DOCSLIB.ORG

MECHANISMS of ADAPTATION in CORAL SNAKE MIMICRY David

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
MECHANISMS of ADAPTATION in CORAL SNAKE MIMICRY David MECHANISMS OF ADAPTATION IN CORAL SNAKE MIMICRY David William Kikuchi A dissertation submitted to the faculty of the University of North Carolina at Chapel Hill in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biology. Chapel Hill 2013 Approved by: David W. Pfennig Allen H. Hurlbert Karin S. Pfennig Maria R. Servedio Kyle Summers ©2013 David William Kikuchi ALL RIGHTS RESERVED ii ABSTRACT DAVID WILLIAM KIKUCHI: MECHANISMS OF ADAPTATION IN CORAL SNAKE MIMICRY (Under the direction of David Pfennig) In Batesian mimicry, an undefended prey species (the mimic) evolves to resemble a defended one (the model) because of the selective advantage of this resemblance in deterring predation. Although Batesian mimicry is one of the oldest known examples of natural selection’s power to produce adaptation, many unanswered questions remain about its evolution, including how mimetic signals coevolve with the perceptual abilities of predators, how mimetic signals are produced, how important shared evolutionary history with a model species is for mimics, and if mimicry can evolve over rough adaptive landscapes. My thesis attempts to address these knowledge gaps by examining the venomous coral snake Micrurus fulvius and its nonvenomous mimic, the scarlet kingsnake Lampropeltis elapsoides. In addition to my empirical studies, I have produced two reviews: one is a general review of mimicry in the form of an annotated bibliography, and the other a review of the hypotheses for imperfect mimicry. In a field experiment, I asked whether or not predators were sensitive to differences between models and mimics in phenotype, that is to say, imperfect mimicry. iii My results revealed that imperfect mimicry was tolerated in some dimensions but not others, and that predators’ cognitive biases play a role in perpetuating imperfect mimicry. Two analytical studies of snake pigmentation revealed that coral snakes, their mimics, and several nonmimetic snakes use the same structures and pigments to produce their coloration. The spectral properties of colors produced by those pigments produce similar perceptual experiences for likely agents of selection in coral snake mimicry. This suggests that sharing developmental systems may facilitate the evolution of mimicry. In another field experiment I tested the assumption that the adaptive landscape between mimicry and crypsis (from which mimicry is thought to evolve) is always rough, featuring an “adaptive valley” of selection against intermediate phenotypes. Under ecological conditions that produce strong selection for precise mimicry, intermediate phenotypes were selected against; however, this was not the case when selection for mimicry was less intense. Therefore, the assumption that the evolution of mimicry always involves a transition through maladaptive intermediate phenotypes may be unwarranted. iv ACKNOWLEDGMENTS Without the contributions of many individuals besides myself, this thesis would never have been finished. I would like to thank my advisor for his objective yet unstintingly enthusiastic involvement with my projects, and my thesis committee for their feedback on the ideas we produced. The many people who have been associated with the Pfennig labs gave me critical advice on everything from statistics to the practicalities of being a graduate student, and indispensable help with experiments. The much larger, less easily defined community of the UNC Biology Department made no less valuable contributions by creating an atmosphere of intellectual curiosity that perpetually fostered new insight. Thanks to my friends for their support, tolerance, and laughter during these five years of reconciling a bunch of abstract notions with the natural world. And most of all, thank you to my family for your unconditional love, without which none of this would mean a thing. v CHAPTER ACKNOWLEDGMENTS Chapter III: We thank K. Pfennig, R. Martin, J. Santos, C. Ledón-Rettig, S. Dhole, A. Leichty, L. Bono, J. Paull, J. Weiss, M. Edmunds, and an anonymous reviewer for helpful comments, and A. Bailey, S. Clarke, and W. Juliantius for lab assistance. This work was funded by the US National Science Foundation and Sigma Xi. All procedures complied with applicable state and US federal regulations. Chapter IV: We thank Karin Pfennig, Verónica Rodriguez-Moncalvo, Lisa Bono, and three anonymous referees for helpful comments. Antonio Serrato helped with specimen collection. Chris Willett and Erin Burch aided with spectroscopy, and Vicky Madden and Steven Ray provided TEM services. Ken Wray kindly furnished coral snake specimens. Animal research was conducted under UNC IACUC permit 11-108. Funding was provided by the National Science Foundation (DEB-1110385 and DEB-1019479). Chapter V: We thank Audrey Kelly, Steve Mullin, Karin Pfennig, Antonio Serrato, Lisa Wünch, and the instructors and students in the herpetology class at SWRS for help in the field. This work was supported by NSF grants 1110385 and 1019479 and the Royster Society of Fellows at UNC-Chapel Hill. Snake collections were licensed under Arizona Game and Fish Scientific Collecting Permit #D12206055. All procedures were carried out in compliance with the Institutional Animal Care and Use Committee at UNC-Chapel Hill, under application #11-108. vi Chapter VI: We thank K. Pfennig, R. Martin, A. Leichty, J. Weiss, and two anonymous reviewers for helpful comments. Y. Choi, E. Christenson, S. Clarke, R. Searles, and G. Zhang assisted in the lab and field. The North Carolina State Museum of Natural Sciences furnished preserved specimens used in this study. This work was funded by the US National Science Foundation and a Theodore Roosevelt Memorial Grant. All procedures complied with applicable state and US