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Jihočeská Univerzita V Českých Budějovicích

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Jihočeská Univerzita V Českých Budějovicích JIHO ČESKÁ UNIVERZITA V ČESKÝCH BUD ĚJOVICÍCH ZEM ĚDĚLSKÁ FAKULTA Studijní program: Prezen ční studium Studijní obor: Biologie a ochrana zájmových organism ů Katedra: Katedra biologických disciplín Vedoucí katedry: doc. RNDr. Ing. Josef Rajchard, Ph.D. MAGISTERSKÁ PRÁCE SKALIMETRICKÉ ZNAKY NEOTROPICKÝCH CICHLID TRIBU HEROINI: FYLOGENETICKÉ INTERPRETACE Vedoucí diplomové práce: doc. RNDr. Ing. Josef Rajchard, Ph.D. Konzultant diplomové práce: RNDr. Jind řich Novák, Ph.D. Autor: Bc. Barbora Stopová České Bud ějovice, leden 2012 Prohlašuji, že svoji magisterskou práci jsem vypracovala samostatn ě pouze s použitím pramen ů a literatury uvedených v seznamu citované literatury. Prohlašuji, že v souladu s § 47b zákona, č. 111/1998 Sb., v platném zn ění souhlasím se zve řejn ěním své diplomové práce, a to v nezkrácené podob ě (v úprav ě vzniklé vypušt ěním vyzna čených částí archivovaných Zem ědělskou fakultou JU) elektronickou cestou ve ve řejn ě p řístupné části databáze STAG provozované Jiho českou univerzitou v Českých Bud ějovicích na jejích internetových stránkách. Datum: 2. 2. 2012 Podpis ……………….. 3 Děkuji panu doc. RNDr. Ing. Josefovi Rajchardovi, Ph.D. za vedení mé práce a RNDr. Jindřichu Novákovi, Ph.D. za zap ůjčení literatury, drahocenné rady, optimismus a laskavý p řístup, za n ěž si ho velmi vážím! Mé díky v neposlední řad ě pat ří i Pavlovi Říhovi za ochotu p ři tvorb ě kladogram ů a neocenitelný p řístup. Milanovi Muškovi, MSc. za nápady, pomoc a kamarádskou podporu. Celé mé rodin ě, nejen za ochotu pomáhat s touto prací, ale i podporu během doby mého studia. Dále bych cht ěla pod ěkovat Old řichovi Říčanovi, Mgr., Ph.D. za poskytnutí materiál ů a pomoci p ři tvorb ě matic. 4 SOUHRN Práce „Skalimetrické znaky u st ředoamerických cichlid tribu Heroini “ si klade za cíl popis znak ů na šupinách u vybraných druh ů cichlid ze St řední Ameriky, zhodnotit vztahy na základ ě t ěchto znak ů ve fylogenezi skupiny a porovnat skalimetrický kladogram s molekulárním. První fosilie zástupc ů tribu Heroini ( Cichlidae: Cichlasomatinae ) jsou známé z dob před 10-15 miliony let. St ředoamerické druhy jsou typickým p říkladem specia čních proces ů typu alopatické speciace, tedy cestou geografické izolace povodí a vodních systém ů. Triby Heroini a Cichlasomatini jsou považovány za blízce p říbuzné, p řičemž Heroini tvo ří druhou nejv ětší skupinu neotropických cichlid a lze na nich demonstrovat nejv ětší rozmanitost těchto ryb ve St řední Americe. Heroini jsou považováni za skupinu monofyletickou. Vývoj tribu ve dvou v ětvích. První je reprezentovaná jihoamerickými rody bazálního postavení, antilským rodem Nandopsis , jihoamerickým odvozeným rodem Caquetaia a st ředoamerickými rody. Druhou v ětev tvo ří jihoamerický komplex ´Cichlasoma´ facetum a vyjmenované jihoamerické rody. Teoretická část této práce je v ěnována fylogenetickým poznatk ům týkajícím se tribu Heroini a popisu skupiny jako takové. V rámci vlastní práce jsou zachyceny jednotlivé znaky na šupinách,jejich mikrofotografie. Výsledky mé práce prokázaly nevhodnost skalimetrických znak