Identification and Characterization of Two Novel Syncytin-Like Retroviral Envelope Genes, Captured for a Possible Role in the At

Total Page:16

File Type:pdf, Size:1020Kb

Identification and Characterization of Two Novel Syncytin-Like Retroviral Envelope Genes, Captured for a Possible Role in the At Identification and characterization of 106 S two novel syncytin-like retroviral SACL envelope genes, captured for a 8 possible role in the atypical structure : 201 of the hyena placenta and in the NNT emergence of the non-mammalian Mabuya lizard placenta Thèse de doctorat de l'Université Paris-Saclay préparée à l'UMR 9196, Gustave Roussy École doctorale n°582 cancérologie: biologie, médecine, santé (CBMS) Spécialité de doctorat: aspects moléculaires et cellulaires de la biologie Thèse présentée et soutenue à Villejuif, le 23 mai 2018, par Mathis Funk Composition du Jury : Uriel Hazan Professeur des université, ENS Paris-Saclay (– UMR 8113) Président Jean-Luc Battini Directeur de recherche, IRIM (– UMR 9004) Rapporteur Olivier Schwartz Directeur de recherche, Institut Pasteur (– UMR 3569) Rapporteur Pascale Chavatte-Palmer Directrice de recherche, INRA (– UMR 1198) Examinatrice François Mallet Directeur de recherche, bioMérieux (– EA 7426) Examinateur Thierry Heidmann Directeur de recherche, CNRS (– UMR 9196) Directeur de thèse Acknowledgments I would first like to thank the members of the jury for taking the time to read the present manuscript, which turned out a bit longer than I had planned. I would like to thank Uriel Hazan for accepting to be the president of this jury, book-ending his involvement in my studies. What had started at the ENS Cachan and continued during my Master’s degree at the Institut Pasteur, finally reaches its culmination with the present work, on a topic that Uriel suggested I look into. I would like to sincerely thank Jean-Luc Battini and Olivier Schwartz for their critical reading and evaluation of the present manuscript and their positive feedback. Finally, I want to thank Pascale Chavatte-Palmer and François Mallet, for taking the time to read my work and for accepting to be a part of the jury. I would, of course, also like to thank Thierry Heidmann for giving me the opportunity to work in his laboratory for the past four years, for being always available when I was in need of a guiding hand and present when I needed some prodding. Thank you for not only helping me to get these results, but also for the countless hours we spent to turn them into actual scientific articles. I am grateful to Anne Dupressoir and Gérard Pierron for spending time reading through my manuscript and their words of encouragement and advice. I also want to thank Christa Kuhn for her enthusiasm and encouragement, which played an important part in getting me to where I am today. Je souhaite remercier l’intégralité de l’unité 9196, en particulier Marie et Anne pour avoir partagé leurs savoirs, leurs conseils ainsi que leurs bureaux, Cécile Vernochet pour son aide inestimable, Seila pour son soutien administratif et émotionnel, Grégoire pour son optimisme inébranlable et infectieux, Caroline, Cécile Lemaître et Anthony pour les bon moments passés ensemble, et ce-dernier tout particulièrement pour nos nombreuses discussions (profondes et/ou inutiles) et son énergie aussi inépuisable qu’insupportable. Un grand merci également à Guil- laume, ex-membre de l’unité, pour m’avoir encadré pendant mes débuts et avoir toujours été disponible pour mes questions. J’aimerais remercier le gang des pasteuriens, premiers parmi eux mes ex-colocataires, Matthieu et Thomas, merci de m’avoir supporté pendant ces trois ans et quelques de vie com- mune au 20 avenue d’Ivry, qui s’est transformé en haut-lieu de la science, accueillant maints et maints congrès réunissant la fine fleur de la virologie et de l’immunologie parisienne. I would iii like to thank Oksana, one of the kindest and most generous persons I have had the pleasure to know