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Diverzita a Fylogenetické Vztahy Bělozubek Rodu Crocidura V Etiopii

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Diverzita a Fylogenetické Vztahy Bělozubek Rodu Crocidura V Etiopii MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Diverzita a fylogenetické vztahy bělozubek rodu Crocidura v Etiopii Bakalářská práce Brno 2014 Jana Táborská Vedoucí bakalářské práce: doc. Mgr. et. Mgr. Josef Bryja, Ph.D. Konzultant: Mgr. Tatiana Aghová Bibliografický záznam Autorka: Bc. Jana Táborská Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Diverzita a fylogenetické vztahy rodu Crocidura v Etiopii Studijní program: Ekologická a evoluční biologie Studijní obor: Zoologie Vedoucí práce: Doc. Mgr. et. Mgr. Josef Bryja, Ph.D. Konzultant: Mgr. Tatiana Aghová Akademický rok: 2013/2014 Počet stran: 60 Klíčová slova: Crocidura, bělozubky, genetická struktura, Soricidae, Etiopie Bibliographic Entry Author: Bc. Jana Táborská Title of Thesis: Diversity and phylogenetic relationships of Crocidura shrews in Ethiopia Degree programme: Ecological and Evolutionary Biology Field of Study: Zoology Supervisor: Doc. Mgr. et. Mgr. Josef Bryja, Ph.D. Co-supervisor: Mgr. Tatiana Aghová Academic Year: 2013/2014 Number of Pages: 60 Keywords: Crocidura, white-toothed shrews, genetic structure, Soricidae, Ethiopia Abstrakt Etiopie se vyznačuje vysokou biodiverzitou a unikátní faunou. Rod Crocidura tvoří nejvíce diverzifikovanou skupinu afrických hmyzoţravců, podle starších prací se na území Etiopie vyskytuje celkem 9 endemických druhů a 20 druhů široce rozšířených na africkém kontinentě. Cílem této práce bylo provést důkladnou literární rešerši týkající se rozšíření, taxonomie a fylogeneze rodu Crocidura v Etiopii a provést genetickou a morfologickou analýzu nově nasbíraného materiálu tohoto rodu. Bylo analyzováno celkem 36 jedinců ze 12 lokalit pokrývajících téměř celé území Etiopie s výjimkou její jihovýchodní části. Pro genetickou analýzu byly pouţity celé sekvence mitochondriálního genu pro cytochrom b (1140 bp). Na základě analýzy byly rozlišeny dvě velké skupiny uvnitř rodu Crocidura, v práci nazvány jako „etiopské endemické druhy“ a „široce rozšířené druhy“, jeţ se liší i preferencí různých nadmořských výšek. V analyzovaném materiálu se vyskytovali zástupci 10 druhů, z endemických druhů C. baileyi, C. lucina, C. glassi, C. thalia, Crocidura sp. A a Crocidura sp. B, z široce rozšířených druhů pak C. parvipes, C. olivieri, Crocidura sp. C a Crocidura sp. D. K jiţ výše zmíněným 9 etiopským endemitům genetická analýza odhalila dva druhy, Crocidura sp. A a Crocidura sp. B, které dosud nebyly blíţe popsány. V rozporu s jinými studiemi si jsou analyzovaní jedinci C. glassi a C. thalia velmi podobní jak na úrovni mitochondriálního genomu, tak v morfologických znacích a habitatech, jeţ obývají. Genetická analýza také odhalila sesterskou skupinu k C. olivieri, v práci nazvanou Crocidura sp. D. Dále byly v práci stanoveny základní morfometrické údaje u geneticky analyzovaných jedinců. Abstract Ethiopia is characterized by a high biodiversity and unique fauna. The genus Crocidura forms the most diversified group of the African insectivores, according to older studies there are 9 endemic species in Ethiopia and 20 widespread species on the African continent. The aim of this study was to carry out detailed research related to the distribution, taxonomy and phylogeny of the genus Crocidura in Ethiopia and to classify the newly collected specimens of this genus by the genetic and morphological analyses. A total of 36 samples from 12 localities, which cover nearly the entire region of Ethiopia, apart from its south-western area, was analyzed. For the genetical analyses complete nucleotide sequence of the mitochondrial gene of cytochrome b (1140 bp) was used. Based on this analysis two large groups were distinguished inside the genus Crocidura, in this study called "Ethiopian endemic species" and "widespread species", which differ also in the preference of different altitude. Representatives of 10 species were found in the analyzed material, from the Ethiopian endemic species C. baileyi, C. lucina, C. glassi, C. thalia, Crocidura sp. A and Crocidura sp. B, from the widespread species then C. parvipes, C. olivieri, Crocidura sp. C and Crocidura sp. D. The genetic analysis has discovered two new species to those 9 Ethiopian endemic species mentioned above, Crocidura sp. A and Crocidura sp. B, which have not been specified to this date. In contradiction to other studies, the analyzed specimens of C. glassi and C. thalia are very similar in the level of the mitochondrial genom, as well as the morphological features and habitats which they occupy. The genetic analysis has also discovered a sister group to C. olivieri, in this study called "Crocidura sp. D". Also the basic morfometric data concerning the genetically