DOCSLIB.ORG

I June 1995 Asiatic Herpetological Research Systematics of the Vipers

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
I June 1995 Asiatic Herpetological Research Systematics of the Vipers I June 1995 Asiatic Herpetological Research Vol. 6, pp. 1-26 i Systematics of the Vipers of the Caucasus: Polymorphism or Sibling Species? G o r a n N i l s o n 1, B o r is S. T u n i y e v 2, N ik o l a i O r l o v 3, m a t s h o g g r e n 4 a n d C l a e s A n d r e n 1 department of Zoology, Goteborg University, Sweden 2Caucasian State Biosphere Reserve, Sochi, Russia 3Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia 4Department of Genetics, University of Uppsala, Russia Abstract. -Inter- and intramorphological variation were examined in sympatric and allopatric polymorphic and monomorphic populations of the Vipera ursinii and Vipera kaznakovi complexes. The alpine Vipera dinniki populations in upper Great Caucasus show a pronounced, and to a certain extent geographical, polymorphism. Color patterns include among others 'kaznakovi', 'tigrina', 'berus', 'bronze', and 'ursinii' types. Several of these patterns can be represented within the same litter in certain populations. Vipera dinniki is sympatric with the Caucasian representative of the Vipera ursinii complex in some areas. This last taxon shows a similar degree of polymorphism, which is unique for this complex, and due to morphological and molecular distinction, we consider it to be a Caucasian evolutionary species within the ursinii complex - Vipera lotievi sp.n. Key Words: Reptilia, Squamata, Viperidae, Vipera dinniki, V. kaznakovi, V. ursinii, V. renardi, V. lotievi sp.n., Caucasus, Russia, Georgia, taxonomy, morphology, polymorphism. Introduction synonym of V. kaznakovi until Vedmederja et al (1986) recognized it as a separate The taxonomy of the vipers of Caucasus species inhabiting alpine and subalpine has for a long time been confusing and meadows in the Caucasus, and thus contradictory. According to the traditional restricting V. kaznakovi to lower altitudes in view a single species, Vipera kaznakovi, is western Caucasus and adjacent moist distributed in the moist and warm lowlands lowland habitats along the eastern Black Sea of the western Caucasus as well as in the coast. Thereby the problem of the gradual mountain valleys towards the east. In the transformation from V. kaznakovi in the east the habitat is drier and along the range west of the Caucasus to V. ursinii in the east the vipers gradually change toward Vipera is restricted to the high Caucasus ursinii in appearance. In the east Caucasus populations, now including V. dinniki and only this last viper was supposed to occur. V. ursinii. Vipera kaznakovi is well defined Thus there seemed to be a somewhat clinal and restricted in distribution, and transformation from "pure" V. kaznakovi in geographically separated from all the other the west to "pure" V. ursinii in the east. viper species in the region. Vipers from the intermediate region could be difficult to determine. Within the same The complex history of nomenclature locality some specimens look like V. and taxonomy has been clarified to a great kaznakovi, other ones are more like V. part in some recent publications (see Orlov ursinii, while still some can be intermediate. & Tuniyev, 1986; 1990), together with hypotheses about the phylogeny of this Nikolsky (1913) separated the alpine group. Concurrently the need of genetic populations into the taxon Vipera berus studies was stressed, and this has led to the d in n iki, which was based on alpine present work where we in a series of papers specimens of the conventional V. kaznakovi intend to clarify the taxonomy and evolution from high altitudes in the western Caucasus of the vipers of this region. The work is (Malaya Laba River- terra typica restricta planned to have a broad perspective and Svanetia) as well as from other places. including phenetic and phylogenetic The name dinniki was long considered as a analyses, habitat choice, niche-breadth, and Vol. 6, p. 2 Asiatic Herpetological Research June 1995 reproduction. Different methods take Altogether about 300 preserved or live different time and in this first paper the specimens of vipers from the Caucasus and morphology is reexamined, based on adjacent regions have been seen during the available material in Museum collections and study. Joint field trips were made in freshly collected material from a number of different parts of the Caucasus in 1990 and new places, as well as on