and someone you can always count on. Merci à Simon l’invisible, ton rétro nous a apporté de nombreuses heures de bonheur. Merci à Vincent le sale cylon pour les côtes de bœuf et à Essia pour ses couscous, toujours aussi bons que son humeur. Merci aux deux Juju pour la bonne ambiance, et je vous souhaite bien du courage, à Juju pour supporter Matthieu et à Juju pour finir ta thèse. Merci aussi à Anaïs, Laura, Cécile, Alix, Marion et tous les autres qui ont rendu la vie parisienne tellement plus agréable. J’aimerais également remercier mes nouveaux ex-colocataires, Raphaël, Ferretti et Viking, pour m’avoir permis de garder l’appartement quelques mois de plus, puis pour m’avoir accueilli sur notre canapé qui, entretemps, était devenu le leur. On a somewhat stranger note I would like to thank all of the friends I have never met, but who have been there to play, talk and just hang out in the virtual parts of this world. Thank you to all sigmas, especially 48, Mori Kaal and koreamax, but also willemd, :goatdrügs:, Cal, and even Thuneral, Slap Chop and Koahi. A great thank you to the Aqueduct Acrobats who have been able to observe my slow and steady descent into placenta and retrovirus induced madness over the past few months, especially Topaz with his awful K/D, expensive haircuts and shared disgust of terrible book adaptations, mrcompson for our academia themed diversions and unmasking the terrible truth behind omics- conferences, and Malloot, everyone’s favorite tactical espionage action ground team leader and platforming liability. I also want to thank Norns and Tarth, our resident Americans, as well as Charisma and Waka. Ich möchte natürlich auch meiner Familie danken, meiner Mutter, meiner Schwester und meinen beiden Brüdern, von deren Hilfe ich immer ausgehen konnte und deren Gesellschaft und Vertrauen mein Leben bereichert haben. Ich danke meiner Oma und meinem Opa für ihre Ermutigungen und Liebe. J’aimerais remercier Patrice, qui a été pour moi pendant ces 25 dernières années le meilleur père que j’aurais pu souhaiter, et qui m’a poussé à être curieux et intéressé en espérant de peut-être, un jour, être aussi instruit et intéressant que lui. Ich möchte hier auch ganz besonders meiner Tante Jutta danken, die innerhalb von ein paar Tagen die ganze Einführung durchgelesen und korrigiert hat. Last but not least, I want to thank all of the people that I have shamefully forgotten to mention here, please forgive me for this oversight. I am still unsure of how I made it this far, but one thing I know: I would not have made it without you. The words above pale in comparison to what I owe you all, I have written and rewritten them many times and I know I’ll never be able to express the feeling appropriately, so all I can do is apologize and say it once more, thank you all so very much. iv Contents Acknowledgments iii Abbreviations xi I Introduction 1 1 Placenta 5 1.1 Challenges of viviparity ............................. 5 1.1.1 Maternofetal exchanges . 5 a. Nutrients 6 b. Oxygen 8 c. Waste 9 1.1.2 Placentation and the maternal immune system . 9 a. Major histocompatibility complex expression 9 b. Uterine natural killer cells 10 c. Adaptive immunity 11 1.1.3 Maternofetal communication . 11 a. Placental hormones 11 b. Extracellular vesicles 12 1.2 Mammalian eutherian placentas ........................ 13 1.2.1 Origin and evolution of the mammalian placenta . 13 a. Monophyletic origin 13 b. Placental diversity 13 1.2.2 Placental types by maternofetal interface . 16 a. Epitheliochorial placentation 18 b. Synepitheliochorial placentation 20 c. Endotheliochorial placentation 22 d. Hemochorial placentation 22 1.3 Viviparity in non-mammalian species ..................... 26 1.3.1 Prevalence of viviparity outside of mammals . 26 v a. Invertebrates 26 b. Non-mammalian vertebrates 27 1.3.2 Examples of non-mammalian placentas . 