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Zootaxa, Taxonomic revision of the Brazilian
Zootaxa 2364: 1–63 (2010)
www.mapress.com / zootaxa/
Copyright © 2010 · Magnolia Press
ISSN 1175-5326 (print edition)
Monograph
ZOOTAXA
ISSN 1175-5334 (online edition)
ZOOTAXA
2364
Taxonomic revision of the Brazilian Atlantic Forest Atractus
(Reptilia: Serpentes: Dipsadidae)
PAULO PASSOS1, 3, RONALDO FERNANDES1, RENATO S. BÉRNILS1
& JULIO C. DE MOURA-LEITE2
1
Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Quinta da Boa Vista,
20940-040, Rio de Janeiro, RJ, Brasil
2
Laboratório de Herpetologia, Museu de História Natural Capão da Imbuia, Prof. Benedito Conceição 407, 82810080 & Núcleo de Estudos do Comportamento, Pontifícia Universidade Católica do Paraná, Imaculada Conceição
1155, Prado Velho, 80215-901, Curitiba, PR, Brasil
3
Corresponding author, E-mail: [email protected]
Magnolia Press
Auckland, New Zealand
Accepted by D. Gower: 21 Jan. 2010; published: 19 Feb. 2010
PAULO PASSOS, RONALDO FERNANDES, RENATO S. BÉRNILS & JULIO C. DE MOURA-LEITE
Taxonomic revision of the Brazilian Atlantic Forest Atractus (Reptilia: Serpentes: Dipsadidae)
(Zootaxa 2364)
63 pp.; 30 cm.
19 February 2010
ISBN 978-1-86977-481-3 (paperback)
ISBN 978-1-86977-482-0 (Online edition)
FIRST PUBLISHED IN 2010 BY
Magnolia Press
P.O. Box 41-383
Auckland 1346
New Zealand
e-mail: [email protected]
http://www.mapress.com/zootaxa/
© 2010 Magnolia Press
All rights reserved.
No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any
means, without prior written permission from the publisher, to whom all requests to reproduce copyright
material should be directed in writing.
This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose
other than private research use.
ISSN 1175-5326
(Print edition)
ISSN 1175-5334
(Online edition)
2 · Zootaxa 2364 © 2010 Magnolia Press
PASSOS ET AL.
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Resumo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Material and Techniques. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Statistical Analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Scope and Style of the Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Species Accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Atractus caete sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Atractus francoi sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Atractus guentheri (Wucherer, 1861) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Atractus maculatus (Günther, 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Atractus pantostictus Fernandes & Puorto, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Atractus paraguayensis Werner, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Atractus potschi Fernandes, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Atractus reticulatus (Boulenger, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Atractus ronnie Passos, Fernandes & Borges-Nojosa, 2007. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Atractus serranus Amaral, 1930 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Atractus thalesdelemai Passos, Fernandes & Zanella, 2005. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Atractus trihedrurus Amaral, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Atractus zebrinus (Jan, 1862) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Key to the Atlantic Forest species of Atractus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Abstract
Dipsadine snakes of the genus Atractus are endemic to the Neotropical region, occurring from Panama to Argentina.
Currently, the taxonomic status of most species of the genus is unclear and previous attempts of taxonomic revisions
have been local in scale. In this paper we evaluate the taxonomic status of the Brazilian Atlantic Forest species of Atractus based on meristic, morphometric, maxillary dentition, and hemipenis characters. Quantitative and qualitative analyses suggest the recognition of one new species (A. caete sp. nov.) from the state of Alagoas, another (A. francoi sp. nov.)
from the mountainous regions of the states of Rio de Janeiro and São Paulo, and the synonymy of A. kangueryensis with
A. thalesdelemai. Specimens previously assigned to A. taeniatus in Argentina and Brazil are here considered A. paraguayensis. A key to the Atlantic Forest Atractus is provided and three new species groups are proposed for some cisAndean Atractus, mainly on the basis of hemipenial morphology: the A. emmeli, A. maculatus, and A. pantostictus species groups.
Key words: Atractus, Atlantic Forest, South America, Morphology, Geographical variation, Taxonomy
Resumo
As serpentes dipsadíneas do gênero Atractus são endêmicas da região Neotropical, ocorrendo do Panamá à Argentina.
