A New Species of Amerotyphlops from Northeastern Brazil, with Comments on Distribution of Related Species
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A Taxonomic Framework for Typhlopid Snakes from the Caribbean and Other Regions (Reptilia, Squamata)
caribbean herpetology article A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata) S. Blair Hedges1,*, Angela B. Marion1, Kelly M. Lipp1,2, Julie Marin3,4, and Nicolas Vidal3 1Department of Biology, Pennsylvania State University, University Park, PA 16802-5301, USA. 2Current address: School of Dentistry, University of North Carolina, Chapel Hill, NC 27599-7450, USA. 3Département Systématique et Evolution, UMR 7138, C.P. 26, Muséum National d’Histoire Naturelle, 57 rue Cuvier, F-75231 Paris cedex 05, France. 4Current address: Department of Biology, Pennsylvania State University, University Park, PA 16802-5301 USA. *Corresponding author ([email protected]) Article registration: http://zoobank.org/urn:lsid:zoobank.org:pub:47191405-862B-4FB6-8A28-29AB7E25FBDD Edited by: Robert W. Henderson. Date of publication: 17 January 2014. Citation: Hedges SB, Marion AB, Lipp KM, Marin J, Vidal N. 2014. A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology 49:1–61. Abstract The evolutionary history and taxonomy of worm-like snakes (scolecophidians) continues to be refined as new molec- ular data are gathered and analyzed. Here we present additional evidence on the phylogeny of these snakes, from morphological data and 489 new DNA sequences, and propose a new taxonomic framework for the family Typhlopi- dae. Of 257 named species of typhlopid snakes, 92 are now placed in molecular phylogenies along with 60 addition- al species yet to be described. Afrotyphlopinae subfam. nov. is distributed almost exclusively in sub-Saharan Africa and contains three genera: Afrotyphlops, Letheobia, and Rhinotyphlops. Asiatyphlopinae subfam. nov. is distributed in Asia, Australasia, and islands of the western and southern Pacific, and includes ten genera:Acutotyphlops, Anilios, Asiatyphlops gen. -
A Case of Albinism in Amerotyphlops Brongersmianus (Vanzolini, 1976) (Serpentes: Typhlopidae) from Southern Bahia, Northeaestern Brazil
Herpetology Notes, volume 10: 131-132 (2017) (published online on 10 March 2017) A case of albinism in Amerotyphlops brongersmianus (Vanzolini, 1976) (Serpentes: Typhlopidae) from southern Bahia, northeaestern Brazil Caio Vinícius de Mira-Mendes1,*, Celso Henrique Varela Rios2, Renato Augusto Martins2, Tadeu Medeiros1, Mirco Solé1 and Antônio Jorge Suzart Argôlo1 Albinism is defined as a congenital decrease or Bahia (16.968567°S; 39.567365°W; datum WGS84). absence of melanin in the skin, mucosa, and eyes, The specimen was photographed, collected and caused by a homozygosis in a recessive mendelian gene deposited in the herpetological collection of Museu (Bechtel, 1995; Griffiths et al., 1998). While albinism de Zoologia da Universidade de São Paulo (MZUSP), is widely spread in several pet snake lineages (Saenko state of São Paulo, Brazil, under the catalogue number et al., 2015), reports of albino snakes from the wild are MZUSP 22790. The snake had a snout-vent length of scattered in the literature (e.g., Sazima and Di-Bernardo, 229 mm and tail length of 4 mm. 1991; Duarte et al., 2005; Krecsák, 2008; Silva et al., Cases of coloration disorder are very rare in 2010, Abegg et al., 2015). Although the exact incidence scolecophidian snakes. Currently, only four reports are of albinism in snakes is not known, estimates suggest known in the literature: Epictia munoai, Amerotyphlops orders of 1:10,000 to 1:30,000 in the general population reticulatus and Grypotyphlops acutus (Nicéforo-Maria, (Bechtel, 1995). Herein, we report a case of total 1958; Orejas-Miranda, 1972; Nivalkar et al., 2012, albinism in Amerotyphlops brongersmianus (Vanzolini, respectively) that represent albinism cases, and one case 1972), from northeastern Brazil. -