inheritance of 1992, but two of us (Tuniyev and Orlov) color pattern. Genetic structures based on have performed extensive research in the phenetic analyses of allozyme data are also region prior to that. For morphometric presented. studies 183 preserved snakes within the ursinii and kaznakovi complexes have been The morphological distinction between examined more carefully, and for most of Vipera dinniki and V. kaznakovi has been these specimens 30 different items of data presented elsewhere (Orlov and Tuniyev, have been collected. This information was 1986; 1990) and will not be repeated in this used, down to population level, in study. In the present paper we are focusing morphological descriptions, taxonomical on patterns of morphological and molecular analyses and conclusions about variation within and between the different zoogeography and range overlap. populations in the Caucasus, and the Inheritance of color pattern was studied taxonomy of these populations. based on 23 pregnant females and their offspring. Material and Methods Data collected were: total length and tail The work has mainly been a study of length; number of preventrals, ventrals, variation in external morphology, allozymes subcaudals, anterior and mid-body dorsal and reproduction in order to reveal patterns scale rows, apical plates, supralabials, of sympatry and sibling species. Additional sublabials, circumocular scales, loreals, genetic studies will follow when material second chinshields, mentals, crown scales becomes available in suitable samples (=intercanthals + intersupraoculars), and (presently delayed due to political reasons). zig-zag windings in dorsal band. Further Thus live and preserved museum material rostral index (height/breadth) and head index has been examined concordantly with (breadth/length) were calculated. Division studies of reproduction in the laboratory. of parietals, frontal, and nasalia was noted, as was the color of dorsal and ventral sides, and iris (in live specimens). Further, upper Additional preserved material used in preocular size; and head, labial and lateral this study originates from the Natural History Museum in Goteborg (GNM); body patterns, as well as distinctiveness of Dipl.-Biol. F. J. Obst, Staatliches Museum canthus rostralis were examined. Details fur Naturkunde, Dresden (MTKD D); Aram about these methods are found in Nilson and Andren (1986). Agasian, Zoological Institute, Academy of Sciences, Eriwan, Armenia. Abbreviations Morphology/phenetics for museums as used in the text are: CNR- Caucasian State Biosphere Reserve, Standard errors accompanying mean Collection of Boris Tuniyev at Yew-box character ratios were used as relative Groove, Sochi; GNM- Goteborg Natural History Museum, Goteborg; MTKD- measurement of dispersion. For the analysis of intra- and interpopulational Staatliches Museum for Naturkunde, morphological variation (phenetic analysis) Dresden; ZIEr- Zoological Institute, Academy of Sciences, Eriwan; ZIG- the samples were divided into subsamples depending on questions raised. Thus Department of Zoology, University of Goteborg; Goteborg- (authors' collection, besides an analysis of morphological variation also a pattern confirming or which later will be incorporated in GNM); ZIN- Zoological Institute, Academy of rejecting the present taxonomic pattern could be achieved. This pattern could also be Sciences, St. Petersburg. June 1995 Asiatic Herpetological Research Vol. 6, p. 3 TABLE 1. Number of specimens used in the genetic analyses and localities (Russia if nothing else is stated) for the examined taxa. kaznakovi: 1. Dagomys, north of Sochi. Six specimens. 2. Rudorova, inland locality, 900 m alt. Four specimens. dinniki: 3. Fisht/Oshten, the westernmost locality of the main Caucasus range. Seven specimens. 4. Lake Impsi, 1,980 m. alt., at a tributary to the Little Laba River on the northern slope of the main range. Seven specimens. 5. Aishkha-П on the southern slope of the main range. Three specimens. 6. Lake Kardyvach at upper Mzymta River on the southern slope of the main range Seventeen specimens. lotieviv. 7. Armkhi, Checheno-Ingushetia. 2,000 m altitude. Seven specimens. berus: 8. Uppsala (terra typica), Sweden. Eleven specimens. eriwanensis: 9. pooled sample from Asbua and Cildir, Kars, east Turkey; and Sevan, Armenia. Six specimens. supported or rejected by the parallel among the eight Caucasus populations biochemical studies. Thereby it is possible studied (see Table 1, for locality data and to state or reject the occurrence of sample sizes). A potential risk of sampling convergent or parallel evolution, i.e. sibling error due to syntopic occurrence of two taxa species. may be avoided by testing observed genotype distribution within a locality Estimation of the different color pattern against Hardy-Weinberg expectations (see frequencies in local populations was based results). Fresh or frozen tissues (-75°C) on observations during the field work. from liver and skeletal muscle were Small museum samples collected by others homogenized in distilled water. The extracts were not included in this analysis due to were centrifuged for 10 min at 10,000 rpm uncertainty of randomness in sampling and 4°C