analyzed specimens were specified. Poděkování Chtěla bych velmi poděkovat svému vedoucímu Josefu Bryjovi a konzultantce Tatianě Aghové za moţnost věnovat se tomuto zajímavému tématu, ale především za jejich ochotu, veškerou pomoc, trpělivost a přátelský přístup. Tatianě bych chtěla navíc poděkovat za pomoc a zaučení jak do laboratorní práce, tak do všech pouţitých programů pro vyhodnocování a zpracování dat. Dále bych chtěla poděkovat Yonasi Meheretu za poskytnuté sekvence, a také virtuální organizaci METACENTRUM (https://metavo.metacentrum.cz/cs/about/index.html), na jejímţ vzdáleném počítači probíhaly fylogenetické analýzy. Tato bakalářská práce byla vypracována na Ústavu biologie obratlovců AV ČR, v. v. i., detašované pracoviště Studenec a byla podpořena projektem Grantové agentury ČR č. P506/10/0983: Srovnávací fylogeografie jihovýchodní Afriky (region Zambezi) na modelu drobných savců. Prohlášení Prohlašuji, ţe jsem svoji bakalářskou práci vypracovala samostatně s vyuţitím informačních zdrojů, které jsou v práci citovány. Souhlasím s uloţením této bakalářské práce v knihovně Ústavu botaniky a zoologie PřF MU v Brně, případně v jiné knihovně MU, s jejím veřejným půjčováním a vyuţitím pro vědecké, vzdělávací nebo jiné veřejně prospěšné účely, a to za předpokladu, ţe převzaté informace budou řádně citovány a nebudou vyuţívány komerčně. Brno 30. dubna 2014 …………………………... Jana Táborská OBSAH 1. ÚVOD ..................................................................................................................... 11 1.1. Formování bioty současné Afriky ................................................................... 11 1.2. Etiopie a její biogeografická struktura ............................................................ 11 1.3. Bělozubky rodu Crocidura v Africe ............................................................... 12 1.4. Cíle práce ........................................................................................................ 14 1.5. Druhy bělozubek rodu Crocidura v Etiopii .................................................... 15 1.5.1. Endemické druhy .................................................................................... 15 1.5.2. Široce rozšířené druhy ............................................................................ 21 2. MATERIÁL A METODY ...................................................................................... 33 2.1. Materiál ........................................................................................................... 33 2.2. Laboratorní analýzy ........................................................................................ 33 2.3. Fylogenetické analýzy .................................................................................... 34 3. VÝSLEDKY ........................................................................................................... 36 3.1. Fylogenetická analýza ..................................................................................... 36 3.2. Základní morfologická analýza ...................................................................... 38 3.3. Geografické rozšíření ...................................................................................... 40 4. DISKUZE ............................................................................................................... 41 4.1. Mezidruhové vztahy u etiopských taxonů rodu Crocidura ............................ 41 4.2. Morfologie a ekologické preference identifikovaných druhů ......................... 42 5. POUŢITÁ LITERATURA ..................................................................................... 46 6. SEZNAM PŘÍLOH ................................................................................................. 54 7. PŘÍLOHY ............................................................................................................... 55 10 1. ÚVOD 1.1. Formování bioty současné Afriky Africký kontinent patří z biologického hlediska mezi nejméně probádané kontinenty na světě. Vyznačuje se vysokou druhovou biodiverzitou, která se ve většině případů dle fosilních nálezů formovala v období Plio-Pleistocénu (přibliţně posledních 5,3 milionů let). Její utváření bylo v tomto období ovlivněno především klimatickými změnami, kdy docházelo ke střídání sušších a vlhčích podmínek během období glaciálů (interpluviálů) a interglaciálů (pluviálů), coţ se podepsalo nejen na vzniku nových druhů, ale i na jejich vnitrodruhové genetické struktuře (DEMENOCAL 2004). Jiţ od Pliocénu bývala vlhčí období střídána těmi aridnějšími, které se vyznačovaly především šířením savan, jeţ vyţadovaly adaptaci organismů na tento typ prostředí. Význačné epizody, při nichţ došlo k výrazným změnám v druhovém sloţení ţivočichů, nastaly před 3,4-3,2; 2,8-2,6; 2,4-2,2 a 2-1,8 miliony let (BOBE & BEHRENSMEYER 2004), kdy bylo
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