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! Osteichthyes ! Osteichthyes ! Osteichthyes ! Osteichthyes = Osteognathostomata- = Osteognathostomata- Ryby s.str.a Tetrapoda Ryby s.str.a Tetrapoda =Teleostomi – Koncoústé =Teleostomi – Koncoústé ryby …ale ryby …ale - Agnatha - Gnathostomata Sarcopterygii Osteognathostomata - Chondrichtyes Actinopterygii Teleostomi - Osteognathostomata †Acanthodii - Actinopterygii Chondrichthyes - Sarcopterygii †Placodermi Osteichthyes = Osteognathostomata (jméno Osteognathostomata: akcentující zahrnutí Tetrapoda) *lepidotrichia (dermální, kostěné), *šupiny s isopedinem (lamelární kost) • Endochondrální osifikace endoskeletu *dermatokranium jednotné • výrazný rozvoj a specialisace exoskeletu základní stavby, *jícnové výchlipky • Malý počet otolitů (3) v blanitém labyrintu (srv. otokonia u paryb) • Žaberní přepážky redukovány, společná žaberní dutina nebo vnější žábra lepidotrichia – • Výchlipky jícnu (plíce +/vs. plynový měchýř) kostěné, dermální, párové • lepidotrichia Osteognathostomata • Monophylum: Actinopterygii+ Sarcopterygii • Jícnová výchlipka (ventr. –plíce, dors. plynový měchýř) • Ztráta interbranchiálních sept • Lepidotrichia Šupiny ryb: • Endoskelet s peri- a endochondrální osifikací Kosmoidní (Sarcopterygii – • Dermální skelet (extrascapularia – kaud okraj zejm.fosilní), stropu lebky, praeoperculare +preoperculární kanál – začátek postr. čáry, operkulární komplex, gularia, Ganoidní (bichiři, kostlíni…) premaxilare, maxillare, qudratojugale, dentale, Leptoidní – kostěnné infrraorbitalia, temporalia, frontale, parietale, (Teleostei): cykloidní, parasphenoid, vomer, interclavicula, clavicula, ktenoidní: annuli, circuli cleithrum, cleithrum, supracleithrum, posttemporale Šupiny basálních Osteichthyes Osteichthyes GANOIDNÍ • Sarcopterygii,"#Svaloploutví:# • Ac.nopterygii,–, Ganoin (=Choanichthyes)# (vitreodentin) / Paprskoploutví# – Ac.nis.a,rec:Coelacanthimorpha# – CladisHa#(bichiři)# dentin/lamel.kost (lalokoploutví)# (isopedin) – Chondrostei#(chrupavči<)# – Choanata#(nozdra<)# – Neopterygii# KOSMOIDNÍ • Dipnoi#(dvojdyšní)# (Sarcopterygii) • Rhipidis.a## • Holostei# – (Osteolepiformes,#Tetrapoda#"# • Teleostomi# čtyřnožci)# • Tradiční3taxon3Crossopterygii3=3 Ac6nista+Rhipidista3(parafylum!)3 Šupiny basálních skupin Osteichthyes Actinopterygium Psarolepis33romeri33 Spodní#devon#Číny# Actinopterygii Archipterygium Sarcopterygii chiropterygium Ac.nopterygii,8,Paprskoploutví, Actinopterygii • volné#ploutve#bez#svalů,#s#kostěnými#lepidotrichiemi# Výrazně moderní • specialisace#dermálního#skeletu#lebky# skupina obratlovců: • autostylie#a#kineHsace,#specialisace#orobranchiálního# Teleostei: Cr –Q prostoru## • šupinovité#uspořádání#tělního#pokryvu#–#ganoidní# šupiny#(primiHvní#skupiny),#leptoidní#šupiny#(Teleostei)# • plynový#měchýř#(výj.bichiři)#postranní#čára## • slabé#čich.