Úvod 3: Fylogenese

• Primární
viscerokranium:
Historie Mammalia:
Mammalia:
Fosilní záznam
mandibularequadratumhyomandibulare
(stapes)
• Synapsida - nejstarší Amniota - od
středniho Karbonu (ca 320 My)
• Cotylosauria vs. Pelycosauria (nejhojnější
skupina v permu a triasu)
• Dermální
skelet:
– Dentale,
coronoideum,
angulare,
suprangulare,
prearticulare
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Pelycosauria
225
THERAPSIDA
Non-Mammalian
Ca
sea
sau
Synapsida
r ia
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Synapsida
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Auta po mo rphies o f the ma in cla des o f sy na psids
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Michel Laurin and Robert R. Reisz The classification of synapsids may be summarized
as follows:
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Eothyrididae ==Caseasauria=| |
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Caseidae | |
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Varanopseidae | | | |
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Ophiacodontidae | | | | | |
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Edaphosauridae | | | | =====| | | | ===================================
Haptodus garnettensis | | | | | ==Eupelycosauria=| | | |
=============================== Palaeohatteria ==A=| | | | ==B=| | |
=========================== Pantelosaurus ==Sphenacodontia=| | | ==C=| |
======================= Cutleria ==D=| | ==E=| === Sphenacodontidae
==Sphenacodontoidea=| === Therapsida This classification and the list of apomorphies
given below is taken from recent studies by Reisz (1986), Hopson (1991), Reisz et al.
(1992), Berman et al. (1995), and Laurin and Reisz (1996), to which the readers should
refer for further information.
Caseasauria (eothyridids and caseids) exhibits the following autapomorphies:
A long external naris with an external narial shelf (Fig. 1A).
A pointed rostrum formed by the dorsal process of the premaxilla (Fig. 1A).
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Temporální jáma mezi
squamosum, jugale, postorbitale,
occipitale orientováno vertikálně
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Michel Laurin and
Robert R. Reisz Tree of Life
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http://tolweb.org/access
ory/Synapsid_Classifica
tion_&_Apomorphies?a
cc_id=466
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Click o n an imag e to v iew larg er v ersio n & d ata in a n ew win d o w
Fig ure 1 . Sy n ap sid sk u lls in lateral v iew. A, Co ty lo rh y n ch u s, a Lo wer Permian caseid . B, Varan o sau ru s, a Lo wer Permian o p h iaco d o n tid . C, Dimetro d o n , a Lo wer Permian sp h en aco d o n tid . D, Titan o p h o n eu s, an Up p er Permian th erap sid . Red rawn fro m B, Berman et al
. (1 9 9 5 ); C, Ro mer an d Price (1 9 4 0 ), an d D, Orlo v (1 9 5 8 ). Ab b rev iatio n s: An , an g u lar; Ar, articu lar; Bc, b rain case; D, d en tary ; E, ep ip tery g o id ; F, fro n tal; J, ju g al; L, lacrimal; M, max illa; N, n asal; P, p arietal; Pm, p remax illa; Po , p o sto rb ital; Po f, p o
stfro n tal; Prf, p refro n tal; Pt, p tery g o id ; Q, q u ad rate; Qj, q u ad rato ju g al; Sa, su ran g u lar; Sp , sp len ial; Sq , sq u amo sal; St, su p ratemp o ral; T, tab u lar. Scale b ars eq u al 1 cm.
Th e main au tap o mo rp h y o f Eo thy ridida e (Eo th y ris an d Casea) is as fo llo ws:
Larg e can in ifo rm to o th is p resen t in an terio r p art o f th e max illa.
Ca seida e p o ssesses th e fo llo win g au tap o mo rp h ies:
A larg e, h ig h ex tern al n aris (Fig . 1 A). Th e ex tern al n aris o f eo th y rid id s is lo n g , b u t it is mo re p rimitiv e in th at it is relativ ely lo w (n arro w).
Po sterio r marg in al teeth b earin g a lo n g itu d in al ro w o f cu sp s. Similar teeth are fo u n d in man y o th er h erb iv o ro u s amn io tes, su ch as h erb iv o ro u s sq u amates (ig u an as) an d man y g ro u p s o f h erb iv o ro u s d in o sau rs.
Red u ced p h alan g eal fo rmu la, ran g in g fro m 2 -3 -4 -4 -3 to 2 -2 -2 -3 -2 . Primitiv ely in sy n ap sid s, th e p h alan g eal fo rmu la was 2 -3 -4 - 5 -3 in th e h an d an d 2 -3 -4 -5 -4 in th e fo o t.
Click o n an imag e to v iew larg er v ersio n & d ata in a n ew win d o w
Fig ure 2 . Sy n ap sid sk u lls in d o rsal v iew. A, Co ty lo rh y n ch u s, a Lo wer Permian caseid . B, Varan o sau ru s, a Lo wer Permian o p h iaco d o n tid . C, Dimetro d o n , a Lo wer Permian sp h en aco d o n tid . D, Titan o p h o n eu s, an Up p er Permian th erap sid . Red rawn fro m B, Berman et al.