30 a. Non-vertebrates: the placenta of salps 30 b. Fish: placentation in sharks 31 c. Lizards: placentation in Mabuya and other Scincidae 32 2 Retroviruses 38 2.1 Exogenous Retroviruses ............................. 39 2.1.1 Genomic orgarnisation . 39 a. Non-coding features 39 b. Gag 41 c. Pro and Pol 42 d. Env 44 e. Regulatory and accessory proteins 44 2.1.2 Retroviral phylogeny . 48 a. Subfamily Orthoretrovirinae 48 b. Subfamily Spumaretrovirinae 51 2.1.3 Viral cycle . 53 a. Viral entry 53 b. Reverse transcription 53 c. Genome integration 57 d. Protein expression 60 e. Assembly and budding 61 f. Maturation 64 2.2 Endogenous Retroviruses ............................ 66 2.2.1 Endogenization of retroviral sequences . 67 a. Infection of germinal cells 67 b. Intra-genome spreading of ERVs 69 c. Groups of ERVs 71 2.2.2 Impact of retroviral integration on the host . 72 a. LTR integration can drive ectopic gene expression 72 b. Impact of retroviral gene acquisition on host physiology 76 2.2.3 Fate of ERVs and control by the host . 78 a. Epigenetic regulation of ERV expression 78 b. Accumulation of mutations and solo LTR formation 79 c. Exaptation and purifying selection 80 3 Syncytins 81 3.1 Retroviral envelopes ............................... 81 vi 3.1.1 Functional domains . 81 a. Domains of the SU 81 b. Domains of the TM 83 3.1.2 Env induced membrane fusion . 86 a. Implication of the cognate receptor in fusion 86 b. Model of viral fusion protein class I induced membrane fusion 89 3.2 Syncytins in mammals .............................. 90 3.2.1 Human syncytins: Syncytin-1 and Syncytin-2 . 90 a. Syncytin-1 92 b. Syncytin-2 94 3.2.2 Mouse syncytins: Syncytin-A and Syncytin-B . 95 a. Discovery 95 b. Expression in 2 segregated layers 95 c. Impact of Syncytin knockout on placental physiology 96 3.2.3 Other mammalian syncytins . 98 a. Rabbit: Syncytin-Ory1 98 b. Cat and dog: Syncytin-Car1 99 c. Old world monkey: EnvV2 99 d. Cattle and sheep: Syncytin-Rum1 and Fematrin-1 99 e.
Recommended publications
  • Reproductive Specializations in a Viviparous African Skink: Implications for Evolution and Biological Conservation Daniel G
    Trinity College Trinity College Digital Repository Faculty Scholarship 8-2010 Reproductive Specializations in a Viviparous African Skink: Implications for Evolution and Biological Conservation Daniel G. Blackburn Trinity College, [email protected] Alexander F. Flemming University of Stellenbosch Follow this and additional works at: http://digitalrepository.trincoll.edu/facpub Part of the Biology Commons Herpetological Conservation and Biology 5(2):263-270. Symposium: Reptile Reproduction REPRODUCTIVE SPECIALIZATIONS IN A VIVIPAROUS AFRICAN SKINK AND ITS IMPLICATIONS FOR EVOLUTION AND CONSERVATION 1 2 DANIEL G. BLACKBURN AND ALEXANDER F. FLEMMING 1Department of Biology and Electron Microscopy Facility, Trinity College, Hartford, Connecticut 06106, USA, e-mail: [email protected] 2Department of Botany and Zoology, University of Stellenbosch, Stellenbosch 7600, South Africa Abstract.—Recent research on the African scincid lizard, Trachylepis ivensi, has significantly expanded the range of known reproductive specializations in reptiles. This species is viviparous and exhibits characteristics previously thought to be confined to therian mammals. In most viviparous squamates, females ovulate large yolk-rich eggs that provide most of the nutrients for development. Typically, their placental components (fetal membranes and uterus) are relatively unspecialized, and similar to their oviparous counterparts. In T. ivensi, females ovulate tiny eggs and provide nutrients for embryonic development almost entirely by placental means. Early in gestation, embryonic tissues invade deeply into maternal tissues and establish an intimate “endotheliochorial” relationship with the maternal blood supply by means of a yolk sac placenta. The presence of such an invasive form of implantation in a squamate reptile is unprecedented and has significant functional and evolutionary implications. Discovery of the specializations of T.