federal regulations. Chapter VII: Our work has been supported by NSF grants DEB-1110385 and DEB- 1019479 and the Royster Society of Fellows at UNC-Chapel Hill. We would like to thank two anonymous reviewers for helpful comments. vii TABLE OF CONTENTS LIST OF TABLES....................................................................................................................xi LIST OF FIGURES.................................................................................................................xii CHAPTER I. GENERAL INTRODUCTION............................................................................................1 II. MIMICRY, AN ANNOTATED BIBLIOGRAPHY................................................................5 INTRODUCTION.........................................................................................................5 GENERAL OVERVIEWS..............................................................................................6 DEFINING MIMICRY..................................................................................................7 TAXONOMIC DISTRIBUTION......................................................................................9 HISTORICAL BACKGROUND....................................................................................12 BATESIAN MIMICRY................................................................................................15 MÜLLERIAN MIMICRY............................................................................................27 FUNCTIONS OF MIMETIC SIGNALS...........................................................................30 MULTIMODAL MIMICRY.........................................................................................31 IMPERFECT MIMICRY..............................................................................................32 GENETICS OF MIMICRY...........................................................................................35 POLYMORPHIC AND SEX-LIMITED MIMICRY...........................................................38 ROLE IN SPECIATION...............................................................................................40 INFLUENCE OF RECEIVER PERCEPTION....................................................................43 viii III. PREDATOR COGNITION PERMITS IMPERFECT CORAL SNAKE MIMICRY...................47 Summary................................................................................................................47 Introduction............................................................................................................47 Methods..................................................................................................................50 Results and Discussion..........................................................................................52 IV. A BATESIAN MIMIC AND ITS MODEL SHARE COLOR PRODUCTION MECHANISMS..........................................................................................61 Summary................................................................................................................61 Introduction............................................................................................................62 Methods..................................................................................................................64 Results....................................................................................................................70 Discussion..............................................................................................................73 V. MIMICRY’S PALETTE: WIDESPREAD USE OF CONSERVED PIGMENTS IN THE APOSEMATIC SIGNALS OF SNAKES.....................................................................87
Load more

Recommended publications
  • Semen Collection and Evaluation in Micrurus Corallinus
    Herpetological Conservation and Biology 15(3):620–625. Submitted: 22 May 2020; Accepted: 12 November 2020; Published: 16 December 2020. SEMEN COLLECTION AND EVALUATION IN MICRURUS CORALLINUS RAFAELA ZANI COETI1,2,4, KALENA BARROS DA SILVA3, GIUSEPPE PUORTO3, SILVIA REGINA TAVAGLIA-CARDOSO3, AND SELMA MARIA DE ALMEIDA-SANTOS1,2 1Laboratório de Ecologia e Evolução, Instituto Butantan, 1500 Avenida Vital Brasil, São Paulo 05503–900, Brazil 2Programa de Pós Graduação em Anatomia dos Animais Domésticos e Silvestres, Universidade de São Paulo, 87 Avenida Professor Doutor Orlando Marques de Paiva, São Paulo 05508–270, Brazil 3Museu Biológico, Instituto Butantan, 1500 Avenida Vital Brasil, São Paulo 05503–900, Brazil 4Corresponding author, e-mail: [email protected] Abstract.—The Painted Coral Snake Micrurus corallinus is one of the Brazilian species kept in captivity to obtain venom for antivenom production. Difficulties in establishing a sizeable breeding colony make it necessary to find alternatives that increase the reproductive efficiency of captive individuals. Here, we tested a semen collection protocol and characterize the seminal parameters of captive M. corallinus. We collected semen during the mating season of the species (spring-summer) and were successful at every first attempt. Spermatozoa of M. corallinus are elongated and filiform, and the midpiece is the longest part. Sperm motility and progressive motility reached values of 80% and 3.6%, respectively, during the reproductive period of this species. Our results will allow further studies to improve husbandry, reproductive rates, and conservation of captive M. corallinus. Key Words.—reproduction; reproductive biotechniques; reptiles; sperm parameters INTRODUCTION and capture rates of individual M. corallinus in the wild (Roze 1996) are worrisome and also make it difficult to Reproductive biotechniques have been useful in establish a breeding colony with a substantial number implementing conservation projects for endangered of animals.