Amerotyphlops Tycherus Townsend, Wilson, Ketzler, and Luque-Montes, 2008
Amerotyphlops tycherus Townsend, Wilson, Ketzler, and Luque-Montes, 2008. This recently described blindsnake is endemic to Honduras and known from two specimens from the vicinity of the type locality (the eastern slope of Cerro Santa Bárbara within Parque Nacional Montaña de Santa Bárbara), in an area supporting remnant cloud forest, and a third from La Florida, a locality within Parque Nacional Sierra de Agalta (McCranie and Valdés-Orellana, 2012) in remnant highland rainforest. This species is one of three members of the family Typhlopidae known from Honduras; the others are A. costaricensis and the also endemic A. stadelmani. Hedges et al. (2014) recently erected the genus Amerotyphlops for a group of blindsnakes largely distributed on the mainland of the Americas, formerly allocated to Typhlops. ' © Josiah H. Townsend. 122 www.mesoamericanherpetology.com www.eaglemountainpublishing.com An updated list of the amphibians and reptiles of Honduras, with comments on their nomenclature JOSÉ M. SOLÍS¹, LARRY DAVID WILSON2, AND JOSIAH H. TOWNSEND3 1Facultad de Ciencias, Escuela de Biología, Universidad Nacional Autónoma de Honduras, Depto. de Francisco Morazán, Tegucigalpa, Honduras. E-mail: [email protected] (Corresponding Author). 2Centro Zamorano de Biodiversidad, Escuela Agrícola Panamericana Zamorano, Departamento de Francisco Morazán, Honduras. E-mail: [email protected] 3Department of Biology, Indiana University of Pennsylvania, Indiana, Pennsylvania 15705-1081, United States. E-mail: josiah. [email protected] ABSTRACT: We provide an updated list of the amphibians and reptiles of Honduras, and include notes on nomenclatural changes. We recognize a total of 396 species (135 amphibians, 261 reptiles), of which 107 are endemic and seven are exotic. Key Words: Checklist, herpetofauna. -
First Record of Egg Development Time And
REVISTA DE CIÊNCIAS AMBIENTAIS RCA (ISSN 1981-8858) http://revistas.unilasalle.edu.br/index.php/Rbca Canoas, v. 14, n. 3, 2020 http://dx.doi.org/10.18316/rca.v14i.6598 FIRST RECORD OF EGG DEVELOPMENT TIME AND HATCHLING MORPHOLOGY OF Amerotyphlops paucisquamus (DIXON & HENDRICKS 1979) (SCOLECOPHIDIA: TYPHLOPIDAE), NORTHEASTERN BRAZIL Jéssica Monique da Silva Amaral1 Paula Araújo1 Vanessa do Nascimento Barbosa1 Frederico Gustavo Rodrigues França1 ABSTRACT We present the rst description of the morphology and mass of hatchlings of the blindsnake Amerotyphlops paucisquamus, as well as information on litter size and egg development time. e clutch size was three eggs and the time of incubation was 37 days. is record seeks to ll gaps in knowledge, since data on the reproduction of this species are not yet known. ese data can be used comparatively for other scolecophidians helping to clarify systematic aspects and morphological variations in geographic scale. Keywords: Off spring; Reproduction; Snake; Blindsnake. RESUMO Primeiro registro do tempo de desenvolvimento dos ovos e morfologia de lhotes de Amerotyphlops paucisquamus (Dixon & Hendricks 1979) (Scolecophidia: Typhlopidae), nordeste do Brasil. Apresentamos a primeira descrição da morfometria e massa de lhotes de Amerotyphlops paucisquamus, além de informações sobre tamanho da ninhada e tempo de desenvolvimento dos ovos, sendo o tamanho da ninhada de três ovos e o tempo de incubação de 37 dias. Esse registro busca preencher lacunas quanto a reprodução desta espécie, visto que dados sobre o assunto são insipientes. Esperamos que esses dados possam ser utilizados de forma comparativa para outras scoleco dias, ajudando a esclarecer aspectos sistemáticos e variações morfológicas em escala geográ ca. -