    [Show full text]
  • The Herpetofauna of the Cubango, Cuito, and Lower Cuando River Catchments of South-Eastern Angola
    Official journal website: Amphibian & Reptile Conservation amphibian-reptile-conservation.org 10(2) [Special Section]: 6–36 (e126). The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola 1,2,*Werner Conradie, 2Roger Bills, and 1,3William R. Branch 1Port Elizabeth Museum (Bayworld), P.O. Box 13147, Humewood 6013, SOUTH AFRICA 2South African Institute for Aquatic Bio- diversity, P/Bag 1015, Grahamstown 6140, SOUTH AFRICA 3Research Associate, Department of Zoology, P O Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth 6031, SOUTH AFRICA Abstract.—Angola’s herpetofauna has been neglected for many years, but recent surveys have revealed unknown diversity and a consequent increase in the number of species recorded for the country. Most historical Angola surveys focused on the north-eastern and south-western parts of the country, with the south-east, now comprising the Kuando-Kubango Province, neglected. To address this gap a series of rapid biodiversity surveys of the upper Cubango-Okavango basin were conducted from 2012‒2015. This report presents the results of these surveys, together with a herpetological checklist of current and historical records for the Angolan drainage of the Cubango, Cuito, and Cuando Rivers. In summary 111 species are known from the region, comprising 38 snakes, 32 lizards, five chelonians, a single crocodile and 34 amphibians. The Cubango is the most western catchment and has the greatest herpetofaunal diversity (54 species). This is a reflection of both its easier access, and thus greatest number of historical records, and also the greater habitat and topographical diversity associated with the rocky headwaters.
    [Show full text]
  • Arrival and Diversification of Mabuyine Skinks (Squamata: Scincidae) in the Neotropics Based on a Fossil-Calibrated Timetree
    Arrival and diversification of mabuyine skinks (Squamata: Scincidae) in the Neotropics based on a fossil-calibrated timetree Anieli Guirro Pereira and Carlos G. Schrago Department of Genetics, Federal University of Rio de Janeiro, Rio de Janeiro, RJ, Brazil ABSTRACT Background. The evolution of South American Mabuyinae skinks holds significant biogeographic interest because its sister lineage is distributed across the African continent and adjacent islands. Moreover, at least one insular species, Trachylepis atlantica, has independently reached the New World through transoceanic dispersal. To clarify the evolutionary history of both Neotropical lineages, this study aimed to infer an updated timescale using the largest species and gene sampling dataset ever assembled for this group. By extending the analysis to the Scincidae family, we could employ fossil information to estimate mabuyinae divergence times and carried out a formal statistical biogeography analysis. To unveil macroevolutionary patterns, we also inferred diversification rates for this lineage and evaluated whether the colonization of South American continent significantly altered the mode of Mabuyinae evolution. Methods. A time-calibrated phylogeny was inferred under the Bayesian framework employing fossil information. This timetree was used to (i) evaluate the historical biogeography of mabuiyines using the statistical approach implemented in Bio- GeoBEARS; (ii) estimate macroevolutionary diversification rates of the South American Mabuyinae lineages and the patterns of evolution of selected traits, namely, the mode of reproduction, body mass and snout–vent length; (iii) test the hypothesis of differential macroevolutionary patterns in South American lineages in BAMM and GeoSSE; and Submitted 21 November 2016 (iv) re-evaluate the ancestral state of the mode of reproduction of mabuyines.
    [Show full text]
  • Reproductionreview
    REPRODUCTIONREVIEW The evolution of viviparity: molecular and genomic data from squamate reptiles advance understanding of live birth in amniotes James U Van Dyke, Matthew C Brandley and Michael B Thompson School of Biological Sciences, University of Sydney, A08 Heydon-Laurence Building, Sydney, New South Wales 2006, Australia Correspondence should be addressed to J U Van Dyke; Email: [email protected] Abstract Squamate reptiles (lizards and snakes) are an ideal model system for testing hypotheses regarding the evolution of viviparity (live birth) in amniote vertebrates. Viviparity has evolved over 100 times in squamates, resulting in major changes in reproductive physiology. At a minimum, all viviparous squamates exhibit placentae formed by the appositions of maternal and embryonic tissues, which are homologous in origin with the tissues that form the placenta in therian mammals. These placentae facilitate adhesion of the conceptus to the uterus as well as exchange of oxygen, carbon dioxide, water, sodium, and calcium. However, most viviparous squamates continue to rely on yolk for nearly all of their organic nutrition. In contrast, some species, which rely on the placenta for at least a portion of organic nutrition, exhibit complex placental specializations associated with the transport of amino acids and fatty acids. Some viviparous squamates also exhibit reduced immunocompetence during pregnancy, which could be the result of immunosuppression to protect developing embryos. Recent molecular studies using both candidate-gene and next-generation sequencing approaches have suggested that at least some of the genes and gene families underlying these phenomena play similar roles in the uterus and placenta of viviparous mammals and squamates.
