from Brazil, Colombia, Ecuador, and Peru. Samples include adults only, with males and females

combined. See ‘‘Materials and Methods’’ for variable abbreviations.

Myotis midastactus Myotis simus

Holotype Paratypes Brazil, Colombia, Ecuador, and Peru

? X Min. Max. n X Min. Max. n

HB 93 90 86 92 9 85 83 87 2

Tail 39 39 36 40 9 33 33 33 2

Hind foot 10 9 8 10 9 8 7 9 3

Ear 13 13 12 13 9 12 11 13 3

Weight 10 6 11 14 7 5 8 2

FA 38.2 39.4 38.5 40.7 16 37.8 35.5 39.7 29

3ML 36.2 36.2 34.5 37.9 16 34.7 32.7 36.9 35

LDH 4.4 4.6 3.4 5.5 16 4.0 3.1 5.3 29

LVH 4.8 3.8 2.9 5.3 16 3.4 2.8 4.2 29

GLS 15.1 14.6 13.9 15.0 26 14.1 13.6 14.8 43

CCL 13.4 13.0 12.4 13.3 25 12.4 11.9 13.0 40

CBL 13.9 13.6 13.2 14.0 25 13.0 12.5 13.5 40

CIL 14.2 13.9 13.3 14.2 25 13.3 12.7 13.9 41

BAL 12.7 12.4 11.8 12.7 25 11.8 11.2 12.4 40

ZB 10.0 9.9 9.6 10.2 15 9.1 8.2 9.5 11

MAB 8.3 8.2 8.0 8.5 26 7.5 6.9 8.1 33

BCB 7.4 7.4 7.1 7.7 26 6.9 6.6 7.3 39

IOB 5.1 5.0 4.8 5.3 26 4.8 4.5 5.0 44

POB 4.0 4.0 3.8 4.3 26 3.8 3.6 4.0 44

BAC 4.2 4.2 4.0 4.5 26 4.1 3.7 4.5 42

BAM 6.1 6.1 5.8 6.4 26 5.7 5.3 6.0 44

MTL 5.5 5.4 5.1 5.6 26 5.1 4.9 5.3 43

M13 3.2 3.2 3.1 3.3 26 3.0 2.7 3.2 44

MAL 10.9 10.7 10.2 11.0 23 10.1 9.5 10.5 15

MAN 5.8 5.8 5.5 6.0 24 5.4 5.2 5.8 43


short fur (LDH, LVH � 5 mm); ZB . 9.5 mm; MAB . 7.8

mm.

Description and comparisons.—Assuming that the Bolivian

specimen studied by Baker and Jordan (1970) represents M.midastactus, its karyotype is indistinguishable from that of all

other Myotis studied (sensu Stadelmann et al. [2004] and

excluding the African Cistugo Thomas, 1912). The diploid (2n)

and fundamental numbers are 44 and 50, respectively. The

autosomal complement has 4 pairs of metacentric

chromosomes (3 large, 1 small) and 17 pairs of acrocentric

chromosomes (graded in size from medium to small). The X is

a medium-sized submetacentric, and the Y is a small

acrocentric (Baker and Jordan 1970). Like all other Myotis—

except M. annectans (Dobson, 1871) and M. ridleyi (Thomas,

1898) from the Oriental region—the dental formula of M.midastactus is 2/3, 1/1, 3/3, 3/3 ¼ 38 (see Koopman 1994),

with the 3rd premolars (P4, p4) larger than the 1st and 2nd

premolars (P2, p2; P3, p3, respectively—see Tate 1941); the

tragus is long and slender (Miller 1907; Koopman 1994). Like

most other New World species, the dorsal surfaces of wing

membranes and uropatagium appear to be naked or nearly

naked.