    [Show full text]
  • Y Coralillos Falsos (Serpentes: Colubridae) De Veracruz, México
    Acta Zoológica ActaMexicana Zool. (n.s.)Mex. 22(3):(n.s.) 22(3)11-22 (2006) CORALILLOS VERDADEROS (SERPENTES: ELAPIDAE) Y CORALILLOS FALSOS (SERPENTES: COLUBRIDAE) DE VERACRUZ, MÉXICO Miguel A. DE LA TORRE-LORANCA1,3,4, Gustavo AGUIRRE-LEÓN1 y Marco A. LÓPEZ-LUNA2,4 1Instituto de Ecología, A. C., Departamento de Biodiversidad y Ecología Animal, Km. 2.5 Carr. Antigua a Coatepec No. 351, Cong. El Haya, C. P. 91070 Xalapa, Veracruz, MÉXICO. [email protected] 2Universidad Juárez Autónoma de Tabasco, División Académica de Ciencias Biológicas, Km. 0.5 Carretera Villahermosa-Cárdenas entronque a Bosques de Saloya, Villahermosa, Tabasco MÉXICO. [email protected] 3Dirección actual: Instituto Tecnológico Superior de Zongolica. Km 4 Carretera a la Compañia s/n, Tepetitlanapa, CP 95005 Zongolica, Veracruz, MÉXICO [email protected] 4Centro de Investigaciones Herpetológicas de Veracruz A. C. RESUMEN En el Estado de Veracruz se distribuyen cinco especies de coralillos verdaderos del género Micrurus y 14 especies de coralillos falsos de diferentes géneros de colúbridos, lo que hace más probables los encuentros con coralillos falsos. Sin embargo, la identificación por patrones de color entre coralillos verdaderos y falsos no es confiable, a causa de la variación del color inter e intraespecífica y a las semejanzas de coloración entre varias especies de estas dos familias de serpientes. Palabras clave: Colubridae, Elapidae, Micrurus, mimetismo, patrón de coloración, Veracruz ABSTRACT Five species of coral snakes genus Micrurus, and 14 species of mimic false coral snakes of different colubrid genera ocurr in Veracruz, making encounters with false coral snakes more likely. However, positive identification by color pattern between coral snakes and their mimics is not reliable because of inter and intraspecific color variation and similarities in coloration between several species of these two snake families.