    [Show full text]
  • An Endogenous Retroviral Envelope Syncytin and Its Cognate Receptor Identified in the Viviparous Placental Mabuya Lizard
    An endogenous retroviral envelope syncytin and its PNAS PLUS cognate receptor identified in the viviparous placental Mabuya lizard Guillaume Cornelisa,b,1,2, Mathis Funka,b,1, Cécile Vernocheta,b, Francisca Lealc,3, Oscar Alejandro Tarazonac,4, Guillaume Meuriced, Odile Heidmanna,b, Anne Dupressoira,b, Aurélien Mirallese, Martha Patricia Ramirez-Pinillac, and Thierry Heidmanna,b,5 SEE COMMENTARY aUnité Physiologie et Pathologie Moléculaires des Rétrovirus Endogènes et Infectieux, CNRS UMR 9196, Gustave Roussy, Villejuif, F-94805, France; bUMR 9196, Université Paris-Sud, Orsay, F-91405, France; cLaboratorio de Biologia Reproductiva de Vertebrados, Escuela de Biologia, Universidad Industrial de Santander, 680002 Bucaramanga, Colombia; dPlateforme de Bioinformatique, INSERM US23/CNRS UMS3655, Gustave Roussy, Villejuif, F-94805, France; and eInstitut de Systématique, Evolution, Biodiversité, Muséum National d’Histoire Naturelle, CNRS UPMC EPHE, Sorbonne Universités, Paris, F-75005, France Edited by R. Michael Roberts, University of Missouri-Columbia, Columbia, MO, and approved October 26, 2017 (received for review August 23, 2017) Syncytins are envelope genes from endogenous retroviruses that Remarkably, placental structures are not restricted to mamma- have been captured during evolution for a function in placentation. lian species. Placentation emerged independently and in a sto- They have been found in all placental mammals in which they have chastic manner in several groups of vertebrates, with the noticeable been searched, including marsupials. Placental structures are not exception of birds (reviewed in refs. 16 and 17). In particular, restricted to mammals but also emerged in some other vertebrates, complex placentas have been described in some South American most frequently in lizards, such as the viviparous Mabuya Scincidae.
    [Show full text]
  • Reproductive Specializations in a Viviparous African Skink: Implications for Evolution and Biological Conservation
    Trinity College Trinity College Digital Repository Faculty Scholarship 8-2010 Reproductive Specializations in a Viviparous African Skink: Implications for Evolution and Biological Conservation Daniel G. Blackburn Trinity College, [email protected] Alexander F. Flemming University of Stellenbosch Follow this and additional works at: https://digitalrepository.trincoll.edu/facpub Part of the Biology Commons Herpetological Conservation and Biology 5(2):263-270. Symposium: Reptile Reproduction REPRODUCTIVE SPECIALIZATIONS IN A VIVIPAROUS AFRICAN SKINK AND ITS IMPLICATIONS FOR EVOLUTION AND CONSERVATION 1 2 DANIEL G. BLACKBURN AND ALEXANDER F. FLEMMING 1Department of Biology and Electron Microscopy Facility, Trinity College, Hartford, Connecticut 06106, USA, e-mail: [email protected] 2Department of Botany and Zoology, University of Stellenbosch, Stellenbosch 7600, South Africa Abstract.—Recent research on the African scincid lizard, Trachylepis ivensi, has significantly expanded the range of known reproductive specializations in reptiles. This species is viviparous and exhibits characteristics previously thought to be confined to therian mammals. In most viviparous squamates, females ovulate large yolk-rich eggs that provide most of the nutrients for development. Typically, their placental components (fetal membranes and uterus) are relatively unspecialized, and similar to their oviparous counterparts. In T. ivensi, females ovulate tiny eggs and provide nutrients for embryonic development almost entirely by placental means. Early in gestation, embryonic tissues invade deeply into maternal tissues and establish an intimate “endotheliochorial” relationship with the maternal blood supply by means of a yolk sac placenta. The presence of such an invasive form of implantation in a squamate reptile is unprecedented and has significant functional and evolutionary implications. Discovery of the specializations of T.