Load more

Recommended publications
  • Rare Species of Shield-Head Vipers in the Caucasus
    Nature Conservation Research. Заповедная наука 2016. 1 (3): 11–25 RARE SPECIES OF SHIELD-HEAD VIPERS IN THE CAUCASUS Boris S. Tuniyev Sochi National Park, Russia e-mail: [email protected] Received: 03.10.2016 An overview is presented on shield-head vipers of the genus Pelias distributed in the post-Soviet countries of the Caucasian Ecoregion. The assessment presents the current conservation status and recommendations to vipers’ ter- ritorial protection. Key words: Caucasian Ecoregion, shield-head vipers, current status, protection. Introduction The Caucasian Ecoregion (the territory south- to-landscape descriptions (Tunieyv B.S. et al., 2009; ward from the Kuma-Manych depression to north- Tuniyev S.B. et al., 2012, 2014). The stationary works eastern Turkey and northwestern Iran) is the centre (mostly on the territory of the Caucasian State Nature of taxonomic diversity of shield-head vipers within Biosphere reserve and Sochi National Park) conducted the genus Pelias Merrem, 1820, of which 13–18 a study of the microclimatic features of vipers’ habitats species are found here. Without exception, all spe- including temperature and humidity modes of air and cies have a status of the different categories of rare- the upper soil horizon. The results were compared with ness, they are included on the IUCN Red list, or in thermobiological characteristics of the animals (Tuni- the current and upcoming publication of National yev B.S. & Unanian, 1986; Tuniyev B.S. & Volčik, and Regional Red Data Books. Besides the shield- 1995). In a number of cases difficult to determine the head vipers the Caucasian Ecoregion inhabit three taxonomic affiliation, in addition to the classical meth- representatives of mountain vipers of the genus ods of animal morphology and statistics, biochemistry Montivipera Nilson, Tuniyev, Andren, Orlov, Joger and molecular-genetic analysis methods have been ap- & Herrman, 1999 (M.
    [Show full text]
  • Status and Protection of Globally Threatened Species in the Caucasus
    STATUS AND PROTECTION OF GLOBALLY THREATENED SPECIES IN THE CAUCASUS CEPF Biodiversity Investments in the Caucasus Hotspot 2004-2009 Edited by Nugzar Zazanashvili and David Mallon Tbilisi 2009 The contents of this book do not necessarily reflect the views or policies of CEPF, WWF, or their sponsoring organizations. Neither the CEPF, WWF nor any other entities thereof, assumes any legal liability or responsibility for the accuracy, completeness, or usefulness of any information, product or process disclosed in this book. Citation: Zazanashvili, N. and Mallon, D. (Editors) 2009. Status and Protection of Globally Threatened Species in the Caucasus. Tbilisi: CEPF, WWF. Contour Ltd., 232 pp. ISBN 978-9941-0-2203-6 Design and printing Contour Ltd. 8, Kargareteli st., 0164 Tbilisi, Georgia December 2009 The Critical Ecosystem Partnership Fund (CEPF) is a joint initiative of l’Agence Française de Développement, Conservation International, the Global Environment Facility, the Government of Japan, the MacArthur Foundation and the World Bank. This book shows the effort of the Caucasus NGOs, experts, scientific institutions and governmental agencies for conserving globally threatened species in the Caucasus: CEPF investments in the region made it possible for the first time to carry out simultaneous assessments of species’ populations at national and regional scales, setting up strategies and developing action plans for their survival, as well as implementation of some urgent conservation measures. Contents Foreword 7 Acknowledgments 8 Introduction CEPF Investment in the Caucasus Hotspot A. W. Tordoff, N. Zazanashvili, M. Bitsadze, K. Manvelyan, E. Askerov, V. Krever, S. Kalem, B. Avcioglu, S. Galstyan and R. Mnatsekanov 9 The Caucasus Hotspot N.
