Copeia 2012, No. 1, 37–48
A New Blind Snake Species of the Genus Tricheilostoma from Espinhaço
Range, Brazil and Taxonomic Status of Rena dimidiata (Jan, 1861)
(Serpentes: Epictinae: Leptotyphlopidae)
Roberta R. Pinto1 and Ronaldo Fernandes1
Designation and description of a neotype and geographic variation of the blind snake Rena dimidiata are presented here
on the basis of a noted sample, providing a question on its generic identity, and proposing a new taxonomic
combination. Rena dimidiata is recognized here as a species of Tricheilostoma, based on putative characters published in a
recent paper, additional external traits and hemipenial characters. Morphometric and meristic characters for all species
of Tricheilostoma are described herein. Additionally, a new species of Tricheilostoma is described from Espinhaço Range,
Brazil.
T
HE purpose of redescription in leptotyphlopid species is mostly explained by the rare exploration of
morphologic characters in different genera and quite
brief historical descriptions. Some recent work illustrates the
need for taxonomic revisions at the generic level and
presents novel insights into a number of putative characters
(Passos et al., 2006; Broadley and Wallach, 2007; Franco and
Pinto, 2009; Pinto and Curcio, 2011).
Adalsteinsson et al. (2009) proposed a molecular phylogeny of the Leptotyphlopidae and resurrected some genera to
recognize the clades within a monophyletic framework.
Most of the Neotropical clades (tribe Epictini, subfamily
Epictinae) correlate with species groups proposed by OrejasMiranda (1967) or Peters and Orejas-Miranda (1970). The
former Leptotyphlops dulcis species group (sensu Peters and
Orejas-Miranda, 1970) was split in two resurrected genera:
Rena Baird and Girard, 1853 and Tricheilostoma Jan, 1860.
According to Adalsteinsson et al. (2009), species of Tricheilostoma are distinguished from species of Rena by having a
brown or pale brown (not white) venter, three supralabials
(also present in Rena bressoni, R. dissecta, and R. myopica),
and a lower average number of middorsal scales. Adalsteinsson et al. (2009) did not analyze all species of both genera in
their phylogenetic hypothesis, so several diagnosed characters are homoplastic when all species of both genera are
taken into account. Passos et al. (2006) and Pinto and Curcio
(2011) proposed some putative characters based on all
species of Tricheilostoma (narrow basal and robust terminal
portions of hemipenial body, middorsal cephalic scales of
moderate size, and enlarged terminal spine). None of those
characters were analyzed in the Adalsteinsson et al. (2009)
phylogeny. So, at present we recognize only the characters
proposed by Passos et al. (2006) and Pinto and Curcio (2011)
to diagnose the genus Tricheilostoma.
Stenostoma dimidiata (5Rena dimidiata) was proposed by
Jan (1861) in a brief description based on a single
unnumbered specimen from ‘‘Brésil’’ (5Brazil). Jan and
Sordelli (1861) figured the holotype and a skull of the
species. Peters and Orejas-Miranda (1970) restricted the type
locality to São Marcos, near the confluence of the Uraricuera
and Tacutu rivers, both tributaries of the Branco River, Boa
Vista Municipality, State of Roraima, Brazil. However, no
type specimen was cited by Peters and Orejas-Miranda
(1970). Hahn (1980) reported that the holotype of Rena
1
dimidiata was deposited in the Museo Cı́vico di Storia
Naturale di Milano and it was later destroyed. According
to official site of the institution (http://www.comune.
milano.it/), all of Jan’s types described from 1853 to 1866
were destroyed, probably during World War II. Actually, the
species is known from two populations (Amazon-Guiana
and Espinhaço Range Region) separated by a large geographic hiatus (Silveira, 2004; Pinto, 2010). Although there
are some data available (Passos et al., 2006; Adalsteinsson
et al., 2009), no evaluation of taxonomic status, hemipenis
description or geographic variation has been proposed.
The absence of a type and an adequate description do not
permit the recognition of the population described by Jan
(1861). For this reason, in agreement with ICZN (1999), we
hereby designate a neotype for Rena dimidiata ( Jan, 1861)
based on the type locality restriction of Peters and OrejasMiranda (1970). Also, we provide data on sexual dimorphism and geographic variation in R. dimidiata and discuss
the generic identity of Stenostoma dimidiata Jan, 1861
(presently in Rena, sensu Adalsteinsson et al., 2009) based
on morphological data. Lastly, we describe a new species of
the genus Tricheilostoma from transitional areas of Tropical
Rainforest and Savanna Domains (Ab’Sáber, 2003; Drummond et al., 2005), and typical Savanna Domain of the
Espinhaço Range region in Brazil.
MATERIALS AND METHODS
Descriptions and comparisons are mainly based on external
morphology; in addition to examination of specimens,
literature data were also considered. Terminology for
cephalic plates, scale features, and measurements follows
Pinto and Curcio (2011), Broadley and Wallach (2007), and
Passos et al. (2006). Meristic and morphometric data follow
Pinto and Curcio (2011). Supralabial terminology follows
Passos et al. (2006), in which an equation is given for
supralabials before ocular scales and supralabials after ocular
scales.
Measurements were taken with a dial caliper to the nearest
0.1 mm, except for total length (TL) and tail length (TAL),
both taken with a graduated ruler to the nearest 1.0 mm.
Variation is expressed by the range, providing the mean
value followed by the standard deviation in parentheses. Sex
was determined through a ventral incision on the base of
Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Rio de Janeiro, Rio de Janeiro 20940-040, Brazil;
E-mail: (RRR) [email protected]; and (RF) [email protected]. Send reprint requests to RF.
Submitted: 23 March 2011. Accepted: 19 October 2011. Associate Editor: B. Stuart.
DOI: 10.1643/CH-11-040
F 2012 by the American Society of Ichthyologists and Herpetologists
38
Copeia 2012, No. 1
the tail. Hemipenes were everted and prepared according to
Pesantes (1994); hemipenis nomenclature follows Branch
(1986).
Sexual dimorphism was tested in some variables (middorsal, midventral, and subcaudal scales) using analysis of
variance (ANOVA) as well as comparative analyses in the
two species: Tricheilostoma dimidiatum and the new species
(Zar, 1999). Assumptions of univariate normality and
homoscedasticity were evaluated using the test of KolmogorovSmirnov and the Levene’s test, respectively (Zar, 1999). All
statistical inference was performed in STATISTICA 7.0 for
Windows (Statistica for Windows version 7.0. Statsoft, Inc.,
Tulsa, Oklahoma, 2004).
Institutional abbreviations follow the list available at
http://www.asih.org/node/204. Acronyms that were absent
or recently changed from the list are as follows: Museu de
História Natural Professor Doutor Adão José Cardoso
(ZUEC), Campinas, Brazil; Instituto Butantan (IBSP), São
Paulo, Brazil; Laboratório de Zoologia de Vertebrados (LZV),
Universidade Federal de Ouro Preto, Ouro Preto, Minas
Gerais, Brazil; Fundação Ezequiel Dias (FUNED), Belo
Horizonte, Minas Gerais, Brazil; Coleção Herpetológica da
Universidade Federal de Minas Gerais (UFMG), Belo Horizonte, Minas Gerais, Brazil; Museu de Zoologia João Moojen
de Oliveira, Universidade Federal de Viçosa (MZUFV),
Viçosa, Minas Gerais, Brazil; Coleção Herpetológica da
Universidade de Brası́lia (CHUNB), Brası́lia, D.F., Brazil;
Museu de Zoologia da Universidade Estadual de Feira de
Santana (MZUEFS), Brazil.