ů jako fylogenetického nástroje; skalimetrické znaky zjevn ě nenesou využitelnou fylogenetickou informaci. Klí čová slova: fylogeneze, skalimetrické znaky, cichlidy, trib Heroini. ABSTRACT Work „Charakteristics of scales neotropical cichlids (tribe Heroini) – phylogenetic interpretation“ has the aim to describe charakteristics of scales selected species fish from the Central America and evaluace relationship with view on the group and compare scalimetric evolutional tree with molecular. The first fossils of species tribe Heroini are known from the age 10-15 millions BC. Cichlids from the Central America are typical example of alopatric evolution, thats mean the way geographical isolation watercourse. Relationship between Heroini and Cichlasomatini is really close and Heroini constitute the second largest tribe of neotropical cichlids. These fish are very varied. Heroini is considered as monofyletic group. Development of the tribe in two branches: The first represent the South American genus which have basal position, antil genus Nandopsis , derivate South America genus Caquetaia and the Central America genus. The second represent the South American complex ´Cichlasoma´ facetum and named South American genus. Theoretic part of this work is focused on phylogenetic knowledge from the other authors and group description. In the part with own issues there are characteristics of scales, microphotographs. Results of this work proved that charakteristics of squamation are not the suitable for phylogenetics analysis. Keywords: phylogeny, (scalimetric) characteristic of scale, cichlids, tribe Heroini 5 OBSAH : 1 Úvod ………………………………………………………………..………………….7 1.1. Čele ď Cichlidae …….………………………………….……………….......8 1.1.1. Charakteristika čeledi……………………………………………...8 1.1.2. Historie systematiky st ředoamerických cichlid……………………9 1.1.3. Fylogeneze čeledi Cichlidae ……………………………………...11 1.2. Fylogeneze tribu Heroini ………………………………………..………..12 1.3. Fylogenetický význam šupinných znak ů ………………………………..…15 2 Cíl práce ………………………………………...………………………………...…..17 3 Materiál a metodika ………………………………...………………………………...17 3.1. Materiál…………………………………………………………………..…17 3.2. Metodika………………………………………………………………....…20 3.2.1. P řehled skalimetrických znak ů…………………………………...21 3.2.2. Kladogramy ……………………………………………………...41 4 Vlastní výsledky ……………………………………………………….……………..42 5 Diskuse ………………………………………...……………………………………..61 6 Záv ěr ……………………………….…………………...…………………………….64 7 Seznam použité literatury ……...……………………………………...……………...65 8 Přílohy ……………………………………………...………………………………...71 8.1. Snímky zpracovaných druh ů ……………………………………………....71 8.2. Tabulka č. 6 - 58 skalimetrických znaků …………………………………100 8.3. Matice č. 1 - kódy jednotlivých stav ů …………………………………...104 6 1 Úvod Skalimetrické znaky, tedy znaky na šupinách, jsou tradi čně využívány jak pro identifikaci, tak i jako výpov ědní znaky fylogenetického postavení a systematického za řazení vyšších taxon ů. Pokusy o spojení skalimetrických (lepidologických) znak ů a představ o fylogenezi nižších taxon ů (trib ů, rod ů, druh ů) jsou relativn ě nové. Základní popisné studie p ředkládá Lippitschová (1988) a obecn ější srovnávací studie šupinných vzorc ů pocházejí od stejné autorky (Lippitsch 1990, 1991, 1995). Otázku, které skalimetrické znaky