    [Show full text]
  • Birds, Reptiles, Amphibians, Vascular Plants, and Habitat in the Gila River Riparian Zone in Southwestern New Mexico
    Birds, Reptiles, Amphibians, Vascular Plants, and Habitat in the Gila River Riparian Zone in Southwestern New Mexico Kansas Biological Survey Report #151 Kelly Kindscher, Randy Jennings, William Norris, and Roland Shook September 8, 2008 Birds, Reptiles, Amphibians, Vascular Plants, and Habitat in the Gila River Riparian Zone in Southwestern New Mexico Cover Photo: The Gila River in New Mexico. Photo by Kelly Kindscher, September 2006. Kelly Kindscher, Associate Scientist, Kansas Biological Survey, University of Kansas, 2101 Constant Avenue, Lawrence, KS 66047, Email: [email protected] Randy Jennings, Professor, Department of Natural Sciences, Western New Mexico University, PO Box 680, 1000 W. College Ave., Silver City, NM 88062, Email: [email protected] William Norris, Associate Professor, Department of Natural Sciences, Western New Mexico University, PO Box 680, 1000 W. College Ave., Silver City, NM 88062, Email: [email protected] Roland Shook, Emeritus Professor, Biology, Department of Natural Sciences, Western New Mexico University, PO Box 680, 1000 W. College Ave., Silver City, NM 88062, Email: [email protected] Citation: Kindscher, K., R. Jennings, W. Norris, and R. Shook. Birds, Reptiles, Amphibians, Vascular Plants, and Habitat in the Gila River Riparian Zone in Southwestern New Mexico. Open-File Report No. 151. Kansas Biological Survey, Lawrence, KS. ii + 42 pp. Abstract During 2006 and 2007 our research crews collected data on plants, vegetation, birds, reptiles, and amphibians at 49 sites along the Gila River in southwest New Mexico from upstream of the Gila Cliff Dwellings on the Middle and West Forks of the Gila to sites below the town of Red Rock, New Mexico.
    [Show full text]
  • Pituophis Catenifer
    COSEWIC Assessment and Status Report on the Gophersnake Pituophis catenifer Pacific Northwestern Gophersnake – P.c. catenifer Great Basin Gophersnake – P.C. deserticola Bullsnake – P.C. sayi in Canada EXTIRPATED - Pacific Northwestern Gophersnake – P.c. catenifer THREATENED - Great Basin Gophersnake – P.c. deserticola DATA DEFICIENT - Bullsnake – P.c. sayi 2002 COSEWIC COSEPAC COMMITTEE ON THE STATUS OF COMITÉ SUR LA SITUATION DES ENDANGERED WILDLIFE IN ESPÈCES EN PÉRIL CANADA AU CANADA COSEWIC status reports are working documents used in assigning the status of wildlife species suspected of being at risk. This report may be cited as follows: Please note: Persons wishing to cite data in the report should refer to the report (and cite the author(s)); persons wishing to cite the COSEWIC status will refer to the assessment (and cite COSEWIC). A production note will be provided if additional information on the status report history is required. COSEWIC 2002. COSEWIC assessment and status report on the Gophersnake Pituophis catenifer in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. vii + 33 pp. Waye, H., and C. Shewchuk. 2002. COSEWIC status report on the Gophersnake Pituophis catenifer in Canada in COSEWIC assessment and status report on the Gophersnake Pituophis catenifer in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. 1-33 pp. For additional copies contact: COSEWIC Secretariat c/o Canadian Wildlife Service Environment Canada Ottawa, ON K1A 0H3 Tel.: (819) 997-4991 / (819) 953-3215 Fax: (819) 994-3684 E-mail: COSEWIC/[email protected] http://www.cosewic.gc.ca Ếgalement disponible en français sous le titre Évaluation et Rapport du COSEPAC sur la situation de la couleuvre à nez mince (Pituophis catenifer) au Canada Cover illustration: Gophersnake — Illustration by Sarah Ingwersen, Aurora, Ontario.
    [Show full text]
  • Identifying Amphibians and Reptiles in Zoos and Aquariums ZOO VIEW
    290 ZOO VIEW Herpetological Review, 2015, 46(2), 290–294. © 2015 by Society for the Study of Amphibians and Reptiles Identifying Amphibians and Reptiles in Zoos and Aquariums PLUS ҫa ChanGe, PluS C’eST la même ChoSe [The more ThinGS ChanGe, Snakes and their allies were traditionally placed in the genus THE MORE THEY STAY THE SAME] Elaphe but were recently referred to Pantherophis based on their —JEAN-BAPTISTE ALPHONSE KARR, 1849 close relationship to other lampropeltine colubrids of the New World (Burbrink and Lawson 2007). Different combinations are Reptiles and amphibians are relatively unique in the sense used by different authors and my colleagues are struggling with of constantly changing taxonomies. That phenomenon simply is these differences; in other words, which names should they use? not a big operative problem for bird and mammal zoo person- Some biologists believe that there is a rule that the most recent nel. To gain a sense of why this is happening, refer to Frost and taxonomic paper should be the one used but there is no such Hillis (1990). There is confusion caused by changes in both stan- established convention in the Code (The International Code of dard and scientific names in herpetology. The general principle Zoological Nomenclature). Recently, a convincing description of in a zoo is that one wants to be talking about the same species the dangers of taxonomic vandalism leading to potential desta- when putting live animals together for breeding or exhibit or bilization has appeared in the literature (Kaiser et al. 2013) and analyzing records for genealogy or research.