Tricheilostoma dimidiatum (Jan, 1861), new combination
Figures 1, 2
Stenostoma dimidiatum Jan, 1861:188.
Stenostoma dimidiatum Jan and Sordelli, 1861:fig. 4, plates V
and VI, holotype and skull.
Glauconia dimidiata.—Boulenger, 1893:64–65 (first combination in the genus Glauconia, description, distribution).
Leptotyphlops dimidiata.—Amaral, 1930 [dated 1929]:139 (first
combination in the genus Leptotyphlops, distribution).
Leptotyphlops dimidiatus.—Orejas-Miranda, 1967:433–435
(correct spelling, description, distribution).
Rena dimidiata.—Hedges, Adalsteinsson, and Branch in
Adalsteinsson et al., 2009:11, 20 (first combination in
the genus Rena).
Neotype.—IBSP 24011 (Fig. 1), adult female, Brazil, State of
Roraima, Boa Vista Municipality, São Marcos, 03u059N,
60u259W, elevation 75 m, Alphonse Richard Hoge expedition, 27 May 1964.
Diagnosis.—Tricheilostoma dimidiatum can be distinguished
from other congeneric species by the following combination
of characters: snout rounded in dorsal and ventral views,
obtusely rounded in lateral view; supraocular present; ocular
subhexagonal with anterior border slightly rounded at eye
level and superior border straight; rostral subtriangular in
dorsal view, not reaching (67%) or reaching (33%) anterior
border of ocular scales; frontal longer than other middorsal
cephalic shields, slightly smaller than supraocular; temporal
not distinct (74%) or distinct (26%); two supralabials (1+1);
four infralabials; robust body width (43.1 6 8.9); 198–215
middorsal in females and 191–214 in males; 185–200 ventral
in females and 175–197 in males; 13–16 subcaudal in
females and 12–18 in males; fused caudals present; 10 scales
Fig. 1. Dorsal (A), lateral (B), and ventral (C) views of the head of
the neotype of Tricheilostoma dimidiatum (IBSP 24011). Scale 5
5 mm.
around middle of tail; and seven dorsal scales rows with pale
brown pigmentation in contrast with seven unpigmented
ventral rows that are cream.
Description of neotype.—Adult female, TL 222 mm, TAL
15 mm; MB 5.1 mm; MT 3.0 mm; TL/TAL 14.8; TL/MB 43.5;
TAL/MT 5.0; HL 6.5 mm, HW 3.4 mm; relative eye diameter
2.4; relative rostral width 0.3. Body subcylindrical, robust,
slightly enlarged on the head and slightly tapered caudally
near tail.
Snout rounded in dorsal and ventral views, obtusely
rounded in lateral view; rostral straight in frontal and
ventral views, dorsal apex subtriangular, not reaching
imaginary transverse line between anterior border of ocular;
rostral contacting supranasal and infranasal laterally, and
frontal dorsally; nasal completely divided horizontally by
Pinto and Fernandes—New Tricheilostoma from Espinhaço Range, Brazil
39
Fig. 2. Geographic distribution of Tricheilostoma dimidiatum (circle) and T. jani (square). Triangle represents the record from Hoogmoed, 1977.
Type localities correspond to open symbols.
oblique suture crossing nostril; nostril roughly elliptical,
obliquely oriented and placed in middle of nasal suture;
supranasal nearly twice as high as long, bordering rostral
anteriorly, infranasal inferiorly, first supralabial and
ocular posteriorly, and frontal and supraocular dorsally;
supranasal base as long as upper border of infranasal scale;
infranasal almost twice as tall as long; upper lip border
formed by rostral, infranasal, anterior supralabial, ocular,
and posterior supralabial; temporal single, not distinct in
size from dorsal scales of lateral rows; two supralabials,
entirely separated by ocular (1+1); anterior supralabial
(first supralabial) as high as long, slightly exceeding nostril
and eyes inferior limits; posterior supralabial (second
supralabial) nearly twice as high as long, crossing nostril
and eye inferior limits, its posterior margin in broad
contact with temporal; ocular enlarged, subhexagonal,
superior border straight, slightly rounded at eye level,
almost twice as tall as long, contacting posterior margins
of supranasal and first supralabial anteriorly, parietal and
second supralabial posteriorly, and supraocular dorsally;
eye distinct (diameter 5 0.5 mm), placed in central area of
expanded upper part of ocular, displaced above nostril
level; supraocular nearly twice as long as wide, slightly
longer and not as wide as frontal, between ocular and
frontal, contacting supranasal anteriorly, frontal and
ocular laterally, and parietal and postfrontal posteriorly;
middorsal cephalic shields (postfrontal, interparietal, and
interoccipital) subequal in size, subhexagonal in dorsal
view, weakly imbricate; frontal longer than wide, longer
than other middorsal cephalic shields, slightly smaller
than supraocular, contacting rostral, supranasal, supraocular, and postfrontal; postfrontal longer than wide,
contacting frontal, supraocular, parietals, and interparietal; interparietal longer than wide, contacting postfrontal, parietals, occipitals and interoccipital; interoccipital
wider than long, contacting interparietal, occipitals, and
first dorsal scale of the vertebral row; parietal and occipital
subequal in shape, irregularly hexagonal; parietal two
times wider than long, lower margin contacting upper
border of second supralabial, posterior margin contacting
respective temporal, occipital, and interparietal, anterior
border in broad contact with ocular, supraocular, and
postfrontal; occipital almost twice wider than long, its
lower limit attaining upper margin of second supralabial,
though separated of latter by temporal; symphysial
trapezoidal, anterior and posterior borders respectively
straight and slightly convex, twice as wide as long; four
infralabials behind symphysial on both sides; first three
infralabials subequal, somewhat taller than long, not
pigmented; third infralabial wider than first two; fourth
infralabial distinctively longer than others, nearly three
times as long as wide, as long as third supralabial base, not
pigmented; head subcylindrical slightly distinguishable
from neck, almost twice as long as wide.
Middorsal scales 202; midventral scales 189; 14 scales rows
around midbody, reducing to ten rows in middle of tail;
cloacal shield triangular, almost three times as wide as long;
14 subcaudals; fused caudals present; terminal spine large,
conical, twice as long as wide. Dorsal scales homogeneous,
cycloid, smooth, weekly imbricate, and nearly twice as wide
as long.
Coloration in preservative (in alcohol).—Seven dorsal scale
rows are uniform pale brown, whereas the seven remaining
ventral scale rows are whitish-cream; at least the lower
margins of the scales that form the upper lip border are
cream colored, following the color of the venter; cloacal
shield light brown, but slightly lighter than dorsal pattern
and darker than general ventral tonality; terminal spine
follows the ventral pattern.
40
Variation.—Middorsal scales 198–215 (205.6 6 4.2, n 5 23)
in females and 191–214 (202.5 6 6.6, n 5 8) in males;
midventral scales 185–200 (191.5 6 4.0, n 5 20) in females
and 175–197 (187.8 6 7.3, n 5 8) in males; subcaudals 13–16
(14.8 6 1.0, n 5 23) in females and 12–18 (16.3 6 2.2, n 5 8)
in males; TL 94–281 mm (211.6 mm 6 45.7, n 5 22) in
females and 26–233 mm (147.3 mm 6 72.6, n 5 7) in males;
TL/TAL ratio 13.3–19.0 (15.1 6 1.3, n 5 23) in females and
5.2–16.8 (12.2 6 3.4, n 5 8) in males; TAL 5.3–7.5% of the TL
(6.7% 6 0.0, n 5 23) in females and 5.9–19.2% (9.1% 6 0.0,
n 5 8) in males; TL/MB ratio 32.9–59.9 (44.7 6 6.4, n 5 23)
in females and 12.5–56.6 (38.5 6 13.4, n 5 8) in males; TAL/
MT ratio 2.4–5.9 (4.0 6 0.7, n 5 23) in females and 2.8–6.0
(4.5 6 1.2, n 5 8) in males; relative eye diameter 1.5–2.9 (2.0
6 0.4, n 5 22) in females and 1.9–2.1 (2.0 6 0.1, n 5 5) in
males; relative rostral width 0.3–0.5 (0.4 6 0.0, n 5 21) in
females and 0.3–0.4 (0.4 6 0.0, n 5 7) in males.