In comparison with other Neotropical Myotis, M. midastac-tus and M. simus have larger upper canines in length and basal

cross-section, and shorter distances between C and P4. In both

species this short distance results in the P2 and P4 being close

or in actual contact, displacing P3 lingually so that P3 is

usually not visible in labial view. Although the P3 is

consistently displaced to the lingual side, as has been pointed

out as a diagnostic trait for simus (see LaVal 1973; Baud and

Menu 1993), the base of the tooth usually is close to alignment

with the other premolars in the tooth row, but its apex is

strongly inclined lingually. Myotis midastactus also fits into

this pattern, with the P3 not clearly visible in labial view (92%,

versus 73% in M. simus). In some M. midastactus the tooth

either is totally displaced lingually and not visible (4%, versus

9% in M. simus), or is aligned and visible in labial view (4%,

versus 18% in M. simus). Myotis midastactus and M. simusalso share other characteristics such as having sagittal and

occipital crests (sagittal crest: midastactus 96%, simus 97%;

occipital crest: midastactus 100%, simus 97%) that vary from

low to moderate in height in midastactus, and moderate to high

in simus; occipital region flattened (midastactus 100%, simus85%); plagiopatagium attached at the base of the toes by a

narrow band of membrane (midastactus 100%; simus 83%,

with the remainder [17%] attached at the ankle); and extremely

short, woolly fur (LDH, LVH � 5 mm). This combination of

characters distinguishes midastactus from all other Neotropical

congeners except M. simus. From M. simus, M. midastactuscan be distinguished by its paler and brighter dorsal and ventral

fur colors. M. midastactus also averages larger than M. simus in

all external and cranial measurements (Table 1). The most

useful cranial traits for identifying specimens in the absence of

FIG. 4.—Map of part of South America illustrating localities of

Myotis midastactus (stars) and Myotis simus (circles) samples used in

the analyses, and records tentatively assigned to M. simus but pending

confirmation (triangles). See Appendix I for localities.

FIG. 5.—A) and B) Semideciduous forest at El Refugio Huanchaca

in the dry season; Noel Kempff Mercado National Park, department of

Santa Cruz, Bolivia. Photographs courtesy of Louise Emmons.


skins is the broader skull, with more laterally projecting

mastoids (midastactus: ZB 9.6–10.2, MAB: 8.0–8.5; simus: ZB

8.2–9.5, MAB 6.9–8.1 [mm]—see Moratelli et al. 2011b:49,

figure 6). With regard to fur color, Bolivian samples are

consistently Ochraceous-orange dorsally, with a slightly paler

venter. Few specimens are slightly paler, with mixed blocks of

grayish and golden fur (Fig. 6). On the other hand, adult and

juvenile M. simus are Tawny, Russet, or Cinnamon-brown

dorsally, and Ochraceous-tawny or Buckthorn-brown on the

venter—all consistently darker than M. midastactus (Fig. 6).

Myotis midastactus can be distinguished from the species in

syntopy—M. nigricans and M. riparius—by its short, woolly,

and paler fur (LDF: 3.4–5.4 mm; fur is dark and longer in M.

nigricans and M. riparius); plagiopatagium attached at foot,

continuing to the base of the toes as a narrow band of

membrane; and more robust skull, mandibles, and teeth. The

low to moderate sagittal crest is most strongly developed over

the interparietals in M. midastactus, but in M. riparius the crest

is highest in the middle of the braincase; in M. nigricans the

sagittal crest is usually absent or very low.

Etymology.—The name M. midastactus is an allusion to the

legend of Midas, the mythical Greek king whose touch turned

everything to gold. It is a Latinized word (Gr. Mida1, Midas; L.

tactus, touch) in apposition to Myotis, and literally means

Midas touch in reference to the golden-yellow fur, which is

unique among New World Myotis. We suggest the English

vernacular name Golden Myotis, and the Spanish vernacular,

murcielago vespertino de oro.