    [Show full text]
  • Patterns of Species Richness, Endemism and Environmental Gradients of African Reptiles
    Journal of Biogeography (J. Biogeogr.) (2016) ORIGINAL Patterns of species richness, endemism ARTICLE and environmental gradients of African reptiles Amir Lewin1*, Anat Feldman1, Aaron M. Bauer2, Jonathan Belmaker1, Donald G. Broadley3†, Laurent Chirio4, Yuval Itescu1, Matthew LeBreton5, Erez Maza1, Danny Meirte6, Zoltan T. Nagy7, Maria Novosolov1, Uri Roll8, 1 9 1 1 Oliver Tallowin , Jean-Francßois Trape , Enav Vidan and Shai Meiri 1Department of Zoology, Tel Aviv University, ABSTRACT 6997801 Tel Aviv, Israel, 2Department of Aim To map and assess the richness patterns of reptiles (and included groups: Biology, Villanova University, Villanova PA 3 amphisbaenians, crocodiles, lizards, snakes and turtles) in Africa, quantify the 19085, USA, Natural History Museum of Zimbabwe, PO Box 240, Bulawayo, overlap in species richness of reptiles (and included groups) with the other ter- Zimbabwe, 4Museum National d’Histoire restrial vertebrate classes, investigate the environmental correlates underlying Naturelle, Department Systematique et these patterns, and evaluate the role of range size on richness patterns. Evolution (Reptiles), ISYEB (Institut Location Africa. Systematique, Evolution, Biodiversite, UMR 7205 CNRS/EPHE/MNHN), Paris, France, Methods We assembled a data set of distributions of all African reptile spe- 5Mosaic, (Environment, Health, Data, cies. We tested the spatial congruence of reptile richness with that of amphib- Technology), BP 35322 Yaounde, Cameroon, ians, birds and mammals. We further tested the relative importance of 6Department of African Biology, Royal temperature, precipitation, elevation range and net primary productivity for Museum for Central Africa, 3080 Tervuren, species richness over two spatial scales (ecoregions and 1° grids). We arranged Belgium, 7Royal Belgian Institute of Natural reptile and vertebrate groups into range-size quartiles in order to evaluate the Sciences, OD Taxonomy and Phylogeny, role of range size in producing richness patterns.
    [Show full text]
  • Reptiles & Amphibians
    AWF FOUR CORNERS TBNRM PROJECT : REVIEWS OF EXISTING BIODIVERSITY INFORMATION i Published for The African Wildlife Foundation's FOUR CORNERS TBNRM PROJECT by THE ZAMBEZI SOCIETY and THE BIODIVERSITY FOUNDATION FOR AFRICA 2004 PARTNERS IN BIODIVERSITY The Zambezi Society The Biodiversity Foundation for Africa P O Box HG774 P O Box FM730 Highlands Famona Harare Bulawayo Zimbabwe Zimbabwe Tel: +263 4 747002-5 E-mail: [email protected] E-mail: [email protected] Website: www.biodiversityfoundation.org Website : www.zamsoc.org The Zambezi Society and The Biodiversity Foundation for Africa are working as partners within the African Wildlife Foundation's Four Corners TBNRM project. The Biodiversity Foundation for Africa is responsible for acquiring technical information on the biodiversity of the project area. The Zambezi Society will be interpreting this information into user-friendly formats for stakeholders in the Four Corners area, and then disseminating it to these stakeholders. THE BIODIVERSITY FOUNDATION FOR AFRICA (BFA is a non-profit making Trust, formed in Bulawayo in 1992 by a group of concerned scientists and environmentalists. Individual BFA members have expertise in biological groups including plants, vegetation, mammals, birds, reptiles, fish, insects, aquatic invertebrates and ecosystems. The major objective of the BFA is to undertake biological research into the biodiversity of sub-Saharan Africa, and to make the resulting information more accessible. Towards this end it provides technical, ecological and biosystematic expertise. THE ZAMBEZI SOCIETY was established in 1982. Its goals include the conservation of biological diversity and wilderness in the Zambezi Basin through the application of sustainable, scientifically sound natural resource management strategies.