Molecular Phylogeny of the Tribe Philodryadini Cope, 1886 (Dipsadidae: Xenodontinae): Rediscovering the Diversity of the South American Racers
ARTICLE Molecular phylogeny of the tribe Philodryadini Cope, 1886 (Dipsadidae: Xenodontinae): Rediscovering the diversity of the South American Racers Juan Camilo Arredondo¹⁶; Felipe G. Grazziotin²; Gustavo J. Scrocchi³; Miguel Trefaut Rodrigues⁴; Sandro Luís Bonatto⁵ & Hussam Zaher⁶⁷ ¹ Universidad CES, Facultad de Ciencias y Biotecnología, Colecciones Biológicas Universidad CES (CBUCES). Medellín, Antioquia, Colombia. ORCID: http://orcid.org/0000-0003-1925-4556. E-mail: [email protected] ² Instituto Butantan, Laboratório Especial de Coleções Zoológicas (LECZ). São Paulo, SP, Brasil. ORCID: http://orcid.org/0000-0001-9896-9722. E-mail: [email protected] ³ Fundación Miguel Lillo, Unidad Ejecutora Lillo (CONICET-UEL). San Miguel de Tucumán, Tucumán, Argentina. ORCID: http://orcid.org/0000-0002-8924-8808. E-mail: [email protected] ⁴ Universidade de São Paulo (USP), Instituto de Biociências (IB-USP), Departamento de Zoologia. São Paulo, SP, Brasil. ORCID: http://orcid.org/0000-0003-3958-9919. E-mail: [email protected] ⁵ Pontifícia Universidade Católica do Rio Grande do Sul (PUC-RS), Escola de Ciências da Saúde e da Vida, Laboratório de Biologia Genômica e Molecular. ORCID: http://orcid.org/0000-0002-0064-467X. E-mail: [email protected] ⁶ Universidade de São Paulo (USP), Museu de Zoologia (MZUSP). São Paulo, SP, Brasil. ⁷ ORCID: http://orcid.org/0000-0002-6994-489X. E-mail: [email protected] (corresponding author) Abstract. South American racers of the tribe Philodryadini are a widespread and diverse group of Neotropical snakes with a complex taxonomic and systematic history. Recent studies failed to present a robust phylogenetic hypothesis for the tribe, mainly due to incomplete taxon sampling. Here we provide the most extensive molecular phylogenetic analysis of Philodryadini available so far, including 20 species (83% of the known diversity) from which six were not sampled previously. -
BULLETIN Chicago Herpetological Society
BULLETIN of the Chicago Herpetological Society Volume 54, Number 1 January 2019 BULLETIN OF THE CHICAGO HERPETOLOGICAL SOCIETY Volume 54, Number 1 January 2019 A Note regarding the Identification of a Snakeskin Belt in the American Civil War Museum, Richmond, Virginia . Brian S. Gray 1 Dietary Notes on the Variable Coral Snake, Micrurus diastema (Duméril, Bibron & Duméril, 1854) . Taylor R. West, Tristan D. Schramer, Yatin Kalki and Daniel B. Wylie 4 Notes on Reproduction of Hurter’s Spadefoot Toads, Scaphiopus hurterii (Anura: Scaphiopodidae), from Oklahoma. Stephen R. Goldberg 9 Book Review: Peterson Field Guide to Western Reptiles and Amphibians, Fourth Edition by Robert C. Stebbins and Samuel M. McGinnis . John G. Palis 12 Lions and Tigers and Bobcats --- Oh My! . Roger A. Repp 14 Minutes of the CHS Board Meeting, December 14, 2018 . 21 In Memoriam . Jessica Wadleigh 22 Chicago Herpetological Society Income Statement: January 1 --- December 31, 2018, and Balance Sheet, December 31, 2018 . 23 Advertisements . 24 New CHS Members This Month . 24 Cover: Mountain pitviper, Ovophis monticola. Drawing (as Trimeresurus monticola) by Hurrish Chunder Khan from The Poisonous Snakes of India by Joseph Ewart, 1878 STAFF Membership in the CHS includes a subscription to the monthly Bulletin. Annual dues are: Individual Membership, $25.00; Editor: Michael A. Dloogatch --- [email protected] Family Membership, $28.00; Sustaining Membership, $50.00; Copy editor: Joan Moore Contributing Membership, $100.00; Institutional Membership, $38.00. Remittance must be made in U.S. funds. Subscribers 2019 CHS Board of Directors outside the U.S. must add $12.00 for postage. Send membership dues or address changes to: Chicago Herpetological Society, President: Rich Crowley Membership Secretary, 2430 N. -