Sexual dimorphism.—Males have significantly more subcaudal scales (F1,29 5 6.1, P , 0.05) than females. No
dimorphism was found in middorsal scales (F1,29 5 2.4, P
5 0.13) or midventral scales (F1,26 5 3.0, P 5 0.09). Females
have a greater relative total length (F1,29 5 11.7, P , 0.001)
than males.
Distribution and habitat.—Brazilian Amazonian lowlands of
the Guiana region and Surinam (Fig. 2). The neotype was
found on the ground under leaves of the ‘‘Buriti’’ palm.
Hoogmoed (1977) found this species in dry regions in a
transition area of Savanna Domain and lower vegetation in
Surinam, suggesting that the species has a Centro-Roraima
distribution as suggested by Müller (1973). Peters and
Orejas-Miranda (1970) recorded this species in southeastern
Venezuela. However, no voucher specimen in this study was
found from this region, so this record is probably doubtful.
Remarks.—Boulenger (1893) listed the species (Glauconia
dimidiata) in a previous description mostly translated from
the original description of Jan (1861). Orejas-Miranda (1967)
redescribed the species (Leptotyphlops dimidiatus) based on
several individuals from septentrional Brazil and Guiana.
The type locality restriction of Peters and Orejas-Miranda
(1970) was not based on any information from the original
description or additional specimens. Possibly, the specimens
studied by Orejas-Miranda (1967) are from São Marcos, Boa
Vista municipality, Roraima State (IBSP 24008–11), which
perhaps influenced the decision made by Peters and OrejasMiranda (1970).
Adalsteinsson et al. (2009) recognized Leptotyphlops dimidiatus as a member of the genus Rena (R. dimidiata),
although the species was not included in their phylogenetic
analysis. According to them, the genus Rena is differentiated
from Tricheilostoma by its higher number of middorsal scales
(on average), white venter (vs. light brown or brown), and
usually two supralabials, so Leptotyphlops dimidiatus was
attributed to the genus Rena. Adalsteinsson et al. (2009)
stated that the average number of middorsals is given in
their table 1; however, in the table only the range is
presented: in Tricheilostoma 152–253 and in Rena 168–312.
Furthermore, both a lower supralabial number and white
venter coloration can be present in each genus. Pinto and
Curcio (2011) cited putative characters based on the
external morphology of Tricheilostoma, such as middorsal
cephalic scales of moderate size and an enlarged terminal
Copeia 2012, No. 1
spine. Based on the characters elucidated by Pinto and
Curcio (2011), we propose the allocation of Rena dimidiata
(sensu Adalsteinsson et al., 2009) to the genus Tricheilostoma
(Tricheilostoma dimidiatum, new combination).
All specimens studied by Orejas-Miranda (1967) were
analyzed in the present study, including additional specimens not seen by that author. We also examined the
specimen cited by the author from Cotinga River, Limão
municipality, State of Roraima, Brazil (AMNH 36065)
identified as Rena affinis (Boulenger, 1884). This individual
is in fact a Tricheilostoma dimidiatum (see Comparisons).
Orejas-Miranda (1967) suggested that R. affinis may be a
synonym of T. dimidiatum or R. affinis could represent a
subspecies of T. dimidiatum.
During this study we found a large geographic hiatus in
the sample, forming two populations: one in GuianaAmazon with specimens just in the north of Amazon near
the Guiana region, the other in Espinhaço Range, Savanna
Domain, and transitional areas of Tropical Rainforest and
Savanna Domain in Minas Gerais State, Brazil (1000–
1750 m).
Hahn (1980) noted that the holotype of Tricheilostoma
dimidiatum was destroyed. In fact, according to the official
website of the Museo Cı́vico di Storia Naturale di Milano, all
Jan’s types deposited during 1853–1866 were destroyed,
probably during World War II. According to the International Code of Zoological Nomenclature, a neotype designation is justified when no name-bearing type specimen is
believe to be extant and an author considers that a namebearing type is necessary to define the nominal taxon
objectively (ICZN, 1999:84, article 75.1). Under these
conditions, a neotype is validly designated when a statement of the characters that the author regards as differentiating from other taxa the nominal species-group taxon for
which the neotype is designated (ICZN, 1999:84, article
75.3.2). Also, when there is evidence that the name-bearing
type specimen was lost or destroyed (ICZN, 1999:84–85,
article 75.3.4). Based upon these arguments, we believed
that the designation of a neotype will bring stability to the
name T. dimidiatum. The criterion adopted here to choose a
neotype follows the restriction of the type locality by Peters
and Orejas-Miranda (1970).
Tricheilostoma jani, new species
Figures 2–7
Leptotyphlops dimidiatus.—Silveira, 2004:411 (distribution).
Leptotyphlops dimidiatus.—Passos et al., 2006:349 (morphometric and meristic variations).
Rena dimidiata.—Hedges, Adalsteinsson, and Branch in
Adalsteinsson et al., 2009:11, 20 (first combination in
the genus Rena).
Holotype.—MNRJ 4263 (Figs. 3–4), adult male, Brazil, State of
Minas Gerais, Belo Horizonte Municipality, Parque das
Mangabeiras, 19u559S, 43u569W, elevation ca. 1000 m, G.
Kisteumacher, 1985.
Paratypes.—(n 5 21, all from Brazil, State of Minas Gerais):
MCN-R 1912, adult female, Municipality of Belo Horizonte,
Santo Antônio, 19u579S, 43u569W, C. E. Benfica, 3 May 2005;
MCN-R 3647, juvenile female, Municipality of Belo Horizonte, Santo Antônio, Barragem Santa Lúcia, 19u579S,
43u569W, C. E. Benfica, 29 October 2002; UFMG 245,
Pinto and Fernandes—New Tricheilostoma from Espinhaço Range, Brazil
41
Fig. 3. Dorsal (A), lateral (B), and ventral (C) views of the head of the
holotype of Tricheilostoma jani (MNRJ 4263). Scale 5 5 mm.
juvenile male, Municipality of Caeté, Serra da Piedade,
19u499S, 43u409W, ca. 1750 m elevation, C. S. Azevedo, 10
May 1999; UFMG 246, juvenile male, Municipality of Caeté,
Serra da Piedade, 19u499S, 43u409W, ca. 1750 m elevation,
C. S. Azevedo, 10 May 1999; UFMG 247, juvenile male,
Municipality of Caeté, Serra da Piedade, 19u499S, 43u409W,
ca. 1750 m elevation, C. S. Azevedo, 10 May 1999; MZUSP
8055, juvenile male, Municipality of Caeté, Serra da Piedade,
19u549S, 43u439W, M. T. Rodrigues, 10 January 1982, field
number MTR 82.0031; MZUSP 7583 (everted hemipenis),
adult male, Municipality of Santana do Riacho, Alto Palácio,
MG-010, km 115, 19u159S, 43u329W, ca. 1200 m elevation,
M. T. Rodrigues, 8 June 1980, field number MTR 80.0716;
MZUSP 7584, adult female, Municipality of Santana do
Riacho, Alto Palácio, MG-010, km 125, 19u159S, 43u329W, ca.
