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Phrynobatrachus Steindachneri (Phrynobatrachidae)

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Phrynobatrachus Steindachneri (Phrynobatrachidae) PŘÍRODOVĚDECKÁ FAKULTA Morfologická diverzifikace afrických bezocasých obojživelníků na úrovni různých taxonomických kategorií Diplomová práce Tadeáš Nečas Vedoucí práce: RNDr. Václav Gvoždík Ph.D. Ústav botaniky a zoologie Brno 2019 Bibliografický záznam Autor: Tadeáš Nečas Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Morfologická diverzifikace afrických bezocasých obojživelníků na úrovni různých taxonomických kate- gorií Studijní program: Zoologie Studijní obor: Ekologická a evoluční biologie Vedoucí práce: RNDr. Václav Gvoždík Ph.D. Akademický rok: 2018/2019 Počet stran: 140 Klíčová slova: Afrika; afroskokan; Amphibia; Anura; Hyperolius; mor- fologie; Kamerun; Kamerunské hory; Kongo; obojživel- nící; Phrynobatrachus; rákosnička; systematika; taxono- mie; žáby. Bibliographic Entry Author Tadeáš Nečas Faculty of Science, Masaryk University Department of Botany And Zoology Title of Thesis: Morphological diversification of African anurans from different taxonomic ranks Degree programme: Zoology Field of Study: Ecological and Evolutionary Biology Supervisor: RNDr. Václav Gvoždík Ph.D. Academic Year: 2018/2019 Number of Pages: 140 Keywords: Africa; Amphibia; amphibians; Anura; Cameroon; Cam- eroon Volcanic Line; Congo; frogs; Hyperolius; mor- phology; Phrynobatrachus; puddle frog; reed frog; sys- tematics; taxonomy. Abstrakt Skrytá diverzita žab (Amphibia: Anura) je již řadu desetiletí předmětem studia mnoha evolučních biologů a taxonomů, jejichž práce se ve velkém zjednodušila se začátkem používání molekulárních metod. Tato práce je zaměřena na dva nepříbuzné taxony žab střední Afriky, které spojuje právě jejich kryptický charakter. Hlavní náplní této práce je zhodnocení jejich morfologických a případně i fylogenetických vztahů v rámci druhového komplexu, či rodu. Prvním studovaným taxonem je druhový kom- plex Phrynobatrachus steindachneri (Phrynobatrachidae). Tento druhový komplex za- hrnující morfologicky velice podobné endemity Kamerunských hor v současnosti za- hrnuje tři druhy popsané a jeden druh kandidátní, jehož popis se připravuje. Cílem první části je za pomoci mnohorozměrných statistických metod zhodnotit morfologic- kou variabilitu, či konzervativnost, v rámci tohoto druhového komplexu. Analýzy 14 morfometrických parametrů odhalily malou, ale signifikantní variabilitu odlišující zejména nejsevernější populace/taxon. Druhá část práce se věnuje druhu Hyperolius robustus, který je v současnosti znám pouze z několika málo jedinců a lokalit v De- mokratické republice Kongo. Jeho zařazení do rodu Hyperolius na základě celkového vzhledu vnější morfologie se zdá být jednoznačné, ale fylogenetické analýzy mito- chondriální a jaderné DNA poukazují na vazbu k rodům Morerella a Cryptothylax. Detailní zhodnocení vnější morfologie za využití mnohorozměrných statistických me- tod a vnitřní morfologie (osteologie) za využití μCT podpořilo výsledky fylogenetic- kých analýz a odlišnost tohoto taxonu od rodu Hyperolius. Práce na těchto dvou pří- kladech ukazuje dva odlišné evoluční modely kryptických druhů žab. První předsta- vuje případ morfologické konzervativnosti, pravděpodobně vlivem konzervativnosti ekologické niky, zatímco druhý představuje případ konvergence. Abstract The cryptic diversity of the order Anura (Amphibia) has been in the focus of evo- lutionary biologists and taxonomists since the expansion of the use of molecular meth- ods in biological sciences. The main objective of this study is to evaluate morpholog- ically and eventually phylogenetically two representatives of Central African frogs that have in common their cryptic character. Though, on two different systematic lev- els. The first studied model is the Phrynobatrachus steindachneri species complex (Phrynobatrachidae). This morphologically very conservative complex contains three nominal species of medium-sized frogs, endemics of the Cameroon Volcanic Line. A description of a fourth species is under a preparation. The main objective of the first part was to assess the morphological variation within this species complex using mul- tivariate statistical analyses. The multivariate analyses of 14 morphometric parameters revealed limited variation, although significantly distinguishing the northern popula- tion/taxon. The second part of this study focused on Hyperolius robustus, an endemic species of the Democratic Republic of the Congo known from only a few specimens and localities. Based on the superficial characters, Hyperolius robustus resembles spe- cies from the genus Hyperolius. On the contrary, phylogenetic analyses of mitochon- drial and nuclear DNA place Hyperolius robustus in a relationship with the genera Morerella and Cryptothylax, which form a