#laloky,#3#otolity# • oplození#vnější,#r"strategie# plynový měchýř, zmnožená basalia a radialia hrudní ploutve homocerkní ocasní ploutev Actinopterygii recentní taxony Teleostei – vrchol evoluce Cladistia – bichiř i čelistnatců 11 spp. Af (nejvyšší podíl Chondrostei – jeseteř i apomorfií, nejvyšší 30 spp. Eu-As-N Am diversita a ekologický Ginglymodi – kostlíni efekt) 7 spp. N Am 3.duplikace genomu, Halecomorphi – kaprouni redukce velikosti 1 sp. N Am genomu, radikální přestavba morfologie lebky, žaberního Teleostei – “kostnatí” aparátu i šupin 33 000 spp. 42 řádů, 430 čeledí, (elasmoidní=leptoidní 4080 rodů šupiny) Bichir: ocasní ploutev difycerkní Jeseter: ocasní ploutev heterocerkní Teleostei Ocasní ploutev Clupeomorpha: uroneurale a přímé ocasní napojení kostěných paprsků ploutev: Salmoniformes : redukce uroneuralií, homocerkní, zvětšení spodních, unikátní centrální jen 2 obratle- element (stegurale) urocentra, Ostariophysi: srůsty a tvarová /funkční uroneuralia Salmo specialisace, parahypurale, epurale etc kinetisce lebky ploutev redukce radiálií ocasní Teleostei: ocasní : ploutev Perca homocerkní Teleostei homocerkní Nejstarší Osteichthyes (svrchní silur-spodní devon) vykazují plesiomorfní znaky Sarcopterygii současně se znaky Acanthodi a Actinopterygií Bichir Polypterus – plochá masivní lebka s velkými kostmi, malá premaxila spojená s maxilárním obloukem, nedělené ploché operculum (symplesiomorfie se Sacropterygii a Paleoniscoidea) Kaproun Amia calva: nepárová gulární deska, diferenciace operkulárního aparátu – interoperculare s hyodiním ligamentem ! rozsáhlé zvětšení ústního prostoru a nasávání potravy (synapomorfie s Teleostei) Plesiomorfní stav: zuby na všech kostech ústního prostoru Elops - Teleostei: extrémní kinetisace lebky: >30 pohyblivých elementů + >50 svalů Perca okoun Teleostei: extrémní kinetisace lebky: >30 pohyblivých elementů + >50 svalů Mechanika čelistních struktur Teleostei:! Teleostei: svaly, kosti, zuby a klouby tvoří (překvapivě kinetický) Teleostei: modulární systém se vzájemně propojenými prvky extrémní extrémní kinetisace lebky: kinetisace lebky: >30 pohyblivých >30 pohyblivých elementů + >50 elementů + >50 svalů svalů Faryngeální čelisti a dentice hyale 1. Cerato-, hypo- 2. basi 3 4 5 Muraena (Elopomorpha) Specialisovaný rozvoj faryngeální dentice u více skupin Teleostei 1478 vs. 576 sp.!!! - Cichlidae + Labridae (suborder Labroidei) - obrovská druhová diversita - klíčová inovace: specialisovaná faryngeální dentice Actinopterygii – plynový měchýř physostomii, physoclistii plynová žláza a oválné políčko Ostariophysi Weberův aparát scaphium přeměna neurapofýz 1.-3. (4.) intercalarium obratle na sluchové kůstky tripus Ostariophysi: Weberův aparát Weberův aparát Cobitidae – redukce plynového měchýře, zvětšení a specialisace přední komory měchýře – diventriculum a chrupavčitou schránkou Srdce: Svalnatý bulbus arteriosus u Teleostei conus arteriosus (tepený násadec) vs. svalnatý bulbus arteriosus – Mozek Actinopterygii – apomorfie Teleostei malý telencephalon, odlišné stavby (evaginace) truncus arteriosus prostorová kompetice s optickým traktem conus arteriosus bulus arteriosus bulbus cordii kostlín kaproun Teleostei Actinopterygii – bazální linie všechna vodní sladkovodní, prostředí primárně mořské: Thermarces cerberus (Zoarcidae) – symbiont Riftia Alternativní fysiologické požadavky: Sladkovodní – vylučování Ekologie#a#Biodiversita# vody ledvinami Absorbce solí žábrami • Primárně#sladkovodní#–#6650#spp.# Ledviny: velké glomeruly, prodloužené tubuly s • Sekundárně#sladkovodní#–#1625#spp.