1250 m elevation, M. T. Rodrigues, 8 June 1980, field
number MTR 80.0717; MZUSP 18526, adult female, Parque
Nacional da Serra do Cipó, 19u159S, 43u339W, ca. 1345 m
elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira, Jr., R. S.
Recoder, and M. A. Sena, 17 November 2010, field
number MTR 19478; MZUSP 18527, adult female, Parque
Fig. 4. Photographs of the holotype of Tricheilostoma jani
(MNRJ 4263) in dorsal (A), lateral (B), and ventral (C) views of the
head. Scale 5 5 mm.
Nacional da Serra do Cipó, 19u169S, 43u319W, ca. 1334 m
elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira, Jr., R. S.
Recoder, and M. A. Sena, 17 November 2010, field
number MTR 19499; MZUSP 18528, adult male (everted
hemipenis), Parque Nacional da Serra do Cipó, 19u169S,
43u319W, ca. 1334 m elevation, M. T. Rodrigues, J.
Cassimiro, M. Teixeira, Jr., R. S. Recoder, and M. A. Sena,
17 November 2010, field number MTR 19500; MZUSP
18529, adult female, Parque Nacional da Serra do Cipó,
19u169S, 43u319W, ca. 1320 m elevation, M. T. Rodrigues, J.
Cassimiro, M. Teixeira, Jr., R. S. Recoder, and M. A. Sena, 17
November 2010, field number MTR 19501; MZUSP 18530,
juvenile female, Parque Nacional da Serra do Cipó, 19u169S,
42
Copeia 2012, No. 1
Fig. 5. Tricheilostoma jani in life in dorsal view (A), lateral view (B–C), and close detail of the head (D). Photos M. Teixeira, Jr.
43u319W, ca. 1320 m elevation, 17 November 2010, M. T.
Rodrigues, J. Cassimiro, M. Teixeira, Jr., R. S. Recoder, and
M. A. Sena, 17 November 2010, field number MTR 19502;
MZUSP 18531, adult male (everted hemipenis), Parque
Nacional da Serra do Cipó, 19u169S, 43u319W, ca. 1340 m
Fig. 6. Hemipenis of Tricheilostoma jani (MZUSP 7583) in asulcate (A)
and sulcate (B) sides. Scale 5 5 mm.
elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira, Jr., R. S.
Recoder, and M. A. Sena, 17 November 2010, field
number MTR 19510; MZUSP 18532, adult male (everted
hemipenis), Parque Nacional da Serra do Cipó, 19u159S,
43u339W, ca. 1317 m elevation, M. T. Rodrigues, J. Cassimiro,
M. Teixeira, Jr., R. S. Recoder, and M. A. Sena, 17 November
2010, field number MTR 19511; MZUSP 18533, juvenile male,
Parque Nacional da Serra do Cipó, 19u159S, 43u339W, ca.
1358 m elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira,
Jr., R. S. Recoder, and M. A. Sena, 18 November 2010, field
number MTR 19521; MZUSP 18534, adult male, Parque
Nacional da Serra do Cipó, 19u159S, 43u339W, ca. 1339 m
elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira, Jr., R. S.
Recoder, and M. A. Sena, 20 November 2010, field
number MTR 19581; MZUSP 18535, adult male (everted
hemipenis), Parque Nacional da Serra do Cipó, 19u159S,
43u339W, ca. 1339 m elevation, M. T. Rodrigues, J. Cassimiro,
M. Teixeira, Jr., R. S. Recoder, and M. A. Sena, 20 November
2010, field number MTR 19582; MZUSP 18536, juvenile male,
Parque Nacional da Serra do Cipó, 19u179S, 43u359W, ca.
1289 m elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira,
Jr., R. S. Recoder, and M. A. Sena, 21 November 2010, field
number MTR 19606; MZUSP 18537, adult male, Parque
Nacional da Serra do Cipó, 19u159S, 43u339W, ca. 1345 m
elevation, M. T. Rodrigues, J. Cassimiro, M. Teixeira, Jr., R. S.
Recoder, and M. A. Sena, 28 November 2010, field
number MTR 19712; ZUEC 1227, adult male, Serra do Cipó,
Municipality of Santana do Riacho, 19u109S, 43u429W, A. A.
Giaretta, 9 November 1991.
Diagnosis.—Tricheilostoma jani can be distinguished from
other congeneric species by the follow combination of
characters: snout truncate in dorsal and ventral view,
rounded in lateral view; supraocular present; ocular subhex-
Pinto and Fernandes—New Tricheilostoma from Espinhaço Range, Brazil
Fig. 7. Habitat of Tricheilostoma jani in Savanna Domain of Serra do
Cipó, Minas Gerais. An overview of the Serra do Cipó (A), their ‘‘campos
rupestres’’ or rocky meadows (B), and the microhabit of T. jani (C).
Photos J. Cassimiro.
agonal with anterior border slightly rounded at eye level, and
superior border straight; rostral triangular in dorsal view, not
reaching (65%) or reaching (35%) anterior border of ocular
scales; frontal as long as supraocular and other middorsal
cephalic shields; temporal not distinct (50%) or distinct
(50%); two supralabials (1+1); four infralabials; slender body
width (37.7 6 4.5); 176–206 middorsal in females and 180–
197 in males; 160–188 ventral in females and 161–180 in
males; 14–18 subcaudal in females and 15–20 in males; fused
caudals present; ten scales around middle of tail; and seven
dorsal scales rows with uniform pale brown pigmentation,
and the seven unpigmented ventral rows, cream.
Comparisons with the Neotropical region leptotyphlopids.—
Neotropical region is represented by four genera of the
43
Leptotyphlopidae family (Epictia, Rena, Tricheilostoma, and
Siagonodon). In this comparison we will provide generic and
specific characters to differ the new species from all those
species in the Neotropical region.
The presence of supraocular scales distinguishes the new
species from all species of Siagonodon, and from Tricheilostoma brasiliensis (Pinto and Curcio, 2011). Tricheilostoma
jani can be distinguished from T. dugandi and some species
of Epictia, except for E. undecimstriata, E. signata, E. columbi,
E. nasalis, E. peruviana, and E. collaris by having no stripes in
the dorsal coloration (Orejas-Miranda, 1969; Hoogmoed,
1977; Cei, 1986, 1994; Pinto et al., 2010; Pinto and Curcio,
2011). The presence of fused caudals distinguishes T. jani
from T. guayaquilensis and most Siagonodon (except S.
borrichianus), and species of Epictia (except E. tenella; Pinto
and Curcio, 2011). A subhexagonal ocular with its border
anteriorly rounded at the eye level distinguishes T. jani from
all species of genera Siagonodon and Epictia. Four infralabials
(vs. three) distinguish the new species from R. unguirostris, S.
borrichianus, and T. nicefori (Pinto et al., 2010). Ten dorsal
scales around the middle of the tail (vs. 12 or 14) separate
the new species from T. joshuai, S. acutirostris, S. septemstriatus, E. albipuncta, E. striatula (12), and S. cupinensis (14)
(Kretzschmar, 2006; Pinto et al., 2010; Pinto and Curcio,
2011). Tricheilostoma jani is distinguished from R. affinis and
R. unguirostris, and all species of Tricheilostoma, except T.
dimidiatum and T. nicefori by having two supralabials (1+1)
instead of three supralabials (2+1). The presence of a
truncate snout in dorsal and ventral views (vs. rounded
snout), slender body (vs. robust), and frontal with same size
as supraocular (vs. smaller) distinguishes the new species
from T. dimidiatum. A frontal scale smaller than the
supraocular separates T. dimidiatum from R. affinis (vs.
frontal with same size as supraocular). Tricheilostoma jani
can be distinguished from R. affinis by less than 210
middorsal scales (vs. 230). For meristic and morphometric
comparisons of all species of Tricheilostoma, see Table 1.
