PTAČÍ FYLOGENEZE Dovnitř Patří Šplhavci

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PTAČÍ FYLOGENEZE Dovnitř Patří Šplhavci E Afroaves – řada linií této skupiny je velmi rozrůzněna v Africe a pro Afriku stepokur kropenatý typická (vlhy, zoborožci, mandelíci), ale fosilie africký původ nepodporují. Mnoho (Pterocles burchelli) linií bylo řazeno do řádu srostloprstých, ale ten není monofyletický, poněvadž PTAČÍ FYLOGENEZE dovnitř patří šplhavci. Patří sem i myšáci (Coliiformes) a trogoni. F Australaves – obdobně jako u Afroaves dnes sice řada linií žije jen FYLOGENEZE NA HRUBÉ ÚROVNI A PTAČÍ ŘÁDY v australasijské oblasti, ale fosilní zástupci byli nalezeni všude možně, takže o původu skupiny těžko něco soudit. Zobrazený fylogenetický stromeček zahrnuje jen velké skupiny, přičemž co je „velká“ skupina, je docela arbitrární. Aby vůbec bylo možné dělat O Passerimorphae (nebo Psittacopasserae) – papoušci korektní fylogenetické analýzy, nesmí být těch skupin moc a zároveň je potřeba, aby to byly skupiny, o nichž se předpokládá, že jsou a pěvci sdílejí řadu znaků, například schopnost učit se zpěvu, která byla monofyletické. Reálně se totiž sekvenují samozřejmě jednotliví zástupci (tedy jeden druh, respektive přímo jeden jedinec z dané skupiny), považovaná za klíčovou pro jejich sloučení (pokládala se za znak vzniklý jen a potřebujeme tedy, aby skutečně skupinu reprezentovali. Proto by nestačilo brát zástupce tradičních ptačích řádů, které známe z učebnic, jednou u jejich společného předka, tedy synapomorfii). Ovšem vzhledem k tomu, jelikož se ukázalo, že řada z nich monofyletických není. Tradiční „dravci“ byli například polyfyletičtí, poněvadž zahrnovali monofyletickou že u pěvců je s jistotou známa jen ve skupině Oscines (zpěvní) a navíc se vyskytuje skupinu složenou z orlů, jestřábů, káňat apod. (včetně kondorů), ale taky sokolovité (poštolky, sokoly apod.), kteří jsou ve skutečnosti blízce také u zcela nezávislých kolibříků, musela vzniknout víckrát konvergentně, a to příbuzní pěvcům a papouškům. Srostloprstí (vlhy, dudci, ledňáčci apod.) zase byli parafyletičtí, poněvadž se ukázalo, že dovnitř celé skupiny buď třikrát (u zpěvných, papoušků a kolibříků), nebo dvakrát (u společného patří šplhavci (datli, žluny, strakapoudi, krutihlav), kteří ovšem byli vyčleňováni jako samostatný řád. Čili bylo potřeba analyzovat například předka papoušků a pěvců a u kolibříků), přičemž u dvou skupin pěvců by pak jednotlivé podskupiny původních srostloprstých zvlášť, poněvadž jinak by nebylo jasné, zástupce jaké skupiny vlastně studujeme. nezávisle zanikla. Pěvci se totiž štěpí primárně na novozélandské pokřovníky (kteří schopnost učit se zpěvu nemají) a zbytek, ten pak na křikavé (Suboscines) 24 29 a zpěvné (Oscines). Pěvci tvoří přibližně polovinu všech ptačích druhů. 10 32 seriema rudozobá slunatec nádherný ostnák trnitý (Jacana spinosa) turako světlý (Corythaixoides concolor) kurol madagaskarský (Leptosomus discolor) (Cariama cristata) (Eurypyga helias) Oscines G Aequornithia (nebo Aequornithes) jsou ptáci vázaní na vodu, kteří 1 zpěvní O byli řazeni do tradičních řádů brodiví (čápi, volavky a ibisové), veslonozí 7 Suboscines (pelikáni a kormoráni), trubkonosí (albatrosi a buřňáci), tučňáci a potáplice. Passeriformes 2 křikaví Passeri- Z tohoto fylogenetického stromu by plynulo, že sesterskou skupinu Acanthisitti morphae 3 pokřovníci k Aequornithia by tvořil klad sestávající z jihoamerického slunatce, Psittaciformes 4 papoušci F novokaledonského kagu a faetonů, ale nelze to stoprocentně určit, jelikož Falconiformes jde taky o bazální rozvětvení někdy na začátku třetihor. 