    [Show full text]
  • Mimicry and Defense
    3/24/2015 Professor Donald McFarlane Mimicry and Defense Protective Strategies Camouflage (“Cryptic coloration”) Diverse Coloration Diversion Structures Startle Structures 2 1 3/24/2015 Camouflage (“Cryptic coloration”) Minimize 3d shape, e.g. flatfish Halibut (Hippoglossus hippoglossus) 3 4 2 3/24/2015 Counter‐Shading 5 Disruptive Coloration 6 3 3/24/2015 Polymorphism – Cepeae snails 7 Polymorphism – Oophaga granuliferus 8 4 3/24/2015 Polymorphism – 9 Polymorphism – Oophaga Geographic locations of study populations and their color patterns. (A) Map of the pacific coast of Colombia showing the three study localities: in blue Oophaga histrionica, in orange O. lehmanni, and in green the pHYB population. (B) Examples of color patterns of individuals from the pHYB population (1–4) and the pattern from a hybrid between Oophaga histrionica and O. lehmanni bred in the laboratory (H) 10 5 3/24/2015 Diversion Structures 11 Startle Structures 12 6 3/24/2015 Warning Coloration (Aposematic coloration) Advertise organism as distasteful, toxic or venomous Problem: Predators must learn by attacking prey; predator learning is costly to prey. Therefore strong selective pressure to STANDARDIZE on a few colors/patterns. This is MULLERIAN MIMICRY. Most common is yellow/black, or red/yellow/black 13 Warning Coloration (Aposematic coloration) Bumblebee (Bombus Black and yellow mangrove snake (Boiga sp.) Sand Wasp (bembix oculata) dendrophila) Yellow‐banded poison dart frog (Dendrobates leucomelas Fire salamander ( Salamandra salamandra) 14 7 3/24/2015 Warning Coloration (Aposematic coloration) coral snakes (Micrurus sp.) ~ 50 species in two families, all venomous 15 Batesian Mimicry 1862 –Henry Walter Bates; “A Naturalist on the River Amazons” 16 8 3/24/2015 Batesian Mimicry Batesian mimics “cheat” –they lack toxins, venom, etc.
    [Show full text]
  • Predatory Behavior of Jumping Spiders
    Annual Reviews www.annualreviews.org/aronline Annu Rev. Entomol. 19%. 41:287-308 Copyrighl8 1996 by Annual Reviews Inc. All rights reserved PREDATORY BEHAVIOR OF JUMPING SPIDERS R. R. Jackson and S. D. Pollard Department of Zoology, University of Canterbury, Christchurch, New Zealand KEY WORDS: salticids, salticid eyes, Portia, predatory versatility, aggressive mimicry ABSTRACT Salticids, the largest family of spiders, have unique eyes, acute vision, and elaborate vision-mediated predatory behavior, which is more pronounced than in any other spider group. Diverse predatory strategies have evolved, including araneophagy,aggressive mimicry, myrmicophagy ,and prey-specific preycatch- ing behavior. Salticids are also distinctive for development of behavioral flexi- bility, including conditional predatory strategies, the use of trial-and-error to solve predatory problems, and the undertaking of detours to reach prey. Predatory behavior of araneophagic salticids has undergone local adaptation to local prey, and there is evidence of predator-prey coevolution. Trade-offs between mating and predatory strategies appear to be important in ant-mimicking and araneo- phagic species. INTRODUCTION With over 4000 described species (1 l), jumping spiders (Salticidae) compose by Fordham University on 04/13/13. For personal use only. the largest family of spiders. They are characterized as cursorial, diurnal predators with excellent eyesight. Although spider eyes usually lack the struc- tural complexity required for acute vision, salticids have unique, complex eyes with resolution abilities without known parallels in animals of comparable size Annu. Rev. Entomol. 1996.41:287-308. Downloaded from www.annualreviews.org (98). Salticids are the end-product of an evolutionary process in which a small silk-producing animal with a simple nervous system acquires acute vision, resulting in a diverse array of complex predatory strategies.