sister clade to the genus Hyperolius. De- tailed assessment of the external morphology, using multivariate statistical analyses, and the internal morphology (osteology), using the μCT, supported results of the phy- logenetic analyses and differentiate H. robustus from the genus Hyperolius. This study shows two different evolutionary models of anuran cryptic species. The first represents a case of the morphological conservatism, probably caused by the conservation of eco- logical niche. While the second represents a case of the convergence. Masarykova univerzita Přírodovědecká fakulta ZADÁNÍ DIPLOMOVÉ PRÁCE Student: Bc. Tadeáš Nečas Studijní program: Ekologická a evoluční biologie Studijní obor: Zoologie Ředitel Ústavu botaniky a zoologie PřF MU Vám ve smyslu Studijního a zku- šebního řádu MU určuje diplomovou práci s tématem: Morfologická diverzifikace afrických bezocasých obojživelníků na úrovni různých taxonomických kategorií Morphological diversification of African anurans from different taxonomic ranks Oficiální zadání: Stupeň morfologické diverzifikace se liší na různých taxonomických úrovních jak vlivem genotypu, tak vnitrodruhová variabilita může být často ovlivněna vnějšími faktory. V rámci projektu budou studovány vybrané taxony afrických oboj- živelníků z různých taxonomických kategorií s cílem zjistit rozsah morfologické di- verzifikace vs. konzervativnosti na vnitrodruhové/mezipopulační úrovni (popř. u blízce příbuzných druhů) a na mezidruhové úrovni v rámci rodu (popř. mezi blízce příbuznými rody). Morfologie je tradiční klasifikační nástroj v systematice a taxono- mii, nicméně ne vždy je morfologie hodnocena v kontextu možné fenotypové variabi- lity či plasticity. Cílem práce bude (i) morfologicky zhodnotit variabilitu vnitrodruho- vou (či blízce příbuzných druhů), (ii) zhodnotit morfologickou diverzifikaci odlišující druhy geneticky více vzdálené (či blízké rody), a (iii) provést porovnání ve fylogene- tickém a taxonomickém kontextu. Jako modelové skupiny budou sloužit vybrané ta- xony bezocasých obojživelníků z rodu Phrynobatrachus a Hyperolius. Literatura: Gvoždík V., Moravec J., Kratochvíl L. (2008). Geographic morphological variation in parapatric Western Palearctic tree frogs, Hyla arborea and Hyla savignyi: are rela- ted species similarly affected by climatic conditions? Biological Journal of the Lin- nean Society 95: 539–556. Laurent R. F. (1979). Description de deux Hyperolius nouveaux du Sankuru (Zaïre) (Amphibia, Hyperoliidae). Revue de Zoologie et de Botanique Africaines 93: 779– 791. Rödel M.-O., Kosuch J., Grafe T. U., Boistel R., Assemian N. G. E., Kouamé N. G. G., Tohé B., Gourène G., Perret J.-L., Henle K., Taffoureau P., Pollet N., Veith M. (2009). A new tree-frog genus and species from Ivory Coast, West Africa (Amphi- bia: Anura: Hyperoliidae). Zootaxa 2044: 23–45. Zimkus B. M. & Gvoždík V. (2013). Sky Islands of the Cameroon Volcanic Line: a diversification hotspot for puddle frogs (Phrynobatrachidae: Phrynobatrachus). Zoo- logical Scripta 42: 591–611. Jazyk závěrečné práce: angličtina Vedoucí diplomové práce: RNDr. Václav Gvoždík, PhD. Podpis vedoucího práce: Konzultant: Mgr. et Mgr. Josef Bryja, Ph.D. Vedoucí pracovní skupiny: doc. Mgr. Tomáš Bartonička, PhD. Podpis vedoucího pracovní skupiny: Datum zadání diplomové práce: 18. 2. 2019 V Brně dne prof. RNDr. Milan Chytrý, Ph.D ředitel Ústavu botaniky a zoologie Zadání diplomové práce převzal dne: Podpis studenta Poděkování Rád bych poděkoval rodině za podporu a trpělivost, kterou mi věnovali během mého studia. Dále bych rád poděkoval panu doktorovi Václavu Gvoždíkovi za věcné připomínky a rady při přípravě této práce, a především za možnost pracovat pod jeho vedením na této zajímavé problematice. Současně bych rád poděkoval kolegům panu magistru Mateji Dolinay a paní magistře Alexandře Hánové za rady a pomoc při labo- ratorní práci, a panu docentovi Michalu Vopálenskému z Laboratoře rentgenové radi- oskopie Ústavu teoretické a aplikované mechaniky AV ČR v Telči, s jehož pomocí probíhalo μCT snímkování. Prohlášení Prohlašuji, že jsem svoji diplomovou práci vypracoval samostatně s využitím in- formačních zdrojů, které jsou v práci citovány. Brno 23. dubna 2019 ……………………………… Jméno Příjmení Obsah 1 Introduction ............................................................................................................. 11 1.1 Objectives ......................................................................................................... 11 1.2 Cryptic species ................................................................................................. 11 1.3 Genus Phrynobatrachus – the first objective ................................................... 12 1.3.1 Systematic position of the genus Phrynobatrachus .................................. 12 1.3.2 Relationships within Phrynobatrachus
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