    [Show full text]
  • Status and Protection of Globally Threatened Species in the Caucasus
    STATUS AND PROTECTION OF GLOBALLY THREATENED SPECIES IN THE CAUCASUS CEPF Biodiversity Investments in the Caucasus Hotspot 2004-2009 Edited by Nugzar Zazanashvili and David Mallon Tbilisi 2009 The contents of this book do not necessarily re ect the views or policies of CEPF, WWF, or their sponsoring organizations. Neither the CEPF, WWF nor any other entities thereof, assumes any legal liability or responsibility for the accuracy, completeness, or usefulness of any information, product or process disclosed in this book. Citation: Zazanashvili, N. and Mallon, D. (Editors) 2009. Status and Protection of Globally Threatened Species in the Caucasus. Tbilisi: CEPF, WWF. Contour Ltd., 232 pp. ISBN 978-9941-0-2203-6 Design and printing Contour Ltd. 8, Kargareteli st., 0164 Tbilisi, Georgia December 2009 The Critical Ecosystem Partnership Fund (CEPF) is a joint initiative of l’Agence Française de Développement, Conservation International, the Global Environment Facility, the Government of Japan, the MacArthur Foundation and the World Bank. This book shows the effort of the Caucasus NGOs, experts, scienti c institutions and governmental agencies for conserving globally threatened species in the Caucasus: CEPF investments in the region made it possible for the rst time to carry out simultaneous assessments of species’ populations at national and regional scales, setting up strategies and developing action plans for their survival, as well as implementation of some urgent conservation measures. Contents Foreword 7 Acknowledgments 8 Introduction CEPF Investment in the Caucasus Hotspot A. W. Tordoff, N. Zazanashvili, M. Bitsadze, K. Manvelyan, E. Askerov, V. Krever, S. Kalem, B. Avcioglu, S. Galstyan and R. Mnatsekanov 9 The Caucasus Hotspot N.
    [Show full text]
  • Biodiversity Assessment for Georgia
    Biodiversity Assessment for Georgia Task Order under the Biodiversity & Sustainable Forestry IQC (BIOFOR) USAID C ONTRACT NUMBER: LAG-I-00-99-00014-00 SUBMITTED TO: USAID WASHINGTON E&E BUREAU, ENVIRONMENT & NATURAL RESOURCES DIVISION SUBMITTED BY: CHEMONICS INTERNATIONAL INC. WASHINGTON, D.C. FEBRUARY 2000 TABLE OF CONTENTS SECTION I INTRODUCTION I-1 SECTION II STATUS OF BIODIVERSITY II-1 A. Overview II-1 B. Main Landscape Zones II-2 C. Species Diversity II-4 SECTION III STATUS OF BIODIVERSITY CONSERVATION III-1 A. Protected Areas III-1 B. Conservation Outside Protected Areas III-2 SECTION IV STRATEGIC AND POLICY FRAMEWORK IV-1 A. Policy Framework IV-1 B. Legislative Framework IV-1 C. Institutional Framework IV-4 D. Internationally Supported Projects IV-7 SECTION V SUMMARY OF FINDINGS V-1 SECTION VI RECOMMENDATIONS FOR IMPROVED BIODIVERSITY CONSERVATION VI-1 SECTION VII USAID/GEORGIA VII-1 A. Impact of the Program VII-1 B. Recommendations for USAID/Georgia VII-2 ANNEX A SECTIONS 117 AND 119 OF THE FOREIGN ASSISTANCE ACT A-1 ANNEX B SCOPE OF WORK B-1 ANNEX C LIST OF PERSONS CONTACTED C-1 ANNEX D LISTS OF RARE AND ENDANGERED SPECIES OF GEORGIA D-1 ANNEX E MAP OF LANDSCAPE ZONES (BIOMES) OF GEORGIA E-1 ANNEX F MAP OF PROTECTED AREAS OF GEORGIA F-1 ANNEX G PROTECTED AREAS IN GEORGIA G-1 ANNEX H GEORGIA PROTECTED AREAS DEVELOPMENT PROJECT DESIGN SUMMARY H-1 ANNEX I AGROBIODIVERSITY CONSERVATION IN GEORGIA (FROM GEF PDF GRANT PROPOSAL) I-1 SECTION I Introduction This biodiversity assessment for the Republic of Georgia has three interlinked objectives: · Summarizes the status of biodiversity and its conservation in Georgia; analyzes threats, identifies opportunities, and makes recommendations for the improved conservation of biodiversity.