5 sokolovití Australaves H 23 Cariamiformes H Telluraves zahrnuje kromě pěvců, papoušků, datlů či vlh i několik 6 seriemy nezávislých linií dravých ptáků, včetně seriem, které byly původně řazeny ke ) Coraciiformes + Piciformes 14 7 srostloprstí + šplhavci krátkokřídlým, nicméně žerou obratlovce a jsou blízce příbuzné vymřelým Bucerotiformes obřím hrůzoptákům (Phorusrhacoidea). Je pravděpodobné, že dravost je zoborožci 8 Telluraves Trogoniformes Coracii- tedy původní znak předka celé skupiny. 9 trogoni morphae Leptosomiformes 10 kurol I Pelecanimorphae obsahuje klady řazené tradičně do řádů Coliiformes 6 11 myšáci E brodivých a veslonohých, které ale byly polyfyletické – pelikáni jsou nejblíže Sarcomphus papaSarcomphus Strigiformes Afroaves příbuzní člunozobci, k nim zřejmě patří kladivouš (a asi volavky), naopak 12 sovy kormoráni jsou zvlášť. Čápi v analýze Jarvise a kolegů nefigurují, ale hadilov písař Accipitridae pravděpodobně také představují samostatnou, časně odvětvenou skupinu (Sagittarius serpentarius) kolibřík zelený 13 jestřábovití Accipitri- Cathartidae morphae Passerea v rámci tohoto kladu. 27 (Colibri thalassinus) 14 kondoři J Pelecanidae Cursorimorphae – jeřábi byli tradičně řazeni do řádu krátkokřídlí, momot černolící ) 15 pelikáni ovšem tento řád se zcela rozpadl, takže není nijak překvapivé, že vycházejí jako G (Momotus momota) Ardeidae příbuzní dlouhokřídlých a bahňáků (tradičně spolu řazených do řádu 16 volavky Pelecani- kondor královský ( kondor královský B Threskiornithidae morphae I Charadriiformes). Ale zase jde o skupiny vzniklé během oné rychlé radiace na 17 ibisové začátku třetihor, a jejich vztahy tedy nejsou jasné. Neoaves – nedávné Phalacrocoracidae molekulární analýzy naznačovaly 18 kormoráni K Hoacin vždycky představoval klasický fylogenetický oříšek. Občas se řadil tinama argentinská rozdělení tohoto kladu na dvě velké Procellariiformes 19 buřňáci Procellarii- ke kukačkám, někdy k hrabavé „drůbeži“, ale nikdy to nebylo moc přesvědčivé. (Eudromia elegans) skupiny, Metaves a Coronaves. Do Sphenisciformes Aequornithia Asi proto, že jde taky o potomka linie odštěpené hned na začátku radiace Centropus toulou Centropus 20 morphae Metaves měli patřit kromě skupin tučňáci Neoaves, kdy přesná sekvence odštěpování nezanechala fylogenetický signál. Gaviiformes Gavii- zde sdružených do kladu Columbea 21 potáplice ještě lelkové a příbuzní (včetně Phaethontiformes morphae L Caprimulgimorphae (dříve Strisores) – lelci a jejich příbuzní 22 faetoni (gvačarové, lelkouni apod.) jsou noční ptáci živící se vesměs v letu vzdušným rorýsů a kolibříků), faetoni, slunatec Eurypygiformes Phaethonti- a kagu, do Coronaves by pak patřila 23 slunatec + kagu morphae planktonem (mají obří, za lovu doširoka rozevřený zobák), jen gvačarové se živí zbývající většina ptačích druhů. Charadriiformes plody a jsou vůbec zvláštní tím, že hnízdí v jeskyních