    [Show full text]
  • Hyperspectral Imaging of Cuttlefish Camouflage Indicates Good Color
    Hyperspectral imaging of cuttlefish camouflage indicates good color match in the eyes of fish predators Chuan-Chin Chiaoa,b,1, J. Kenneth Wickiserc, Justine J. Allena,d, Brock Genterc, and Roger T. Hanlona,e aMarine Biological Laboratory, Woods Hole, MA 02543; bDepartment of Life Science, National Tsing Hua University, Hsinchu, Taiwan 30013; cDepartment of Chemistry and Life Science, United States Military Academy, West Point, NY 10996; and Departments of dNeuroscience and eEcology and Evolutionary Biology, Brown University, Providence, RI 02912 Edited* by A. Kimball Romney, University of California, Irvine, CA, and approved April 13, 2011 (received for review December 30, 2010) Camouflage is a widespread phenomenon throughout nature and hyperspectral image is typically captured by scanning the 2D an important antipredator tactic in natural selection. Many visual sensor either spectrally or spatially in the third dimension to predators have keen color perception, and thus camouflage acquire the 3D data cube of which the z axis normally represents patterns should provide some degree of color matching in addition the reflectance spectrum of the corresponding point in the scene. to other visual factors such as pattern, contrast, and texture. Camouflage is the primary defense of coleoid cephalopods Quantifying camouflage effectiveness in the eyes of the predator (octopus, squid, and cuttlefish) and their rapidly adaptable body is a challenge from the perspectives of both biology and optical patterning system is among the most sophisticated in the animal imaging technology. Here we take advantage of hyperspectral kingdom (21–23). The expression of camouflaged body patterns imaging (HSI), which records full-spectrum light data, to simulta- in cuttlefish is a visually driven behavior.
    [Show full text]
  • Mimicry - Ecology - Oxford Bibliographies 12/13/12 7:29 PM
    Mimicry - Ecology - Oxford Bibliographies 12/13/12 7:29 PM Mimicry David W. Kikuchi, David W. Pfennig Introduction Among nature’s most exquisite adaptations are examples in which natural selection has favored a species (the mimic) to resemble a second, often unrelated species (the model) because it confuses a third species (the receiver). For example, the individual members of a nontoxic species that happen to resemble a toxic species may dupe any predators by behaving as if they are also dangerous and should therefore be avoided. In this way, adaptive resemblances can evolve via natural selection. When this phenomenon—dubbed “mimicry”—was first outlined by Henry Walter Bates in the middle of the 19th century, its intuitive appeal was so great that Charles Darwin immediately seized upon it as one of the finest examples of evolution by means of natural selection. Even today, mimicry is often used as a prime example in textbooks and in the popular press as a superlative example of natural selection’s efficacy. Moreover, mimicry remains an active area of research, and studies of mimicry have helped illuminate such diverse topics as how novel, complex traits arise; how new species form; and how animals make complex decisions. General Overviews Since Henry Walter Bates first published his theories of mimicry in 1862 (see Bates 1862, cited under Historical Background), there have been periodic reviews of our knowledge in the subject area. Cott 1940 was mainly concerned with animal coloration. Subsequent reviews, such as Edmunds 1974 and Ruxton, et al. 2004, have focused on types of mimicry associated with defense from predators.
    [Show full text]
  • Snakes of South-East Asia Including Myanmar, Thailand, Malaysia, Singapore, Sumatra, Borneo, Java and Bali
    A Naturalist’s Guide to the SNAKES OF SOUTH-EAST ASIA including Myanmar, Thailand, Malaysia, Singapore, Sumatra, Borneo, Java and Bali Indraneil Das First published in the United Kingdom in 2012 by Beaufoy Books n n 11 Blenheim Court, 316 Woodstock Road, Oxford OX2 7NS, England Contents www.johnbeaufoy.com 10 9 8 7 6 5 4 3 2 1 Introduction 4 Copyright © 2012 John Beaufoy Publishing Limited Copyright in text © Indraneil Das Snake Topography 4 Copyright in photographs © [to come] Dealing with Snake Bites 6 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publishers. About this Book 7 ISBN [to come] Glossary 8 Edited, designed and typeset by D & N Publishing, Baydon, Wiltshire, UK Printed and bound [to come] Species Accounts and Photographs 11 Checklist of South-East Asian Snakes 141 Dedication Nothing would have happened without the support of the folks at home: my wife, Genevieve V.A. Gee, and son, Rahul Das. To them, I dedicate this book. Further Reading 154 Acknowledgements 155 Index 157 Edited and designed by D & N Publishing, Baydon, Wiltshire, UK Printed and bound in Malaysia by Times Offset (M) Sdn. Bhd. n Introduction n n Snake Topography n INTRODUCTION Snakes form one of the major components of vertebrate fauna of South-East Asia. They feature prominently in folklore, mythology and other belief systems of the indigenous people of the region, and are of ecological and conservation value, some species supporting significant (albeit often illegal) economic activities (primarily, the snake-skin trade, but also sale of meat and other body parts that purportedly have medicinal properties).