nepodléhají u ur čitého druhu ontogenetické variabilit ě řeší ve své práci Muška et al. (2005). Ve své diplomové práci se zabývám p říbuzností neotropických druh ů cichlid a fylogenezí v rámci tribu Heroini (pod čele ď Cichlasomatinae ) (Nelson 1976, Hanel 1998, Hanzák 1969). Tento tribus je rozd ělen do 27 st ředo a jihoamerických rod ů. Zahrnuje 145 druh ů a 93 zatím nepopsaných populací, kde lze v pr ůběhu času předpokládat povýšení na druhy (Muška 2003). Skupina Heroini je považována za monofyletickou (zahrnuje p říslušníky jediné fylogenetické linie) (Kullander 1998, Keenleyside 1991). Fylogenezi tribu Heroini na základ ě molekulárních dat rekonstruují Martin a Bermingham (1998), Říčan et al. (2008), Říčan (2005), Roe (1997), López-Fernández et al. (2010). P ředstavy o fylogenezi tribu na základ ě nemolekulárních dat p ředkládá Ritschelová (2003). Další práce na téma fylogenetických vztah ů skupiny Cichlidae přinášejí práce Martin (1998), Kullander (1983), Stiassny (1981, 1991), Farias (1999), Říčan (1999, 2001, 2005). Má magisterská práce voln ě navazuje na práci bakalá řskou, kterou jsem obhájila pod tématem „Srovnání morfoanatomických aspekt ů rané ontogeneze ryb skupin Physostomi a Physoclisti na p říkladu vybraných druh ů“. Po seznámení se s některými aspekty ontogeneze ryb mne za čalo zajímat, zda n ěkteré z ontogenetických znak ů lze popsat stavy, z nichž by bylo možné vyt ěžit i aspekt fylogenetický. Fylogenetické studie jsou v sou časné dob ě provád ěny u ryb, zejména s využitím genetických, resp. molekulárních znak ů. U znak ů morfologických a eko-etologických bývá obtížné odlišit ty, které fylogenetickou informaci nenesou. Jedním z posledních pokus ů, jak zohlednit časnou ontogenezi zbarvení pro p ředstavy fylogenetické představuje práce Říčana et al. 7 (2005). Sou časn ě je známo, že n ěkteré pokusy o vysv ětlení významu reproduk čního chování pro fylogenezi skon čily konstatováním, že jednotlivé typy chování (nap ř. typy pé če o potomstvo) vznikaly vícenásobn ě nezávisle na sob ě (p řehled uvádí Soukalová 2009). 1.1. Čeled Cichlidae 1.1.1. Charakteristika čeledi Obecn ě je pro cichlidy typické pom ěrn ě vysoké , laterárn ě zplošt ělé t ělo. Oproti tělu je p řítomna velká hlava, na níž bývá tukový hrbol (Obr. č. 1), velmi často p řízna čný pro samce (Baruš, Oliva et al. 1995). Postranní čára je dvouv ětevná a čichové jamky nemají p řepážku (Nelson 1976, Hanel 1998). B řišní ploutve jsou v hrdelní poloze. Hřbetní a řitní ploutev se skládají z dob ře odlišitelné p řední části s ostny a ze zadní části, která nese m ěkké paprsky. Typický je sexuální dimorfismus a barvom ěna podle momentálního rozpoložení, prost ředí, pohlavní dosp ělosti, stá ří či sociálního postavení (Hofmann a Novák 1998). Mnohé druhy jsou speciáln ě p řizp ůsobeny danému biotopu a druhu potravy v něm. Tato skute čnost se jasn ě odráží v rozší ření zejména t ěch cichlid, které jsou vázány na skalnatý region. To vede k vytvá ření mnoha barevných geografických poddruh ů, které pravd ěpodobn ě historicky vzešly z jediného p ředka. Ukázkovým příkladem je variabilita rodu Tropheus (Kraut 2008). Podle Keenlyside (1991) mají cichlidy pohyblivé čelisti se složitým svalovým aparátem s mnoha zuby.
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