Natural history and conservation.—Most of the information

available on the ecology, reproduction, and natural history of

M. midastactus comes from museum labels and from L.

Aguirre’s fieldwork in Bolivia. This information is synthesized

under accounts for M. simus in Espinoza (2007) and Moratelli

(2012). Myotis midastactus has day roosts in holes in the

ground, under thatched roofs, and hollow trees (Espinoza 2007;

Moratelli 2012). In the first 2 roosts, it was found together with

the Argentine brown bat, Eptesicus furinalis d’Orbigny and

Gervais, 1847 (Espinoza 2007). According to Aguirre et al.

(2003), 1 or 2 individuals of M. midastactus have been found

in hollow trees (Handroanthus heptaphyllus: Bignoniaceae) in

mixed groups with the lesser bulldog bat, Noctilio albiventrisDesmarest, 1818; and 1 was found sharing a roost with N.albiventris and the Pallas’s mastiff bat, Molossus molossus(Pallas, 1766). Individuals of M. midastactus and N. albiventrisfrom these roosts had similarly colored fur, texture, and smell;

on the basis of the number of individuals, M. midastactus is

solitary or forms very small groups (Aguirre et al. 2003).

Myotis simus is another species that shares roosts with N.albiventris; cohabitation records are from Paraguay (Myers and

Wetzel 1979) and northern Brazil (Patterson 1992). Population

samples from northern Brazil unquestionably represent M.simus, but those from Paraguay were not included here and

their identification is pending. According to L. Emmons (pers.

comm.), the adult female from PNNKM was caught during the

dry season in a deciduous secondary forest adjacent to an

airstrip; M. midastactus is the 77th species of bat identified as

occurring in the PNNKM—see Emmons et al. (2006) for a

description of PNNKM, and for a partial list of the bat

assemblage currently recognized from PNNKM. In a study by

Aguirre (1994), M. midastactus were found feeding on

Orthoptera (Gryllidae), Hemiptera (Corixidae), Homoptera

(Cicadellidae, Delphacidae), Coleoptera (Carabidae,

Dytiscidae, Staphylinidae, Scarabaeidae, Curculionidae),

Lepidoptera, and Diptera (Nematocera), with most items

measuring less than 10 mm. In the same study, 10 pregnant

females were captured in September, each with 1 embryo. In a

study of resource partitioning by bats in a Bolivian savanna,

Aguirre et al. (2002) found that M. midastactus—for its body

weight, and height and length of head (8.2 g, 9.8 mm, 14.4

mm, respectively [n¼ 1])—has a bite force (2.88 N) expected

for an animal of such proportions. Tarifa and Aguirre (2009)

considered M. simus from Bolivia (¼ M. midastactus) Near

Threatened, but we do not have enough evidence to comment

on its conservation status.

Nomenclatural statement.—A life science identifier (LSID)

number was obtained for the new species (Myotis midastactus):

FIG. 6.—A) Dorsal and B) ventral pelage of Myotis simus (AMNH

76252 [left]) and Myotis midastactus (AMNH 211181 [center],

AMNH 211190 [right]).


urn:lsid:zoobank.org:pub:5E2233E3-F7CA-42E2-8676-

4D99C6EED1FD.

DISCUSSION

The type locality of M. guaycuru Proenca, 1943, is in Mato

Grosso do Sul, Brazil, geographically closer to Bolivia than to

the Amazon basin localities to the north, but the type

specimen is morphologically and morphometrically like

Amazon basin populations (Moratelli et al. 2011b). Therefore,

the name M. guaycuru is not applicable to our Bolivian

populations, and is not a senior synonym of M. midastactus.

We did not examine Argentinean and Paraguayan samples,

but on the basis of descriptions of M. simus by Barquez et al.

(1999) and Lopez-Gonzalez (2005), southernmost popula-

tions in these countries appear to represent M. simus rather

than M. midastactus. On the basis of locality records from the

Brazilian Pantanal and Cerrado, we speculate that the

southernmost populations of M. simus (sensu stricto) in

Argentina and Paraguay are connected with populations in the

Amazon basin through the Brazilian Pantanal and Cerrado

(see Fig. 4).