    [Show full text]
  • National Biodiversity Strategy and Action Plan of Georgia
    Biodiversity Strategy and Action Plan - Georgia – Tbilisi, 2005 Foreword Georgia signed the Convention on Biological Diversity in 1994, thus accepting responsibility to safeguard the nation’s rich diversity of plant, animal, and microbial life, to begin using biological resources in sustainable way, and to ensure equitable sharing of benefits from biodiversity. Later the country joined other conventions including the Convention on Climate Change, the Ramsar Convention on Wetlands, CITES and the Bonn Convention. As a signatory to these important international environmental treaties, Georgia enters the world scene with the potential for joining the most advanced nations in the field of environmental protection. At the present moment of transition, Georgia has a unique opportunity to use the early experiences of other countries, and avoid irreversible changes in the quality of its environment. The national legislation on environmental protection adopted over the past few years provides an adequate legal basis for this, although further elaboration and reinforcement of the existing legislation is needed. With the Ministry of Environment being currently reorganised and assuming broader responsibilities, Georgia’s institutional arrangements for environmental protection already has the necessary structure for improving the quality of the environment throughout the country. The role of non-governmental groups has been very important in resolving problems related to nature conservation. Georgia has shown an excellent example of co-operation between governmental and non-governmental organizations in the field of environment, and particularly in the field of biodiversity conservation. After signing the Convention on Biological Diversity, the Georgian Government immediately acted to develop a Biodiversity Country Study, in partnership with UNEP, and implemented by NACRES, a local conservation organisation.
    [Show full text]
  • Expert Advice on Terrestrial Biodiversity Conservation, Land Take and Compensation Report
    Dariali Hydropower Plant Project Expert Advice on Terrestrial Biodiversity Conservation, Land Take and Compensation Report Tbilisi 2013 INTRODUCTION Botanical and Zoological surveys have been carried in order to address the key data gap existing in ESIA of Dariali HPP Project from the Biodiversity standpoint that is provided in the “Expert Advice on Terrestrial Biodiversity Conservation, Land Take and Compensation Report” that includes two Annexes: I. Survey and comparative analysis of flora and vegetation of Dariali Hydropower Plant Project Corridor and compensation sites (carried out by Botanists: Dr Mariam Kimeridze and Mr David Chelidze) and II. Survey and comparative analysis of fauna of Dariali Hydropower Plant Project Corridor and compensation sites (carried out by Zoologists: Dr Alexander Bukhnikashvili, Dr Teimuraz Kokosadze and Mrs Marine Gioshvili). Three small areas of land were removed from the Kazbegi National Park for the Dariali HPP construction totaling 8,737 ha that belonged to the area within the Boundaries of Traditional Use Zone of the KNP. Three territories have been added to the Protected Areas as compensation areas for the land lost at Dariali due to HPP: Nature Monument of Sakhiznari Cliff Columns-335,7ha, Nature Monument of the Abano Mineral Lake-0,04 ha and Nature Monument of the Truso Travertines-4,2 ha. For additional information with regard to impact of Dariali HPP construction on KNP please see the report prepared by Dr Mariam Kimeridze “Impact of Dariali HPP on Kazbegi National Oark Traditional USE Zone” dated 31.05.2013). The detailed botanical and zoological studies were carried out in the river Tergi gorge within the borders of the Project Corridor and Compensation Sites.