Individuals of Tricheilostoma dimidiatum have a higher
number of middorsal scales (Females: F1,27 5 51.7, P ,
0.0001; Males: F1,19 5 33.9, P , 0.0001) and midventral
scales (Females: F1,29 5 26.0, P , 0.0001; Males: F1,27 5 67.5,
P , 0.0001) than T. jani. Males of T. jani have a higher
number of subcaudals (F1,27 5 8.7, P , 0.01) than T.
dimidiatum. Moreover, females of T. dimidiatum exhibits a
greater proportion of body width (TL/MB; F1,32 5 17.0, P ,
0.001) than females of T. jani. The relative total length (TL/
TAL) is greater in T. dimidiatum (F1,32 5 28.6, P , 0.0001)
than in T. jani, indicating that the latter species has a smaller
proportional width than T. dimidiatum. However, females of
T. jani present a greater relative tail length (TAL % in TL;
F1,32 5 34.2, P , 0.0001) than T. dimidiatum.
Description of holotype.—Adult male, TL 187 mm, TAL
19 mm; MB 4.2 mm; MT 3.8 mm; TL/TAL 9.8; TL/MB
45.1; TAL/MT 5.1; HL 6.2 mm, HW 3.6 mm; relative eye
diameter 1.6; relative rostral width 0.3. Body subcylindrical,
slender, head slightly enlarged and slightly tapered caudally
near tail.
Snout truncate in dorsal and ventral views, rounded in
lateral view; rostral straight in frontal and ventral views,
dorsal apex triangular, reaching imaginary transverse line
between anterior border of ocular; rostral contacting
supranasal and infranasal laterally, and frontal dorsally;
nasal completely divided horizontally by oblique suture
(2)
(2)
(7)
(23)
(17)
(2)
(1)
(23)
(8)
(6)
(5)
(3)
(26)
(39)
(1)
(1)
(11)
(21)
(4)
(10)
(10)
(20)
(16)
(22)
(9)
(1)
(1)
(16)
(18)
(1)
F
M
?
F
M
M
?
F
M
?
F
M
F
M
?
?
F
M
F
M
F
M
F
M
?
M
?
F
M
?
T. anthracinum
T. salgueiroi
T. nicefori
T. macrolepis
T. koppesi
T. joshuai
T. jani
T. guayaquilensis
T. fuliginosum
T. dugandi
T. dimidiatum
T. brevissimum
T. brasiliensis
Sex (n)
Species
176–206
180–197
184–199
174–193
195–213
185–212
211–255
218–252
231–254
167
168
209–233
200–223
–
185–193
184–187
182–189
194–224
193–209
152–162
162
198–215
191–214
190–197
181–184
171–174
208–225
198–218
–
253
DO
160–188
161–180
172–187
169–181
182–196
169–195
213–233
201–237
207–233
153
–
188–213
183–207
–
170–176
167–168
–
178–212
171–196
141–152
151
185–200
175–197
174–186
158–160
172
191–208
173–201
210
233
VE
14–18
15–20
13–15
13–17
13–16
13–19
16–22
17–24
18–23
16
13
16–23
17–24
23
16–18
17–18
15–19
13–19
17–20
12–14
13
13–16
12–18
14–15
9–10
12–13
13–17
14–20
15
20
SC
10.3–14.1
7.4–13.7
13.6–24.6
10.8–17.0
13.1–24.2
9.9–16.9
11.3–15.8
8.5–15.9
8.2–13.8
12.9
14.7
10.0–14.5
5.7–12.3
32.4–34.5
36.9–38.3
12.2–16.6
11.3–15.7
9.2–14.6
12.6–13.2
12.6
13.3–19.0
5.2–16.8
14.0–17.4
18.4–30.0
15.5–19.3
11.9–18.4
10.1–14.8
11.7
13.1
TL/TAL
29.9–42.4
31.4–45.4
34.0–55.2
37.8–50.0
29.0–44.6
33.9–47.8
32.2–59.5
37.1–68.3
34.0–53.1
42.9
–
38.2–58.8
25.3–53.4
32.4–34.5
36.9–38.3
31.7–43.7
31.3–59.2
34.5–63.6
32.9–41.3
30.3
32.9–59.9
12.5–56.6
35.0–51.6
27.0–37.5
33.7–52.5
34.8–50.2
34.4–55.1
–
48.6
TL/MB
SL
3.0–5.3
3.2–6.2
2.9–3.5
2.8–3.6
1.7–4.5
2.9–4.8
3.8–5.7
4.4–6.7
4.8–7.9
–
–
4.0–5.8
4.9–7.4
4
3
4
2+1
1+1
2+1
present
4
4
present
4
2+1
present
4
4
present
4
2+1
present
4
present
present
present
present
present
present
absent
4
4
present
SO
4
IL
1+1
3.6–4.6
2+1
5.1–10.1
–
3.2–4.6 2+1 (90.4%)
3.7–5.6 1+1 (4.8%)
3.3
2+1
2.8
2.4–5.9
1+1
2.8–6.0
–
2.4
2+1
2.5–2.6
3.0–4.6
2+1
2.9–5.2
–
–
2+1
TAL/MT
10
10
10
10
12
10
12
10
10
10
10
10
10
Midtail
scales
1
3
5
3, 4
1
3
6
4
7
3
6
3, 4
2
Color
pattern
Passos et al., 2005, 2006;
Costa et al., 2009;
This paper
Passos et al., 2006;
Pinto et al., 2010
Passos et al., 2005,
2006; This paper
Passos et al., 2005, 2006;
Pinto et al., 2010;
This paper
Pinto et al., 2010
Orejas-Miranda and
Peters, 1970;
This paper
This paper
Passos et al., 2006;
Pinto et al., 2010
Hoogmoed, 1977; Passos
et al., 2005, 2006;
This paper
Passos et al., 2006;
Pinto et al., 2010
Passos et al. 2006;
This paper
Pinto and Curcio, 2011
Bailey, 1946; Passos et al.,
2006; This paper
Source of data
Table 1. Meristic and Morphometric Variation in Species of Tricheilostoma. DO = middorsal scales; VE = midventral scales; SC = subcaudals; SL = supralabials; IL = infralabials; SO = supraocular. Color
pattern modified from Passos et al. (2006): 1 = uniform dark brown dorsally and pale brown ventrally; 2 = uniform violet black dorsally and ventrally; 3 = uniform brown dorsally and cream ventrally; 4 =
uniform dark brown dorsally and cream ventrally; 5 = reticulate dark brown dorsally and reticulate pale brown ventrally; 6 = uniform brown dorsally and pale brown ventrally; 7 = brown with lighter
striping dorsally and cream ventrally.