Cerrophidion Wilsoni Jadin, Townsend, Castoe, and Campbell, 2012. The
Cerrophidion wilsoni Jadin, Townsend, Castoe, and Campbell, 2012. The Honduran Montane Pitviper is a priority one species with an EVS of 15, placing it in the high vulnerability category (see this paper). This pitviper is distributed primarily in lower montane rainforest at elevations from 1,400 to 3,491 m, but can occur peripherally in premontane rainforest and pine-oak forest as low as 1,220 m (Jadin et al. 2012). As indicated by Jadin et al. (2012: 10), this snake “occurs in at least 13 isolated highland forest areas across Eastern Nuclear Central America…and all known populations…are found within the borders of Honduras and El Salvador.” This juvenile individual was found in Refugio de Vida Silvestre Texíguat, in north-central Honduras. One of the describers of this taxon is the dedicatee of this paper, and the snake was named in honor of one of the authors. Photo by Josiah H. Townsend. Amphib. Reptile Conserv. 1 January 2019 | Volume 13 | Number 1 | e168 DEDICATION We are happy to dedicate this paper to our friend and Josiah H. Townsend. 2018. An integrative assessment colleague, Josiah H. Townsend, Associate Professor of of the taxonomic status of putative hybrid leopard frogs Biology at Indiana University of Pennsylvania, in Indiana, (Anura: Ranidae) from the Chortís Highlands of Central Pennsylvania. Over the last two decades, since he was America, with description of a new species. Systematics a student in one of Larry Wilson’s classes, Joe has built and Biodiversity 2018: 1–17. This paper is an example an imposing reputation as the principal authority on the of the seminal work being conducted by Joe Townsend herpetofauna of the biogeographically significant Chortís and his colleagues, which is exposing the underestimated Highlands of northern Central America. -
Escola De Ciências Programa De Pós-Graduação Em Zoologia Doutorado Em Zoologia
ESCOLA DE CIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA DOUTORADO EM ZOOLOGIA FIDÉLIS JÚNIO MARRA SANTOS A REVISION OF THE SMALL SNAKES OF THE FAMILY ANOMALEPIDIDAE (REPTILIA: SQUAMATA: SERPENTES), USING HIGH RESOLUTION COMPUTERIZED TOMOGRAPHY Porto Alegre 2018 PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL ESCOLA DE CIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA A REVISION OF THE SMALL SNAKES OF THE FAMILY ANOMALEPIDIDAE (REPTILIA: SQUAMATA: SERPENTES), USING HIGH RESOLUTION COMPUTERIZED TOMOGRAPHY Fidélis Júnio Marra Santos TESE DE DOUTORADO PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL Brasil 2018 PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL ESCOLA DE CIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA A REVISION OF THE SMALL SNAKES OF THE FAMILY ANOMALEPIDIDAE (REPTILIA: SQUAMATA: SERPENTES), USING HIGH RESOLUTION COMPUTERIZED TOMOGRAPHY Fidélis Júnio Marra Santos Orientador: Roberto E. Reis TESE DE DOUTORADO PORTO ALEGRE - RS - BRASIL “A onisciência é negada ao homem”. Ludwig von Mises AVISO Esta tese é parte dos requisitos necessários para obtenção do título de doutor, área de Zoologia, e como tal, não deve ser vista como uma publicação no senso do Código Internacional de Nomenclatura Zoológica (capítulo 3 – artigo 9) (apesar de disponível publicamente sem restrições). Desta forma, quaisquer informações inéditas, opiniões e hipóteses, assim como nomes novos, não estão disponíveis na literatura zoológica. Pessoas interessadas devem estar cientes de que referências públicas ao conteúdo desse estudo, na sua presente forma, somente devem ser feitas com aprovação prévia do autor. NOTICE This thesis is a partial requirement for the PhD degree in Zoology and, as such, should not be considered as a publication in the sense of the International Code of Zoological Nomenclature (chapter 3 – article 9) (although it is available without restrictions). -