    [Show full text]
  • Red-Tailed Green Ratsnake Gonyosoma Oxycephalum
    SEAVR 2019: 033‐034 ISSN : 2424‐8525 Date of publication: 24 February 2019 Hosted online by ecologyasia.com Red‐tailed Green Ratsnake Gonyosoma oxycephalum predation on White‐bellied Woodpecker in the Philippines Allan Gil S. FERNANDO & Emerson Y. SY [email protected] (Fernando), [email protected] (Sy) Observer: Allan Gil S. Fernando. Photograph by: Allan Gil S. Fernando. Subjects identified by: Allan Gil S. Fernando, Emerson Y. Sy. Location: : Nabasan Trail, Municipality of Subic, Zambales Province, Luzon Island, Philippines. Elevation: 40 metres. Habitat: Lowland secondary forest. Date and time: 26 January 2019, 10:14 hrs. Identity of subjects: i) Red‐tailed Green Ratsnake, Gonyosoma oxycephalum (Reptilia: Squamata: Colubridae). ii) White‐bellied Woodpecker, Dryocopus javensis (Aves: Piciformes: Picidae) Description of record: A green snake was observed hanging from a horizontal branch approximately three meters above the ground, whilst constricting a White‐bellied Woodpecker. Another White‐bellied Woodpecker nearby was making non‐stop alarm calls. The bird ceased struggling after three minutes and the snake started its attempt to swallow it head first. The snake was able to swallow the head of the bird, but regurgitated it after a few minutes. It made another attempt to swallow the bird, but this was also unsuccessful. The two swallowing attempts lasted for 17 minutes. Observation was discontinued immediately after the second regurgitation, thus the actual consumption of the bird by the snake was not observed. Remarks: The snake was identified as Gonyosoma oxycephalum based on (i) uniform green colour, (ii) tail is light reddish brown colour and (iii) head is elongated and distinct from the neck (Taylor, 1922).
    [Show full text]
  • The Speciation History of Heliconius: Inferences from Multilocus DNA Sequence Data
    The speciation history of Heliconius: inferences from multilocus DNA sequence data by Margarita Sofia Beltrán A thesis submitted for the degree of Doctor of Philosophy of the University of London September 2004 Department of Biology University College London 1 Abstract Heliconius butterflies, which contain many intermediate stages between local varieties, geographic races, and sympatric species, provide an excellent biological model to study evolution at the species boundary. Heliconius butterflies are warningly coloured and mimetic, and it has been shown that these traits can act as a form of reproductive isolation. I present a species-level phylogeny for this group based on 3834bp of mtDNA (COI, COII, 16S) and nuclear loci (Ef1α, dpp, ap, wg). Using these data I test the geographic mode of speciation in Heliconius and whether mimicry could drive speciation. I found little evidence for allopatric speciation. There are frequent shifts in colour pattern within and between sister species which have a positive and significant correlation with species diversity; this suggests that speciation is facilitated by the evolution of novel mimetic patterns. My data is also consistent with the idea that two major innovations in Heliconius, adult pollen feeding and pupal-mating, each evolved only once. By comparing gene genealogies from mtDNA and introns from nuclear Tpi and Mpi genes, I investigate recent speciation in two sister species pairs, H. erato/H. himera and H. melpomene/H. cydno. There is highly significant discordance between genealogies of the three loci, which suggests recent speciation with ongoing gene flow. Finally, I explore the phylogenetic relationships between races of H. melpomene using an AFLP band tightly linked to the Yb colour pattern locus (which determines the yellow bar in the hindwing).
    [Show full text]