44
Copeia 2012, No. 1
Pinto and Fernandes—New Tricheilostoma from Espinhaço Range, Brazil
crossing nostril; nostril roughly elliptical, obliquely oriented
and placed in middle of nasal suture; supranasal higher than
longer, bordering rostral anteriorly, infranasal inferiorly,
first supralabial and ocular posteriorly, and frontal and
supraocular dorsally; supranasal base as long as upper border
of infranasal scale; infranasal taller than long; upper lip
border formed by rostral, infranasal, anterior supralabial,
ocular, and posterior supralabial; temporal distinct in size
from dorsal scales of lateral rows; two supralabials, entirely
separated by ocular (1+1); first supralabial almost twice as
tall as long, exceeding nostril and eye levels, higher than
second supralabial; second supralabial taller than long,
crossing nostril and contact eye levels, its posterior margin
in broad contact with temporal; ocular enlarged, subhexagonal, superior border straight, slightly rounded at eye
level in anterior border, higher than long, contacting
posterior margins of supranasal and first supralabial anteriorly, parietal and second supralabial posteriorly, and
supraocular dorsally; eye distinct (diameter 5 0.9 mm),
placed in central area of expanded upper part of ocular,
displaced above nostril level; supraocular longer than wide,
as long as frontal, between ocular and frontal, contacting
supranasal anteriorly, frontal and ocular laterally, and
parietal and postfrontal posteriorly; middorsal cephalic
shields (frontal, postfrontal, interparietal, and interoccipital) subequal in size, pentagonal, weakly imbricate; frontal
longer than wide, contacting rostral, supranasal, supraocular, and postfrontal; postfrontal longer than wide, contacting frontal, supraocular, parietals, and interparietal; interparietal wider than long, contacting postfrontal, parietals,
occipitals, and interoccipital; interoccipital wider than long,
contacting interparietal, occipitals, and first dorsal scale of
vertebral row; parietal and occipital subequal in shape,
irregularly hexagonal; parietal twice as wide as long, lower
margin contacting upper border of second supralabial,
posterior margin contacting respective temporal, occipital,
and interparietal, anterior border in broad contact with
ocular, supraocular, and postfrontal; occipital almost twice
as wide as long, longer than parietal but less wide, its lower
limit attaining upper margin of second supralabial, though
separated of the latter by temporal; symphysial trapezoidal,
anterior and posterior borders respectively straight and
slightly convex, twice as wide as long; four infralabials; first
two infralabials subequal, somewhat twice as long as wide,
not pigmented; third infralabial wider than first two; fourth
infralabial not pigmented, distinctively longer than others,
almost twice as long as wide, slightly shorter than third
supralabial base; head subcylindrical, slightly distinguishable from neck, nearly two times longer than wide.
Middorsal scales 191; midventral scales 169; 14 scales rows
around middle of body, reducing to ten rows in middle of
tail; cloacal shield triangular, almost two times wider than
long; 20 subcaudals; fused caudals present; terminal spine
large, conical, longer than wide. Dorsal scales homogeneous, cycloid, smooth, weekly imbricate, and wider than
long. Femur claws are externally visible on both sides.
Coloration in preservative (in alcohol).—Seven dorsal scale
rows uniform pale brown, and seven ventral scale rows
cream or light brown pigmented in the center of ventral
scales; head coloration follows the dorsal color dorsally and
venter color ventrally, or spotted brown pigmented ventrally; lower margins of the scales that form the upper lip border
cream colored, as is the venter; cloacal shield cream,
45
following the ventral pattern; terminal spine follows the
body pattern.
Coloration in life.—Similar color pattern of body to preserved
specimens (Fig. 5). Seven dorsal scale rows uniform brown,
and seven ventral scale rows pale brown in center of scale
with cream border; lower margins of upper lip border cream
colored; cloacal shield following the belly pattern; terminal
spine follows the body pattern.
Hemipenis.—(Everted organ, n 5 5; Fig. 6). Organ single,
trumpet-shaped, narrow at base and robust at apex; hemipenial body ornamented with flounces; basal portion
covered by one flounce, sometimes small, slightly demarcated from terminal region by a distinctively large flounce
on mid-portion of the asulcate side; two well developed
flounces, and two other small flounces may be present
distally on terminal region of the asulcate side; sulcate side
with three to four flounces distributed from the middle to
the distal portion of the hemipenial body; sulcus spermaticus single, entering the organ on the basal surface and
extending towards the tip; sulcal folds raised and unadorned; apex of distal portion of the organ slightly
concave, covered by small papillae.
Variation.—Middorsal scales 176–206 (196.8 6 7.9, n 5 11)
in females and 180–197 (187.9 6 4.2, n 5 21) in males;
midventral scales 160–188 (180.7 6 7.8, n 5 11) in females
and 161–180 (169.3 6 4.6, n 5 21) in males; subcaudals 14–
18 (15.6 6 1.5, n 5 11) in females and 15–20 (18.3 6 1.5, n 5
21) in males; TL 93–262 mm (182.8 mm 6 58.3, n 5 11) in
females and 92–225 mm (182.0 mm 6 41.6, n 5 21) in
males; TL/TAL ratio 10.3–14.1 (12.6 6 1.2, n 5 11) in females
and 7.4–13.7 (10.2 6 1.3, n 5 21) in males; TAL 7.1–9.7% of
the TL (8.0% 6 0.0, n 5 11) in females and 7.3–13.4% (9.9%
6 0.0, n 5 21) in males; TL/MB ratio 29.9–42.4 (35.8 6 4.3, n
5 11) in females and 31.4–45.4 (37.0 6 3.9, n 5 21) in males;
TAL/MT ratio 3.0–5.3 (3.9 6 0.8, n 5 11) in females and 3.2–
6.2 (4.6 6 0.7, n 5 21) in males; relative eye diameter 1.4–2.3
(1.8 6 0.3, n 5 10) in females and 1.4–2.3 (1.7 6 0.2, n 5 20)
in males; relative rostral width 0.3–0.4 (0.4 6 0.0, n 5 10) in
females and 0.3–0.4 (0.4 6 0.0, n 5 20) in males.
Sexual dimorphism.—Females have a higher number of
middorsal scales (F1,30 5 17.9, P , 0.001) and midventral
scales (F1,30 5 27.7, P , 0.0001) than males; in contrast,
males have more subcaudals (F1,30 5 23.4, P , 0.0001) than
females. Males have greater diameter of tail in proportion of
tail length than females (F1,30 5 5.6, P , 0.05). Females
exhibit a greater relative total length (F1,30 5 24.1, P ,
0.0001) than males, but males have longest tail in
proportion to body length (F1,30 5 18.7, P , 0.001).
Distribution and habitat.—Tricheilostoma jani is known to live
in different aspects of the transition areas of Savanna and
Tropical Rainforest domains and Savanna Domain, ‘‘Campos Rupestres’’ (5montane savanna, rocky fields, or rocky
meadows) (Ab’Sáber, 2003; Drummond et al., 2005) in
southeastern Brazil. At the present, this new species is
limited to the Espinhaço Range region, Brazil (Fig. 2). The
records suggest that T. jani is a snake of Tropical Rainforest
and Savanna transition areas and typical Savanna Domain,
from rocky meadows, characteristic formations of elevations
from 1000–1750 m (Fig. 7). One specimen from municipality
46
of Mucugê, state of Bahia (MZUEFS 1232) apparently was
identified as the new species; however, this only specimen is
broken in parts, making the meristic and morphometric
analyses difficult, so we preferred not to consider it in this
study. As for most species of leptotyphlopids, the new species
is apparently strictly fossorial, constructing tunnels (MZUSP
7583–84; M. Rodrigues, pers. obs.) and can be found in
cavities under stones, alone or frequently in pairs. They
eventually come out when rain starts and their tunnels
apparently become flooded. Some individuals were caught
crossing the street during the rain (MCN-R 1912, 3647; C. E.
Benfica, pers. obs.).
Etymology.—The specific epithet is in homage to Giorgio Jan
(in memoriam), an important naturalist and curator of Museo
Cı́vico di Storia Naturale di Milano that contributed to the
knowledge of these rare snakes. Jan also proposed the name
Tricheilostoma as a subgenus of Stenostoma (oldest generic
name of leptotyphlopids) in Jan and Sordelli (1860) and Jan
(1861), recently resurrected by Adalsteinsson et al. (2009).