    [Show full text]
  • Calibrating the Tree of Vipers Under the Fossilized Birth-Death Model Jiří Šmíd 1,2,3,4 & Krystal A
    www.nature.com/scientificreports OPEN Calibrating the tree of vipers under the fossilized birth-death model Jiří Šmíd 1,2,3,4 & Krystal A. Tolley 1,5 Scaling evolutionary trees to time is essential for understanding the origins of clades. Recently Received: 18 September 2018 developed methods allow including the entire fossil record known for the group of interest and Accepted: 15 February 2019 eliminated the need for specifying prior distributions for node ages. Here we apply the fossilized birth- Published: xx xx xxxx death (FBD) approach to reconstruct the diversifcation timeline of the viperines (subfamily Viperinae). Viperinae are an Old World snake subfamily comprising 102 species from 13 genera. The fossil record of vipers is fairly rich and well assignable to clades due to the unique vertebral and fang morphology. We use an unprecedented sampling of 83 modern species and 13 genetic markers in combination with 197 fossils representing 28 extinct taxa to reconstruct a time-calibrated phylogeny of the Viperinae. Our results suggest a late Eocene-early Oligocene origin with several diversifcation events following soon after the group’s establishment. The age estimates inferred with the FBD model correspond to those from previous studies that were based on node dating but FBD provides notably narrower credible intervals around the node ages. Viperines comprise two African and an Eurasian clade, but the ancestral origin of the subfamily is ambiguous. The most parsimonious scenarios require two transoceanic dispersals over the Tethys Sea during the Oligocene. Scaling phylogenetic trees to time is one of the major challenges in evolutionary biology. Reliable estimates for the age of evolutionary events are essential for addressing a wide array of questions, such as deciphering micro- and macroevolutionary processes, identifying drivers of biodiversity patterns, or understanding the origins of life1.
    [Show full text]
  • Review Species List of the European Herpetofauna – 2020 Update by the Taxonomic Committee of the Societas Europaea Herpetologi
    Amphibia-Reptilia 41 (2020): 139-189 brill.com/amre Review Species list of the European herpetofauna – 2020 update by the Taxonomic Committee of the Societas Europaea Herpetologica Jeroen Speybroeck1,∗, Wouter Beukema2, Christophe Dufresnes3, Uwe Fritz4, Daniel Jablonski5, Petros Lymberakis6, Iñigo Martínez-Solano7, Edoardo Razzetti8, Melita Vamberger4, Miguel Vences9, Judit Vörös10, Pierre-André Crochet11 Abstract. The last species list of the European herpetofauna was published by Speybroeck, Beukema and Crochet (2010). In the meantime, ongoing research led to numerous taxonomic changes, including the discovery of new species-level lineages as well as reclassifications at genus level, requiring significant changes to this list. As of 2019, a new Taxonomic Committee was established as an official entity within the European Herpetological Society, Societas Europaea Herpetologica (SEH). Twelve members from nine European countries reviewed, discussed and voted on recent taxonomic research on a case-by-case basis. Accepted changes led to critical compilation of a new species list, which is hereby presented and discussed. According to our list, 301 species (95 amphibians, 15 chelonians, including six species of sea turtles, and 191 squamates) occur within our expanded geographical definition of Europe. The list includes 14 non-native species (three amphibians, one chelonian, and ten squamates). Keywords: Amphibia, amphibians, Europe, reptiles, Reptilia, taxonomy, updated species list. Introduction 1 - Research Institute for Nature and Forest, Havenlaan 88 Speybroeck, Beukema and Crochet (2010) bus 73, 1000 Brussel, Belgium (SBC2010, hereafter) provided an annotated 2 - Wildlife Health Ghent, Department of Pathology, Bacteriology and Avian Diseases, Ghent University, species list for the European amphibians and Salisburylaan 133, 9820 Merelbeke, Belgium non-avian reptiles.