    [Show full text]
  • Guide to the Reptiles and Amphibians of the Maasai Mara
    Male Boomslang Dispholidus typus N OLARE MOTOROGI CONSERVANCY OLARRO CONSERVANCY Copyright: Stephen Spawls 2017. Further Copies of this book can be ob- tained from Gamewatchers Safaris, P.O. Box 388 -00621, Village Market, Nairobi, Kenya, (United Nations Crescent; near Village Market, Nairobi); Email [email protected] or from Stephen Spawls, 7 Crostwick Lane, Spixworth, Norwich NR10 3PE, U.K.; Email [email protected] Cover picture: Mara Black-necked Spitting Cobra by Anthony Childs Designed by: Norfolk Educational Services, City College Norwich A Guide to Amphibians and Reptiles of the Maasai Mara Ecosystem By Printed by Stephen Spawls Norwich City College Print Room Contents Introduction 2 Foreword by Jake Grieves-Cook, 3 Chairman of Gamewatchers Safaris & Porini Camps Why conservation and the conservancies are important 4 Observing and Collecting Reptiles and Amphibians 5 Safety and Reptiles/Amphibians 7 Snakebite 8 The Maasai-Mara Ecosystem and its reptile 9 and amphibian fauna The Reptiles and Amphibians of the Maasai Mara ecosystem 10 Some species that might be found in the Maasai Mara 34 but are as yet unrecorded Acknowledgements 38 Further Resources 39 Introduction “The natural resources of this country – its wildlife which offers such an attraction to beautiful places…the mighty forests which guard the water catchment areas so vital to the survival of man and beast – are a priceless heritage for the future. The government of Kenya…pledges itself to conserve them for posterity with all the means at its disposal.” Mzee Jomo Kenyatta, First President of the Republic of Kenya (inscription at the entrance of Nairobi National Park 1964) As far as I am aware, this book is the first guide to the reptiles and amphibians of any large conservation area in Kenya.
    [Show full text]
  • DNA Barcoding Assessment of Genetic Variation in Two Widespread Skinks from Madagascar, Trachylepis Elegans and T
    Zootaxa 3755 (5): 477–484 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3755.5.7 http://zoobank.org/urn:lsid:zoobank.org:pub:039C3026-D3B9-47B2-8340-6B916C636665 DNA barcoding assessment of genetic variation in two widespread skinks from Madagascar, Trachylepis elegans and T. gravenhorstii (Squamata: Scincidae) MIGUEL VENCES1,6*, ALEXANDRA LIMA2,3*, AURÉLIEN MIRALLES4 & FRANK GLAW5 1Zoological Institute, Technische Universität Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany 2Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO), Campus Agrário de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal 3Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, R. Campo Alegre s/n, 4169-007 Porto, Portugal 4CNRS-UMR5175 Centre d'Ecologie Fonctionnelle et Evolutive, 1919 route de Mende, 34293 Montpellier cedex 5, France 5Zoologische Staatssammlung München, Münchhausenstr. 21, 81247 München, Germany 6Corresponding author. E-mail: [email protected] *These authors contributed equally to the paper Abstract Trachylepis elegans and T. gravenhorstii are two of the most widespread reptiles in Madagascar, inhabiting a wide variety of habitats. Previous studies have indicated a considerable mitochondrial DNA (mtDNA) variation within these species, but the geographic distribution of the major haplotype lineages is poorly known. Herein we analyse the phylogeography of these lizards based on 107 sequences of the mitochondrial cytochrome oxidase subunit I gene, 101 of which newly de- termined. As in previous mtDNA assessments, T. elegans and T. gravenhorstii were not reciprocally monophyletic, al- though recent analyses including nuclear markers indicated their probable monophyly, respectively.