MATERIAL EXAMINED
Literature data are cited in parentheses.
Epictia albipuncta: Argentina: Tucumán, IBSP 58886; between Tucumán and Velez, USNM 164255; Salta: Yakulica,
MZUSP 8051.
Epictia australis: Argentina: Rı́o Negro: Valcheta, MACN
12526 (paratype); Buenos Aires: Tandil, USNM 345499.
Epictia goudotti: Colombia: Magdalena: Valley of Rı́o Magdalena, MNHN 1068 (holotype); Cienaga, USNM 144173;
Parque Nacional Isla de Salamanca, ICN 7127; PNN Tayrona,
Santa Marta, ICN 6196; Cundinamarca: Apulo, MLS 18;
Tolima: Ambalema, MLS 19; Honda, MLS 20. Without
specific locality, MLS 21 (Pinto et al., 2010).
Epictia magnamaculata: Colombia: Arquipielago de San
Andrés, Providencia y Santa Catalina, Isla de Providencia,
ICN 2629–31, ICN 11172, MLS 1972; Isla de San Andrés,
MLS 2038–40. Without specific locality, ICN 2730–34 (Pinto
et al., 2010).
Epictia munoai: Uruguay: Tacuarembó: Tambores, Pozo
Hondo, MNRJ 3306 (paratype; MHNM 068C).
Epictia nasalis: Nicaragua: Managua, USNM 16134 (holotype;
Pinto and Curcio, 2011).
Epictia signata: Colombia: Without specific locality, IBSP
7204. Ecuador: Gualaquiza: Morono-Santiago, USNM
232404 (Pinto et al., 2010).
Epictia striatula: Bolivia: Beni: Vaca Dı́ez, Tumi Chucua,
USNM 280427; Chuquisaca: Yotala, USNM 281801.
Rena affinis: Venezuela: Tachira, BMNH 75.2.26.4 (holotype).
Rena unguirostris: Argentina: Córdoba: Cruz Del Eje, BMNH
1946.1.11.52 (holotype); Catamarca: Capital, Águas Coloradas, FML 0689; Póman, Puesto Rı́o Blanco, FML 1399;
Tinogasta, Tinogasta, Palo Blanco, FML 1773; 35 km de
Tinogasta, Villa Luján (5Rio Colorado), FML 1910; Rı́o
Negro: Conesa, General Conesa, FML 1829; Salta: Anta,
Finca Pozo Largo, FML 2075, FML 2308; La Rioja: Castro
Barros, Chuquis, FML 9620; Paraguay: Chaco: Fortin
Guachalla, Rio Pilcomayo, FMNH 44174 (Pinto and Curcio,
2011).
Siagonodon borrichianus: Argentina: Patquia, Estancia de
Breyer, La Rioja, USNM 73500; Mendonza, MCZ 15900
(Pinto and Curcio, 2011).
Copeia 2012, No. 1
Tricheilostoma anthracinum: Ecuador: Baños, UMMZ 92531–
32, 88897; Abitagua, UMMZ 90817, 92533–35 (Bailey,
1946).
Tricheilostoma brasiliensis: Brazil: Maranhão: Carolina,
CHUNB 52064; Bahia: Barreiras, IBSP 50436, UMMZ
108817; Cocos, CHUNB 51368; Correntina, MNRJ 18392–
93; Piauı́: Estação Ecológica Uruçuı́-Una, MZUSP 12189–92;
Tocantins: Babaçulândia, MZUSP 12668; Palmeiras do
Tocantins, MZUSP 17713–15; Goiás: PCH Santa Edwiges I
[region of Mambaı́], MZUSP 17761–62; Posse, CHUNB
50870; Minas Gerais: Buritizeiro, CHUNB 44545; Formoso,
MZUSP 12881–85; João Pinheiro, MNRJ 17802; PARNA
Cavernas do Peruaçu, Januária, MZUSP 17716–25; Três
Marias, MNRJ 4616; Mato Grosso: APM-Manso, Chapada
dos Guimarães, UFMT 0683; Mato Grosso do Sul: Corumbá,
UFMT 1159–60, 1162–63 (Pinto and Curcio, 2011).
Tricheilostoma brevissimum: Colombia: Antioquia: Sonsón,
MCZ 38950 (paratype); Caquetá: Florencia, MLS 3, 1311
(holotype; Pinto et al., 2010).
Tricheilostoma dimidiatum: Surinam: Brokopondo: Railway
Km 62, RMNH 13479–80; Para: Powakka, RMNH 17835–36,
ZMAN 12974, CM 44284 (Hoogmoed, 1977).
New data.—Guiana: Upper Takutu-Upper Essequibo:
Lethem, FMNH 189352; Karanambo Ranch (river Rupununi), North Savanna Rupununi, Rupununi, USNM 566291;
between Wichabai and Isheartun, AMNH 60908; Saint
Ignatius, MCZ 115573. Brazil: Roraima: Marco fronteira
BV-8 (BR-174), MZUSP 8616, 8812; Boa Vista, MZUSP 9101,
9740–42, 9744, 9836, 9847, 9849–50, 9959, 10090, 10120;
Ilha de Maracá, MZUSP 9262–63; São Marcos, Boa Vista, IBSP
24008–10; Baixo Rio Mucujaı́, MZUSP 3731; Maloca Mangueira, MZUSP 9800; Fazenda Salvamento, MZUSP 9987;
Fazenda Monte Cristo, MZUSP 9846; Colônia do Apiauı́,
Igarapé Serrinha, MZUSP 9244; Rio Cotinga, Limão, AMNH
36065.
Tricheilostoma dugandi: Colombia: Atlántico: Barranquilla,
MCZ 58785, MLS 1992, 2337, ICN 2025–28, CBB unnumbered (paratype), CBB unumbered (holotype; Pinto et al.,
2010).
Tricheilostoma fuliginosum: Brazil: Tocantins: Lajeado, IBSP
64368 (paratype); Palmas, CHUNB 11538–39, 12014, 12550,
18319–20, 21854, 21955 (paratypes); U.H.E. Luı́s Eduardo
Magalhães, IBSP 64368 (paratype); U.H.E. Lajeados, IBSP
65050–55 (paratypes); Porto Nacional, IBSP 65358, 65581–
83, 66168 (paratypes); Goiás: Minaçú, CHUNB 6055–56,
6101 (paratypes), IBSP 56732–34 (paratypes); Peixe, IBSP
66101–02 (paratypes); Serra da Mesa, MZUSP 11017–18,
11020 (paratypes), 11019, MZUSP 11039–40 (paratypes);
Goiás: Minas Gerais: U.H.E. Queimados, MNRJ 10034
(holotype), MNRJ 10686, CHUNB 20348 (paratypes), MCNR 1019; Minas Gerais: Unaı́, CHUNB 20348, 30892, 32869
(paratypes; Passos et al., 2006).
New data.—Brazil: Tocantins: Palmas, MZUSP 11513–14;
U.H.E. Luı́s Eduardo Magalhães, MZUSP 14618, 14864,
15731; Pedro Afonso, CHUNB 50812; Porto Nacional, IBSP
65368, 65685. Goiás: Colinas do Sul, CHUNB 50422; Minaçú
CHUNB 6098; Luziânia, CHUNB 40846–48, 40746–47,
40749, 40789; Serra da Mesa, MZUSP 11041; Três Ranchos,
CHUNB 44726, 44750; Monte Alegre de Goiás, CHUNB
50869; São Domingos, CHUNB 40843; Rio Verde, CHUNB
43619. Without specific locality, CHUNB 24608.
Tricheilostoma guayaquilensis: Ecuador: Guaylas, Guayaquil,
ZMB 4508 (holotype).