    [Show full text]
  • On Placing and Thermobiology of Polymorphic Population of Vipera Dinniki (Nikolsky, 1913) (Serpcntes, Viperidae) at Uppermost Msymta-River Valley
    Llorente et al. (Eds). Scientia Herpelologica 1995:244-249 On placing and thermobiology of polymorphic population of Vipera dinniki (Nikolsky, 1913) (Serpcntes, Viperidae) at uppermost Msymta-river valley B. TUNIYEV1 & S. VOLCIK2 ' Caucasian State Biosphere Reserve, K.Marx sir. 8a, Sochi, RU-354341, Russia 2 Praha University, post. pr. 79, 12000, Praha 2, posta Praha 2, Ceska republica Abstract: Investigated area has about 5 hectares in subalpine belt (1750-1850 m) of uppermost Mzymta-river valley, West Caucasus. This area has two morains, rocky outcrops, high-grass and and mix elfin woodland-meadows. Both morains and rochs arc the places of hibernation. There are distinguished 4 phenotypes o f Vipera dinniki. Among thrm "tigrina"-morph prevails over "nebulosa" - and "bronze"-morphs. The most rare morph is "kaznakovi"-morph. Sexes ratio close to 1 (28 cW : 33 ¥¥). Adult vipers predominante in the age structure of population. Most not-numerous group is semiadults. Depend on slope exposition the begining and the end ofdaily activity have distinctions on the various places of this area but total length of surface activity was similar in each places and had 35% ofdaily cycle. It was picked out the distinctions in territorial placing between sexes and different age-grades. Thermobiological data of this area demonstrated the exceeding of the body temperature of active males than those of females. Body temperature of representatives of both sexes has stricty dependence on ground temperature than on air temperature. Results testify to existence of differences in thermobiology of various morphs. Key words: Caucasus, Vipers, Morphs, Thermobiology. Introduction M a t e r i a l a n d m e t h o d s Snake's relations with environmental Vipera dinniki was studied at the begining of constrains (as poikilothems) are interesting river Mzymta, vicinity of Sochi, Western vvithing temperature balance firstly.
    [Show full text]
  • BIOGEOGRAFÍA Y ECOLOGÍA DE LAS VÍBORAS IBÉRICAS (Vipera Aspis, V
    TESIS DOCTORAL BIOGEOGRAFÍA Y ECOLOGÍA DE LAS VÍBORAS IBÉRICAS (Vipera aspis, V. latastei y V. seoanei ) EN UNA ZONA DE CONTACTO EN EL NORTE PENINSULAR BIOGEOGRAPHY AND ECOLOGY OF THE IBERIAN VIPERS (Vipera aspis, V. latastei, V. seoanei ) IN A CONTACT ZONE IN NORTHERN IBERIAN PENINSULA FERNANDO MARTÍNEZ FREIRÍA UNIVERSIDAD DE SALAMANCA FACULTAD DE BIOLOGÍA DEPARTAMENTO DE BIOLOGÍA ANIMAL, PARASITOLOGÍA, ECOLOGÍA, EDAFOLOGÍA Y QUÍMICA AGRÍCOLA BIOGEOGRAFÍA Y ECOLOGÍA DE LAS VÍBORAS IBÉRICAS (Vipera aspis, V. latastei y V. seoanei ) EN UNA ZONA DE CONTACTO EN EL NORTE PENINSULAR BIOGEOGRAPHY AND ECOLOGY OF THE IBERIAN VIPERS (Vipera aspis, V. latastei and V. seoanei ) IN A CONTACT ZONE IN NORTHERN IBERIAN PENINSULA Memoria para optar al grado de Doctor en Biología por la Universidad de Salamanca, presentada por Fernando Martínez Freiría Fdo. Fernando Martínez Freiría Salamanca, 2009 NOTAS PREVIAS Para la realización de esta tesis doctoral he sido beneficiario durante cuatro años, desde febrero de 2004 hasta enero de 2008, de una beca pre-doctoral para la Formación de Profesorado Universi- tario (FPU) del Ministerio de Educación, Ciencia y Deporte (referencia AF 2633-2003). En la elaboración de esta memoria de tesis doctoral se han utilizado varios artículos publicados en revistas científicas y, por ello, esta memoria presenta capítulos en lengua inglesa y otras partes en lengua castellana. En cumplimiento del acuerdo adoptado por la Comisión de Doctorado y Pos- grado de la Universidad de Salamanca, por el que se permite la presentación de la memoria de te- sis doctoral en dos idiomas, adjunto un resumen en castellano, antes de cada capítulo en lengua inglesa.