Ecological Diversity of a Snake Assemblage from the Atlantic Forest at the South Coast of Paraíba, Northeast Brazil
A peer-reviewed open-access journal ZooKeys 787: 107–125Ecological (2018) diversity of a snake assemblage from the Atlantic Forest... 107 doi: 10.3897/zookeys.787.26946 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research Ecological diversity of a snake assemblage from the Atlantic Forest at the south coast of Paraíba, northeast Brazil Ivan L. R. Sampaio1, Claudileide P. Santos2, Rafaela C. França3,4, Isabella M. M. C. Pedrosa5, Mirco Solé4,6, Frederico G. R. França7 1 Programa de Pós-graduação em Ecologia e Monitoramento Ambiental, Universidade Federal da Paraíba – UFPB, Av. Santa Elizabete, s/n – Centro, CEP 58297-000 Rio Tinto, PB, Brazil 2 Programa de Pós-graduação em Ciências Biológicas – Zoologia, Universidade Federal da Paraíba, Castelo Branco, CEP 58051-900 João Pes- soa, PB, Brazil 3 Programa de Pós-graduação em Ecologia e Conservação da Biodiversidade, Universidade Esta- dual de Santa Cruz, Rodovia Jorge Amado, Km 16, CEP 45662-900 Ilhéus, Bahia, Brazil 4 Herpetology Sec- tion, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany 5 Núcleo de Ecologia de Serpentes UFPB, Laboratório de Ecologia Animal, Universidade Federal da Paraíba – UFPB, CCAE, Av. Santa Elizabete, s/n – Centro, CEP 58297-000 Rio Tinto, PB, Brazil 6 Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km. 16, 45662-900 Ilhéus, Bahia, Brazil 7 Departamento de Engenharia e Meio Ambiente, Centro de Ciências Aplicadas e Educação, Universi- dade Federal da Paraíba – UFPB, Av. Santa Elizabete, s/n – Centro, CEP 58297-000 Rio Tinto, PB, Brazil Corresponding author: Frederico G. R. França ([email protected]) Academic editor: R. -
The Reptiles of Paraguay: Literature, Distribution, and an Annotated Taxonomic Checklist Pier Cacciali1, Norman J
Reptiles of Paraguay SPECIAL PUBLICATION OF THE MUSEUM OF SOUTHWESTERN BIOLOGY NUMBER 11, pp. 1–373 25 June 2016 The Reptiles of Paraguay: Literature, Distribution, and an Annotated Taxonomic Checklist Pier Cacciali1, Norman J. Scott2, Aida Luz Aquino Ortíz3, Lee A. Fitzgerald4, and Paul Smith5 1 Instituto de Investigación Biológica del Paraguay. Del Escudo 1607, Asunción, Paraguay; and Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany. E-mail: [email protected] 2 Corresponding Author: Research Associate, Department of Biology, University of New Mexico; Research Associate, Smithsonian Institution; Research Associate, Los Angeles County Museum of Natural History, P. O. Box 307, Creston, California 93432, USA. E-mail: [email protected] 3 Director, World Wildlife Fund Country Office, 150 c/ Peron, Edificio Opa Rudy 150, 4to piso, Asunción, Paraguay. E-mail: [email protected] 4 Biodiversity Research and Teaching Collections, Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, Texas 77843, USA. E-mail: [email protected] 5 Fauna Paraguay, Encarnación, Departamento Itapúa, Paraguay, www.faunaparaguay.com; E-mail: [email protected]; and: Para La Tierra, Municipalidad de Santa Barbara, Departamento San Pedro, Paraguay, www.paralatierra.org; E-mail: [email protected]. Special Publication of the Museum of Southwestern Biology 1 Reptiles of Paraguay TABLE OF CONTENTS DEDICATION ........................................................................................................................... -