Atualmente, o posicionamento taxonômico da maioria das espécies do gênero é confuso e as tentativas de revisões taxonômicas foram limitadas à escala regional. Neste trabalho, nós avaliamos o posicionamento taxonômico das espécies de
Atractus da Floresta Atlântica brasileira por meio de caracteres merísticos, morfométricos, de dentição maxilar e morfologia do hemipênis. Análises quantitativas e qualitativas sugerem o reconhecimento de uma nova espécie (A. caete sp.
nov.) do estado de Alagoas e outra (A. francoi sp. nov.) da região serrana dos estados do Rio de Janeiro e São Paulo e a
REVISION OF BRAZILIAN ATLANTIC FOREST ATRACTUS
Zootaxa 2364 © 2010 Magnolia Press ·
3
sinonimização de A. kangueryensis com A. thalesdelemai. Ademais, os espécimes previamente assinalados a A. taeniatus
na Argentina e Brasil são aqui considerados A. paraguayensis. Uma chave é fornecida para as Atractus da Floresta Atlântica e três novos grupos de espécies são propostos para algumas Atractus cisandinas, baseado principalmente na morfologia do hemipênis: os grupo de espécies de A. emmeli, A. maculatus e A. pantostictus.
Palavras-chave: Atractus, Floresta Atlântica, América do Sul, Morfologia, Variação geográfica, Taxonomia
Introduction
The dipsadine genus Atractus is endemic of the Neotropical region, occurring primarily on the mainland from
11º North to 35º South, from sea level to 4,500 meters elevation, in almost all South American biomes (Passos
2008). The genus comprises small to moderate-sized snakes, having secretive (semi-fossorial or cryptozoic)
lifestyles and feeding on earthworms, arthropods, and mollusks (Cunha & Nascimento 1993; Martins &
Oliveira 1993, 1999; Cisneros-Heredia 2005). Atractus is a diverse genus of snakes closely related to Adelphicos and Geophis (Savage 1960; Downs 1967; Fernandes 1995b; Passos 2008), including about 130 valid species, many of them known only from their type specimens and sometimes having restricted distributions
(Passos & Fernandes 2008; Prudente & Passos 2008; Passos & Arredondo 2009; Passos et al. 2009a,b,c,d,e,f).
To date, the taxonomic status of several species remains unclear, and all attempts of taxonomic revisions have
been limited to a local scale (Savage 1960; Roze 1961; Cunha & Nascimento 1993; Martins & Oliveira 1993;
Giraudo & Scrocchi 2000; Myers 2003; Silva 2004; Esqueda & La Marca 2005). The major problem with
Atractus taxonomy is that geographical variability data as well as sexual and/or ontogenetic variations are
unknown for most currently recognized taxa (Passos et al. 2009 a,c,e). Lack of comparative specimens and
poor knowledge of intraspecific variation led some authors to recognize new taxa on the basis of slight
nuances in colour pattern and/or meristic characters (e.g., Scrocchi & Cei 1991; Alvarez et al. 1992; BernalCarol & Roze 1996). These taxa, in many cases, were shown to not be diagnosable from previous recognized
species after the examination of large series of specimens (e.g., Passos et al. 2009d; Fernandes 1996; Passos et
al. 2009c).
The Brazilian Atlantic Forest is one of the most diverse and threatened ecosystems on Earth, with only
about 7% of its orginal coverage remaining (Morellato & Haddad; Oliveira-Filho & Fontes 2000). Its remarkable diversity has been attributed to the presence of a rich flora with unique physiological adaptations to
unusual biophysical characteristics (Pessenda et al. 2009). The biome is composed of two major vegetation
types, the coastal forest or Atlantic Rainforest (sensu strictu) and the tropical seasonal forest or Atlantic Semidecidous forest (Morellato & Haddad 2000). The Atlantic Rainforest covers mostly low to medium elevations
(≤ 1000 m) of the eastern slopes of the mountain chain that runs subparallel and close to the coastline from
southern to northeastern Brazil; the Atlantic Semi-deciduous forest extends across the plateau (usually > 600
m) in the centre and southeastern interior of the country (Morellato & Haddad 2000). The Atlantic Rainforest
experiences warm and wet climate without a well defined dry season, while a seasonal climate with relative
severe dry season (generally from April to September) predominates over the distribution of the Atlantic
Semi-deciduous forest (Morellato & Haddad 2000). The biome lies almost entirely within Brazil, with a small
portion extending to northeastern Argentina and southeastern Paraguay (Oliveira-Filho & Fontes 2000). Currently, 11 species of Atlantic Forest Atractus are recognized (A. guentheri, A. kangueryensis, A. maculatus, A.
paraguayensis, A. potschi, A. ronnie, A. serranus, “A. taeniatus”, A. thalesdelemai, A. trihedrurus, and A.
zebrinus), and two other are distributed on transitional zones between Atlantic Forest and general open formations in central and southern Brazil (A. pantostictus and A. reticulatus) (Fernandes 1996; Passos et al. 2005b;
Cacciali et al. 2007; Passos et al. 2007b).
The aim of this study is to provide a comprehensive revision of the Atlantic Forest Atractus in order to
evaluate the validity of the currently know taxa, and to present accurate diagnostic characters and data on
intraspecific and interspecific variation for all recognized species in the area.
4 · Zootaxa 2364 © 2010 Magnolia Press
PASSOS ET AL.
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