# aktivní resorbcí NaCl, exkret dilutní (vysoký podíl • Mořské#ryby#konHnetálního#šelfu#–#9130# NH3) spp.# • Mořské#konHnentálních#prahů#–#1280#spp.# Mořské – vylučování solí • Epipelagiál#moří#–#225#spp.# ledvinou (ledvina malá, glomeruly u mnohých • Hlubokomořské#–#1100#spp.# chybí), exkret močovina Aktivní sekrece NaCl žábrami Mimořádný#ekonomický#význam# • Lov#sladkovodních#ryb#–#ca#11#mil.#tun# ročeně# • Tažné#–#ca#3#mil.###t/rok# • Mořské#–#ca#80#mil.#t/rok# • Ekonomický#chov#sladkovodních#ryb,# rybníkářství##"#výtěžnost#ca#0.4#t#/ha# • tropické#akvakultury#–#až#8#t#/ha# FAO,sta.s.ka, Actinopterygii recentní taxony rybolovu:, # celkem#ročně#ca# 100#milionů#tun,# Cladistia – bichiř i # od#2000#plynulý# 11 spp. Af pokles#výtěžku# Chondrostei – jeseteř i mořského#rybolovu# 30 spp. Eu-As-N Am Ginglymodi – kostlíni 7 spp. N Am Halecomorphi – kaprouni 1 sp. N Am Teleostei – “kostnatí” 33 000 spp. 42 řádů, 430 čeledí, 4080 rodů třída Actinopterygii Bichiři – Polypteriformes (Cladistia) =Mnohoploutní – Polypteriformes (Brachyopterygii) • tělo protáhlé, 5-18 hřbetních ploutví, svaln.násadec pár.ploutví, dva plicní vaky, typhlosolis, 11 spp. Polypterus – bichir Af ! + Paleonscoidei (devon-křída) Jeseteři – Accipenseriformes (Chondrostei) chrupačitá kostra, heterocerkní oc. ploutev ganoidní šupiny, prim.sladkovodní jeseterovití - Accipenseridae 30 spp. Eu-As-NAm • veslonosovití - Polyodontidae - 2 spp. NAm, SE As Kostnatí - Neopterygii • 29 000 spp. 450 fam. ca 40 řádů třída Actinopterygii Jeseteři – Accipenseriformes (Chondrostei) jeseterovití - Accipenseridae 30 spp. Eu-As-NAm • veslonosovití - Polyodontidae - 2 spp. NAm, SE As chrupačitá kostra, heterocerkní oc. ploutev, ganoidní šupiny, spodní ústa vousky, rypec, spir.řasa, K-strategové, až 100 let, velká vejce – kaviár, prim.sladkovodní, polyploidní populace 4n,8n,16n Acipenser - jeseter, 16 spp. Eu-Sib- N Am Huso huso – vyza až 7 m, 1,5 t. (dnes již jen Volha, Ural) Scaphirhynchus - lopatonosi (3 spp. Am Pseudoscaphirhynchus – 3 spp. Amudarja Polyodon3spathula3333 středovýchod#N#Am,# Veslonos#americký# Missisipi,#Missouri# 1,5#m###filtrátor# zooplanktonu,# “americký#kaviár”# na#farmách#I#v#Evropě,#nyní# volně#v#Dunaji# Psephurus gladius veslonos čínský Psephurus3gladius3.# Veslonos#čínský# Jang-c-jang, Až 8 m (největší sladkovodní ryba?) – nyní asi již vyhubena do roku 1980 – výlov 75 tun tření#v#přítocích,#migrace# do#dolního#toku# Neopterygii ! “HOLOSTEI – Mnohokostné“ • ganoidní šupiny, opistocélní obratle, typhlosolis, plyn.měchýř jako příd.dýchací orgán, hyoidní žábra, složité dermatokranium (např. opercularní komplex + cleithrum + angulare) – lebka akinetická, heterocekní oc. pl. homocerkní, plyn měch s dých.fcí, draví, ! Kostlíni Semionotiformes • 7 spp. NAm (fosilní Cr kosmop.) Lepisosteus ! Kaprouni - Amiiformes • 1 sp. NAm (jura kosmop.) Amia calva ! TELEOSTEI - Koncoústé Gynglimodi, # produkt#spodnotriasové#radiace# "#značná#diversifikace#koncem# triasu#a#v#juře# López"Arbarello#A#(2012)#PhylogeneHc#InterrelaHonships#of#Ginglymodian#Fishes# (AcHnopterygii:#Neopterygii).#PLoS#ONE#7(7)# Teleostei – vrchol evoluce TELEOSTEI - Kostnaté ryby čelistnatců (nejvyšší podíl ! Ocasní ploutev: malý počet zvětšených a spec. apomorfií,
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