REVIEW Confronting Taxonomic Vandalism in Biology: Conscientious
applyparastyle “fig//caption/p[1]” parastyle “FigCapt” Biological Journal of the Linnean Society, 2021, 133, 645–670. With 3 figures. REVIEW Confronting taxonomic vandalism in biology: conscientious community self-organization can preserve nomenclatural stability Downloaded from https://academic.oup.com/biolinnean/article/133/3/645/6240088 by guest on 30 June 2021 WOLFGANG WÜSTER1,*, , SCOTT A. THOMSON2, MARK O’SHEA3 and HINRICH KAISER4 1Molecular Ecology and Fisheries Genetics Laboratory, School of Natural Sciences, Bangor University, Bangor LL57 2UW, UK 2Museu de Zoologia da Universidade de São Paulo, Divisão de Vertebrados (Herpetologia), Avenida Nazaré, 481, Ipiranga, 04263-000, São Paulo, SP, Brazil; and Chelonian Research Institute, 401 South Central Avenue, Oviedo, FL 32765, USA 3Faculty of Science and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton WV1 1LY, UK 4Department of Vertebrate Zoology, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; and Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, CA 92395, USA Received 28 October 2020; revised 17 January 2021; accepted for publication 19 January 2021 Self-published taxon descriptions, bereft of a basis of evidence, are a long-standing problem in taxonomy. The problem derives in part from the Principle of Priority in the International Code of Zoological Nomenclature, which forces the use of the oldest available nomen irrespective of scientific merit. This provides a route to ‘immortality’ for unscrupulous individuals through the mass-naming of taxa without scientific basis, a phenomenon referred to as taxonomic vandalism. Following a flood of unscientific taxon namings, in 2013 a group of concerned herpetologists organized a widely supported, community-based campaign to treat these nomina as lying outside the permanent scientific record, and to ignore and overwrite them as appropriate. -
Table S3.1. Habitat Use of Sampled Snakes. Taxonomic Nomenclature
Table S3.1. Habitat use of sampled snakes. Taxonomic nomenclature follows the current classification indexed in the Reptile Database ( http://www.reptile-database.org/ ). For some species, references may reflect outdated taxonomic status. Individual species are coded for habitat association according to Table 3.1. References for this table are listed below. Habitat use for species without a reference were inferred from sister taxa. Broad Habitat Specific Habit Species Association Association References Acanthophis antarcticus Semifossorial Terrestrial-Fossorial Cogger, 2014 Acanthophis laevis Semifossorial Terrestrial-Fossorial O'Shea, 1996 Acanthophis praelongus Semifossorial Terrestrial-Fossorial Cogger, 2014 Acanthophis pyrrhus Semifossorial Terrestrial-Fossorial Cogger, 2014 Acanthophis rugosus Semifossorial Terrestrial-Fossorial Cogger, 2014 Acanthophis wellsi Semifossorial Terrestrial-Fossorial Cogger, 2014 Achalinus meiguensis Semifossorial Subterranean-Debris Wang et al., 2009 Achalinus rufescens Semifossorial Subterranean-Debris Das, 2010 Acrantophis dumerili Terrestrial Terrestrial Andreone & Luiselli, 2000 Acrantophis madagascariensis Terrestrial Terrestrial Andreone & Luiselli, 2000 Acrochordus arafurae Aquatic-Mixed Intertidal Murphy, 2012 Acrochordus granulatus Aquatic-Mixed Intertidal Lang & Vogel, 2005 Acrochordus javanicus Aquatic-Mixed Intertidal Lang & Vogel, 2005 Acutotyphlops kunuaensis Fossorial Subterranean-Burrower Hedges et al., 2014 Acutotyphlops subocularis Fossorial Subterranean-Burrower Hedges et al., 2014