    [Show full text]
  • Winter Activity of the Hime-Habu (Ovophis Okinavensis) in the Humid Subtropics: Foraging on Breeding Anurans at Low Temperatures
    WINTER ACTIVITY OF THE HIME-HABU (OVOPHIS OKINAVENSIS) IN THE HUMID SUBTROPICS: FORAGING ON BREEDING ANURANS AT LOW TEMPERATURES AKIRA MORI1, MAMORU TODA1,3, AND HIDETOSHI OTA2 ABSTRACT: Snakes are ectothermic and rely on external heat sources to elevate their body temperature to levels required for various activities. Because of this constraint, snakes living at high latitudes and/or elevations cease their activity during cold weather, and in some cases exhibit physiological adaptations to the cold climate. Ovophis okinavensis is a small, nocturnal pitviper found in the subtropical region of the Ryukyu Archipelago, Japan, where the ambient temperature has a distinct seasonal fluctuation. We studied seasonal activity patterns, body temperatures, and food habits of O. okinavensis along a mountain stream from December 1996 to March 2000. Preliminary observations were also made with captive snakes to test the hypothesis that O. okinavensis has a physiological adaptation to efficiently digest prey at low temperatures. We found that O. okinavensis is most active at night during the cool season (from December to February), when two species of ranid frogs (Rana narina and_ R. okinavana) aggregate at a stream for reproduction. The body temperature of O. okinavensis ranged from 11.2 to 23.8°C (x = 16.1°C). Ninety-eight percent of the stomach contents consisted of frogs, and 37.0 % and 54.7 % were identified as R. narina and R. okinavana, respectively. The appearance of O. okinavensis coincided with breeding aggregations of these frogs, and extensive predation on the frogs by the snakes strongly suggests that winter activity in O. okinavensis at low body temperatures is associated with foraging.
    [Show full text]
  • Table S1. Number of Photographs, Rational for Species Exclusion And
    Table S1. Number of photographs, rational for species exclusion and source of distribution maps for each of the 35 venomous species and 87 non-venomous species of the Western Palearctic snakes, plus the 80 foreign species of the foreign class. Number of Rational for Venomousness Species name Distribution map sources images species exclusion Atractaspis engaddensis 41 (1) Bitis arietans 40 (2) Cerastes cerastes 53 (3) Cerastes gasperettii 40 (1) Cerastes vipera 50 (1) Daboia mauritanica 48 (1) Daboia palaestinae 54 (1) Echis carinatus 45 (3) Echis coloratus 40 (3) Echis pyramidum 52 (1) Gloydius halys 40 (3) Macrovipera lebetina 45 (3) Montivipera bornmuelleri 40 (1) Montivipera bulgardaghica 81 (1) Montivipera raddei 40 (1) Montivipera wagneri 40 (1) Montivipera xanthina 47 (1) Naja haje 40 (3) Naja nubiae 41 (3) Pseudocerates fieldi 40 (1) Vipera ammodytes 84 (1) Venomous Vipera anatolica 45 (1) Vipera aspis 84 (1) Vipera berus 64 (3) Vipera darevskii 40 (1) Vipera dinniki 41 (1) Vipera eriwanensis 40 (1) Vipera graeca 40 (4) Vipera kaznakovi 58 (1) Vipera latastei 81 (1) Vipera renardi 52 (5) Vipera seoanei 70 (1) Vipera ursinii 40 (1) Walterinnesia aegyptia 40 (1) Walterinnesia morgani 26 (3) Foreign species class : Acanthophis cryptamydros Acanthophis laevis Acanthophis pyrrhus Acanthophis wellsei Antairoserpens warro Antaresia childreni Antaresia maculosa 80 Antaresia perthensis Austrelaps ramsayi Austrelaps superbus Boiga irregularis Brachyurophis australis Brachyurophis fasciolatus Brachyurophis incinctus Brachyurophis morrisi Brachyurophis
    [Show full text]