On the basis of phylogenetic affinities retrieved by

Stadelmann et al. (2007) and additional morphological

evidence, Moratelli et al. (2013) proposed dividing the

assemblage of Venezuelan and Colombian Myotis spp. into

the albescens and ruber groups. Using the more compre-

hensive and better-resolved phylogeny derived by Ruedi et

al. (2013) as support, the same rationale can be applied to

the remaining Neotropical Myotis. In this phylogeny, a

previous polytomy among species in the ruber group is now

resolved, with keaysi [ruber [riparius [simus, elegans]]]. But

with the recent recognition of M. keaysi pilosatibialis at the

species level (Mantilla-Meluk and Munoz-Garay 2014), the

material assigned by Stadelmann et al. (2007) and Ruedi et

al. (2013) to M. keaysi (from Yucatan, Mexico) must be

reassigned to M. pilosatibialis, with M. keaysi (sensu stricto)

pending positioning. Myotis midastactus is also pending

positioning but both species are unquestionably allied in the

ruber-group. This group is sister to a lineage including M.vivesi and other species currently distributed in the

Neotropical and Nearctic regions. All species in the rubergroup are restricted to the Neotropics. Myotis elegans is

restricted to Mexico and Central America; M. keaysi is now

restricted to high elevations in the Andes from Colombia

southward to Bolivia and Argentina; M. midastactus is

endemic to Bolivia (the 2nd species of bat endemic to the

country [see Siles et al. 2013]); M. pilosatibialis occurs from

Mexico southward to Colombia; M. ruber is found only in

eastern Brazil; and M. simus apparently occurs widely in the

Amazon basin southward to Paraguay and northern Argenti-

na through the Brazilian Pantanal and Cerrado.

New World Myotis remain a puzzle, and more morpholog-

ical and molecular research is necessary to understand species

limits and phylogenetic relationships among Neotropical

species. Examining each piece of this puzzle moves us closer

to understanding the geological and ecological processes that

have shaped this assemblage.

NOMENCLATURAL STATEMENT

This article represents an electronic publication conforming

to the International Code of Zoological Nomenclature. This

electronic work represents an immutable copy published in a

journal with an ISSN number and has been archived in the

following digital repositories: LOCKSS and Portico. This

published work and the nomenclatural acts herein have been

registered in ZooBank and assigned a LSID. The LSID for this

publication is: urn:lsid:zoobank.org:pub:5E2233E3-

F7CA-42E2-8676-4D99C6EED1FD. The information can be

a c c e s s e d a t h t t p : / / z o o b a n k . o r g / R e f e r e n c e s /

5E2233E3-F7CA-42E2-8676-4D99C6EED1FD.

ACKNOWLEDGMENTS

The following curators and collection staff provided access to

specimens under their care: A. Peracchi (ALP), M. de Vivo, J.

Barros (MZUSP), N. Simmons, E. Westwig (AMNH), R. Baker, H.

Garner (TTU), K. Helgen, D. Lunde, and L. Gordon (NMNH). L.

Emmons (NMNH) provided information on a specimen she

collected in Santa Cruz, Bolivia and photos of its habitat, and M.

Tschapka (University of Ulm, Germany) provided the photograph of

this animal. L. Aguirre (Universidad Mayor de San Simon, Bolivia)

provided information on natural history and bibliography. A.

Gardner (United States Geological Survey Patuxent Wildlife

research Center) reviewed a draft of the manuscript, and E.

Gutierrez (USNM) reviewed the abstract in Spanish. This work

was supported by the National Council for Scientific and

Technological Development, Brazil (CNPq 202612/2012) and the

Smithsonian Institution.

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Submitted 16 May 2013. Accepted 2 June 2014.

Associate Editor was Ryan W. Norris.

APPENDIX IListed below are localities with specimens examined and those

pending confirmation. Abbreviations for collections are in

‘‘Material and Methods.’’ Localities are arranged by species

and alphabetically by country and by major political unit

(department/state, province/county). For locality numbers see

Fig. 4. Coordinates for M. midastactus are from Anderson

(1997 [AMNH material]) and L. Emmons (pers. comm.