a ve tmě se orientují kukačka tulu ( 24 bahňáci (dlouhokřídlí) pomocí echolokace. Všechny tyto skupiny byly dlouho považovány za příbuzné 13 Jenže toto rozdělení ve skutečnosti Gruiformes Cursori- J podporuje jen jediný intron 25 jeřábi morphae svišťounů, tedy rorýsů a kolibříků. Svišťouni nicméně patří dovnitř celého kladu, beta-fibrinogenu, a když se Opisthocomiformes jsou to tedy vlastně takoví odvození lelkové (vzpomeňte si na to, až uvidíte ve analyzovaly větší soubory genů, 26 hoacin K městě proletovat hejna rorýsů těsně před setměním, když ostatní ptáci už 30 ukázalo se, že to nesedí. Caprimulgiformes Caprimulgi- usínají). Kolibříci jsou samozřejmě v řadě ohledů zcela extrémní a je k nevíře, že 27 lelkové + svišťouni morphae z velkých dinosaurů mohlo evolucí přes „lelky“ vzniknout zvíře o velikosti Otidiformes L 28 dropi čmeláka, svým způsobem letu (a vlastně i výživy) odpovídající spíše hmyzu. 16 B NEOAVES Musophagiformes Otidi- turakové M Otidimorphae – kukačky se tradičně dávají dohromady s turaky, 29 morphae Cuculiformes M nicméně příbuznost těchto dvou skupin s dropy může působit zvláštně. Ovšem 30 kukačky zemní kukačky jsou velké, běhají a loví na zemi jako dropi. Navíc jde o starou ara zelenokřídlý Mesitornithiformes 4 31 mesiti D skupinu vzniklou během oné explozivní radiace Neoaves na začátku třetihor čili (Ara chloroptera) Pterocliformes na rozrůznění měla spoustu času. 38 32 stepokuři Columbi- Columbiformes morphae NEOGNATHAE 33 měkkozobí Phoenicopteriformes 36 25 34 plameňáci Phoenicopteri- Columbea Podicipediformes 35 potápky morphae Galiiformes čája obojková 36 hrabaví 37 C (Chauna torquata) Anseriformes GALLOANSERAE 37 vrubozobí D Tinamiformes + Struthioniformes Columbea sdružuje měkkozobé 38 běžci (včetně tinam) 2 (holuby a hrdličky) s potápkami A PALAEOGNATHAE křída paleogén ) a plameňáky, což by málokoho napadlo, kladivouš africký jeřáb popelavý ostnák africký zvlášť když měkkozobí jsou v tradičních (Scopus umbretta) (Grus grus) 24 (Actophilornis africanus) 110 100 90 80 70 60 50 40 30 20 10 0 příručkách řazeni poblíž pěvců. Jsou to však docela zvláštní ptáci: nestaví si miliony let C hnízdo, co by stálo za řeč, jen plošinku hýl obecný zoborožec kaferský Galloanserae – sem patří A bělolící drop z klacíků, a mláďata krmí zvláštním (Pyrrhula pyrrhula) (Bucorvus vrubozobí spolu s hrabavými ptáky, Palaeognathae byli tradičně označovaní jako „běžci“, jenže kromě Dvoustranu připravil Eupodotis afra „mlékem“ vytvářeným ve voleti, přičemž leadbeateri) ( včetně tabonů z australasijské oblasti pštrosa, nanduů, emuů, kasuárů a kiviů sem patří i tinamy, které sice létají David Storch; něco podobného dělají i plameňáci! Navíc (ti nesedí na vejcích, ale inkubují je nerady, ale umějí to. Tinamy navíc patří někam dovnitř celé skupiny, čímž padá snímky © Čestmír Číhalík, není divu, že tak stará skupina jako v hromadách tlejícího listí nebo teplého představa původní nelétavosti předka všech dnes nelétavých forem (to by Tomáš Grim, Zdeněk Columbea je ekologicky a morfologicky sopečného popela). Patzelt, Petr Pokorný, diverzifikovaná (nesmí nás mást, že je tinamy musely schopnost letu zase zpátky získat, což není moc pravděpodob- Ondřej Sedláček,
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