    [Show full text]
  • An Endogenous Retroviral Envelope Syncytin and Its Cognate Receptor
    An endogenous retroviral envelope syncytin and its PNAS PLUS cognate receptor identified in the viviparous placental Mabuya lizard Guillaume Cornelisa,b,1,2, Mathis Funka,b,1, Cécile Vernocheta,b, Francisca Lealc,3, Oscar Alejandro Tarazonac,4, Guillaume Meuriced, Odile Heidmanna,b, Anne Dupressoira,b, Aurélien Mirallese, Martha Patricia Ramirez-Pinillac, and Thierry Heidmanna,b,5 aUnité Physiologie et Pathologie Moléculaires des Rétrovirus Endogènes et Infectieux, CNRS UMR 9196, Gustave Roussy, Villejuif, F-94805, France; bUMR 9196, Université Paris-Sud, Orsay, F-91405, France; cLaboratorio de Biologia Reproductiva de Vertebrados, Escuela de Biologia, Universidad Industrial de Santander, 680002 Bucaramanga, Colombia; dPlateforme de Bioinformatique, INSERM US23/CNRS UMS3655, Gustave Roussy, Villejuif, F-94805, France; and eInstitut de Systématique, Evolution, Biodiversité, Muséum National d’Histoire Naturelle, CNRS UPMC EPHE, Sorbonne Universités, Paris, F-75005, France Edited by R. Michael Roberts, University of Missouri-Columbia, Columbia, MO, and approved October 26, 2017 (received for review August 23, 2017) Syncytins are envelope genes from endogenous retroviruses that Remarkably, placental structures are not restricted to mamma- have been captured during evolution for a function in placentation. lian species. Placentation emerged independently and in a sto- They have been found in all placental mammals in which they have chastic manner in several groups of vertebrates, with the noticeable been searched, including marsupials. Placental structures are not exception of birds (reviewed in refs. 16 and 17). In particular, restricted to mammals but also emerged in some other vertebrates, complex placentas have been described in some South American most frequently in lizards, such as the viviparous Mabuya Scincidae.
    [Show full text]
  • Here Additional SEcies EEcted in Undavala Ere Recorded
    © University of Hamburg 2018 All rights reserved Klaus Hess Publishers Göttingen & Windhoek www.k-hess-verlag.de ISBN: 978-3-933117-95-3 (Germany), 978-99916-57-43-1 (Namibia) Language editing: Will Simonson (Cambridge), and Proofreading Pal Translation of abstracts to Portuguese: Ana Filipa Guerra Silva Gomes da Piedade Page desing & layout: Marit Arnold, Klaus A. Hess, Ria Henning-Lohmann Cover photographs: front: Thunderstorm approaching a village on the Angolan Central Plateau (Rasmus Revermann) back: Fire in the miombo woodlands, Zambia (David Parduhn) Cover Design: Ria Henning-Lohmann ISSN 1613-9801 Printed in Germany Suggestion for citations: Volume: Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N. (eds.) (2018) Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions. Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek. Articles (example): Archer, E., Engelbrecht, F., Hänsler, A., Landman, W., Tadross, M. & Helmschrot, J. (2018) Seasonal prediction and regional climate projections for southern Africa. In: Climate change and adaptive land management in southern Africa – assessments, changes, challenges, and solutions (ed. by Revermann, R., Krewenka, K.M., Schmiedel, U., Olwoch, J.M., Helmschrot, J. & Jürgens, N.), pp. 14–21, Biodiversity & Ecology, 6, Klaus Hess Publishers, Göttingen & Windhoek. Corrections brought to our attention will be published at the following location: http://www.biodiversity-plants.de/biodivers_ecol/biodivers_ecol.php
    [Show full text]