[USNM material]). Those for M. simus were obtained from

Gardner’s (2008) Gazetteer of marginal localities or directly

from labels.Myotis midastactus (n ¼ 28; 26 //, 2 ??): BOLIVIA: Beni,

Cercado, about 23 km W of San Javier ([locality 1] 148340S,

64855 0W—AMNH 211155, 211156 [holotype], 211167–211176,

211178–211184, 211190, 211192–211198); Santa Cruz, El Refugio

Huanchaca ([loc. 4] 14846002 00S, 61802005 00W—USNM 584502). The

material reported by Anderson (1997) from the vicinities of the type

locality—near Espıritu ([loc. 3] » 148130S, 668400W) and Curiraba

River ([loc. 2] 148500S, 668230W)—were not examined but we

assumed they correspond to M. midastactus.

Myotis simus (n ¼ 79; 45 // [including 12 subadults], 30 ??

[including 10 subadults], 4 sex undetermined): BRAZIL: Amazonas,

Borba ([loc. 10] 068530S, 528020W—AMNH 91886–91892, 94224,

94225, 94227, 94230–94234); Amazonas, Itacoatiara ([loc. 9] 038080

S, 588260W—MZUSP 3472); Amazonas, Manaus ([loc. 11] 01840S,

638360W—AMNH 79534, 91472–91478, 91500); Amazonas, Par-

intins ([loc. 8] 028380S, 568440W—AMNH 92983, 93489–93497,

93922–93925); Amazonas, Rio Jurua ([loc. 12] 048480S, 688670W—

MZUSP 638, 1074); Mato Grosso, Taiama ([loc. 6] 168480S,

578280W—MZUSP 13815); Mato Grosso, Barao de Melgaco ([loc.

7] 168430S, 568110W—MN #); Mato Grosso do Sul, Salobra ([loc. 5]

218580S, 568310W—ALP 9277); Unknown locality (MZUSP 1062).

COLOMBIA: Amazonas, unknown locality (TTU 9073, 9076–9078).

ECUADOR: Quito, Pastaza ([loc. 15] 18270S, 768400W—AMNH

71483, 71485–71488, 71490–71494). PERU: Loreto, Maynas ([loc.

14] 03819 0S, 728070W—AMNH 74105, 74109, 74110, 74378–

74381); Loreto, Ucayali ([loc. 16] 068440S, 758060W—AMNH

76240–76249, 76252, 76253 [topotypes of M. simus]); Pasco, San

Juan ([loc. 17] 108300S, 748530W—USNM 364481, 364482).

Myotis cf. simus (literature records): ARGENTINA: Corrientes,

Ituzaingo, Isla Apipe ([loc. 24] 278300S, 568530W—Barquez et al.

1999); Formosa, Pilcomayo, Parque Nacional Rıo Pilcomayo ([loc.

20] 258000S, 588150W—Barquez et al. 1999); Formosa, Pilcomayo,

Estancia Las Mercedes ([loc. 21] 258110S, 578540W—Barquez et al.

1999). COLOMBIA: Amazonas, Leticia ([loc. 13] 04809 0S,

698570W—Gardner 2008). PARAGUAY: Presidente Hayes, Villa

Hayes 230 km by road NW ([loc. 19] » 238230S, 568500W—Lopez-

Gonzalez 2005; Gardner 2008); Central, Luque, 17 km by road E

([loc. 22] » 258160S, 578260W—Lopez-Gonzalez 2005; Gardner

2008); Misiones, Ayolas, Barrio San Antonio ([loc. 23] 27823.420S,

56850.150W—Lopez-Gonzalez 2005). PERU: Ucayali, Balta, Cash-

inaua indian village ([loc. 18] 108080S, 718130W—Voss and Emmons

1996).


a new species of myotis (chiroptera, vespertilionidae) from bolivia

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