Pinto and Fernandes—New Tricheilostoma from Espinhaço Range, Brazil
Tricheilostoma jani, new species: Brazil: Minas Gerais: Distrito
da Chapada, Ouro Preto, LZV 813S; Estação Ecológica do
Tripuı́, Ouro Preto, LZV 372S; Pioneiros, Ouro Branco, LZV
578S; Ouro Branco, LZV 778S; Caeté, MZUFV 750; Belo
Horizonte, FUNED 339; Ibirité, FUNED 356; Vespasiano,
MZUSP 2780. Without specific locality, FUNED 879, MNRJ
16990.
Tricheilostoma joshuai: Colombia: Antioquia: Jericó, MLS 11,
13 (holotype), 2646–47 (paratypes), UMMZ 84093, IBSP
7206, 7223, 8919, MHNCSJ 266; Cauca river, MLS 12
(paratype); San Juan river, IBSP 9188; without specific
locality, MLS 14 (paratype); Risaralda: Santa Rosa de Cabal,
MLS 2048; Valle del Cauca: San Antonio, MVZ 68688;
Caldas: Villamaria, MLS 15 (paratype); Salamina, MLS 1857;
Quindı́o: Salento, MLS unnumbered (Pinto et al., 2010).
Tricheilostoma koppesi: Brazil: Tocantins: Palmas, CHUNB
28150; Goiás: U.H.E. Corumbá, Mineiros, CHUNB 24548,
25712–16, 28151–52, IBSP 58708; São Paulo: Brotas, IBSP
43663; Itirapina, IBSP 58706; Mogi-Guaçú, ZUEC 577;
Pirassununga, IBSP 70032, MNRJ 11206; Mato Grosso do
Sul: Parnaı́ba, IBSP 45616–17; Terenos, IBSP 8883 (holotype;
Passos et al., 2005, 2006).
New data.—Brazil: Tocantins: Porto Alegre do Tocantins,
CHUNB 38928; Paranã, CHUNB 40813; Goiás: Caldas Novas,
MZUSP 11111–12; Mineiros, CHUNB 34117, 28152; PARNA
das Emas, MZUSP 15046; PCH Mosquitão, Caiapó river,
between Arenópolis and Iporá, MZUSP 17683; São Paulo:
Águas de Santa Bárbara, MZUSP 14976–77; Brotas, IBSP
45864.
Tricheilostoma macrolepis: without specific locality, ZMB
5722 (Paralectotype). Colombia: Antioquia: Capitan, Briceño, MHUA 14509; Santa Isabel, Remedios, MHUA 14059;
Caldas: Victoria, MHUA 14598, Norcasia, El Valle, MHUA
14615; Chocó: without specific locality, MCZ 39705;
Córdoba: Pueblo Nuevo, ICN 11337; Urrá, ICN 7677; Huila:
Palermo, MLS 2261; Norte de Santander: Ocaña, MLS 16;
Zulia, MLS 4; Santander: Bucaramanga, ICN 7299, MLS
1991; Sangil, IBSP 8588; Tolima: Chaparral, MLS 10, 1900;
Guamo, MLS 1; Valle del Cauca: Buenaventura, USNM
154031; Calima river, 10 km N El Pailón, USNM 267261;
near Cali, 1 km W Yumbo, MCZ 140118; Cali, USNM
151738; Vaupés: Estación Biológica Caparú, ICN 8142–43.
Venezuela: Carabobo: Puerto Cabello, ZMB 1434 (lectotype),
5294 (paralectotype; Pinto et al., 2010).
New data.—Venezuela: Carabobo, Puerto Cabello, ZMB 5722
(Paralectotype); El Vale, AMNH 59406, UIMNH 94016;
Caracas, USNM 62205, 107891; Vargas: Carayaca, IBSP
25801; Falcon: Acosta, MCZ 48752–73, 48918; Mérida:
Mérida, MCZ 18862. Brazil: Pará: Canindé, MZUSP 4286,
4308, MPEG 10167; Jacundá, MPEG 16725; São Domingos
do Capim, MPEG 8728; Santarém, MPEG 19043, 19094,
19109; Benevides, MPEG 8638; Tucuruı́, IBSP 46915, 47081;
Amazonas: Maués, MNRJ 385; Cururu river, MNRJ 386;
Rondônia: Ji-Paranã River, MZUSP 7317–18.
Tricheilostoma nicefori: Colombia: Santander: Charalá, ICN
5727; Mogotes, MLS 17 (holotype; Pinto et al., 2010).
Tricheilostoma salgueiroi: Brazil: Minas Gerais: Governador
Valadares, MZUFV 1216; Recreio, MNRJ 7856, 12239; São
José da Safira, MZUSP 14262–63; Aimorés, MCN-R 1468–69;
Paté, MZUFV 1214; Muriaé, MZUFV 1397, 1519; Espı́rito
Santo: Governador Lindemberg, MNRJ 12131–32; Pinheiros,
MNRJ 14014–15; Viana, MZUSP 15052–53; Sooretama,
47
MZUSP 2463; Guarapari, MZUSP 15050–51; Domingos
Martins, MNRJ 1925; Linhares, MNRJ 1926; Baixo Guandu,
IBSP 8876 (holotype); Rio de Janeiro: Cambuci, MNRJ
14487; Niterói, MNRJ 13124, 14245–46, 15422; Rio de
Janeiro-Minas Gerais: Pirapetinga–Bom Jesus de Itabapoana,
MNRJ 15825 (Passos et al., 2005, 2006; Costa et al., 2009).
New data.—Bahia: Fazenda Unacu, São José, MZUSP 9098;
Minas Gerais: Resplendor, IBSP 42606; Parque Nacional das
Sempre Vivas, MCN-R 2561; Espı́rito Santo: Aracruz, MNRJ
4856.
ACKNOWLEDGMENTS
We are thankful to V. Wallach (MCZ) and J.Cassimiro (IB/
USP) for helpful comments on the manuscript; V. Wallach
(MCZ) for english revision on the manuscript; D. Frost
(AMNH), C. McCarthy (BMNH), H. Voris and A. Resetar
(FMNH), J. Rosado (MCZ), K. de Queiroz and R. Wilson
(USNM), C. Phillips and J. Petzing (UIMNH), J. Faivovich
(MACN), A.Prudente (MPEG), P. Garcia (UFMG), F. Franco
(IBSP), H. Zaher and C. Mello (MZUSP), G. Colli and M. Zatz
(CHUNB), I. Sazima and P. Manzani (ZUEC), G. Cotta
(FUNED), M. Pires (LZV), F. Junca (MZUEFS), P. Garcia
(UFMG), R. Feio (MZUFV), L. Nascimento (MCN-R) for
allowing and facilitating the examination of specimens. We
thank R. Mark-Oliver and F. Tillack (ZMB) for sending
photos of the holotype of T. guayaquilensis and paralectotype of T. macrolepis (ZMB 5722). We thank M. Rodrigues for
sending field information about T. jani; and H. Pinto, M.
Rodrigues, M. Teixeira, Jr., R. Recoder, M. de Sena, and J.
Cassimiro for specimens recently collected. We also are
thankful to J. Cassimiro and M. Teixeira, Jr. for live
specimen and habitat photos of T. jani; P. Nascimento
(MNRJ) for the line art. We are grateful to Fundação de
Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) and
Conselho Nacional de Ciência e Tecnologia (CNPq) for
financial support.
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