(1 9 9 5 ); C, Ro mer an d Price (1 9 4 0 ), an d D, Orlo v (1 9 5 8 ). Ab b rev iatio n s: Bc, b rain case; F, fro n tal; J, ju g al; L, lacrimal; M, max illa; N, n asal; P, p arietal; Pa, p alatin e; Pm, p remax illa; Po , p o sto rb ital; Po f, p o stfro n tal; Pp , p o stp arietal; Prf, p refro n tal
; Qj, q u ad rato ju g al; Sq , sq u amo sal; St, su p ratemp o ral; T, tab u lar. Scale b ars eq u al 1 cm.
Au tap o mo rp h ies o f E upely co sa uria :
A lo n g , n arro w su p ratemp o ral (Fig . 2 B-D). Th e su p ratemp o ral o f caseasau rs is almo st as wid e as lo n g (len g th to wid th ratio less th an two ).
Th e fro n tal co n trib u tes to at least o n e th ird o f th e d o rsal o rb ital marg in (th e o rb ital co n trib u tio n o f th e fro n tal is n arro wer in caseasau rs) (Fig . 2 B-D).
Au tap o mo rp h ies o f V a ra no pseida e:
Marg in al d en titio n co mp o sed o f stro n g ly cu rv ed , med io laterally flatten ed teeth .
Occip ital flan g e o f sq u amo sal red u ced .
Narro w zy g o matic arch .
Au tap o mo rp h ies o f th e clad e in clu d in g Op h iaco d o n tid ae, Ed ap h o sau ria, an d Sp h en aco d o n tia (u n n amed cla de A in th e ab o v e classificatio n ):
Nasal lo n g er th an p arietal at mid lin e (it is sh o rter in o th er sy n ap sid s) (Fig . 2 B-D).
A n arro w p o sterio r p ro cess o f th e p o sto rb ital (th is p ro cess is b ro ad in o th er sy n ap sid s).
A lo n g co n trib u tio n o f th e ju g al to th e v en tral ed g e o f th e sk u ll (th is co n trib u tio n is n arro w, if p resen t, in o th er sy n ap sid s).
A co n trib u tio n o f th e sq u amo sal to th e zy g o matic arch (in o th er sy n ap sid s, th is arch is co mp o sed ex clu siv ely o f th e ju g al an d th e q u ad rato ju g al) (Fig . 1 B-D).
Ecto p tery g o id teeth ab sen t.
Au tap o mo rp h ies o f Ophia co do ntida e in clu d e:
An to rb ital reg io n o f sk u ll at least twice as lo n g (Fig . 2 B-D) as th e p o sto rb ital reg io n (it is sh o rter in o th er sy n ap sid s).
Nasal lo n g er th an fro n tal (it is sh o rter th an th e fro n tal in o th er sy n ap sid s).
Paro ccip ital p ro cess sh o rt, n o t ex ten d in g to th e ch eek (it is lo n g er an d reach es th e q u ad rate o r th e sq u amo sal in o th er sy n ap sid s) (Fig . 3 B).
Tab u lar n o t ex ten d in g v en trally b ey o n d th e lev el o f th e su p rao ccip ital (in o th er sy n ap sid s, it ex ten d s farth er v en trally ) (Fig . 3 B).
Click o n an imag e to v iew larg er v ersio n & d ata in a n ew win d o w
Fig ure 3 . Sy n ap sid sk u lls in o ccip ital v iew. A, Co ty lo rh y n ch u s, a Lo wer Permian caseid . B, Varan o sau ru s, a Lo wer Permian o p h iaco d o n tid . C, Dimetro d o n , a Lo wer Permian sp h en aco d o n tid . D, Titan o p h o n eu s, an Up p er Permian th erap sid . Red rawn fro m B, Berman et
al. (1 9 9 5 ); C, Ro mer an d Price (1 9 4 0 ), an d D, Orlo v (1 9 5 8 ). Ab b rev iatio n s: Bc, b rain case; J, ju g al; P, p arietal; Pp , p o stp arietal; Ps, p arasp h en o id ; Pt, p tery g o id ; Q, q u ad rate; Qj, q u ad rato ju g al; S, stap es; Sq , sq u amo sal; St, su p ratemp o ral; T, tab u lar. Sc
ale b ars eq u al 1 cm.
Au tap o mo rp h ies o f U nna med cla de B (Ed ap h o sau rid ae an d Sep h en aco d o n tia) in clu d e:
Orb ital p ro cess o f fro n tal p resen t (in o th er sy n ap sid s, th e fro n tal may co n trib u te to th e o rb it, b u t it d o es n o t ex ten d farth er laterally in th is area th an elsewh ere) (Fig . 2 C, D).
Lateral su rface o f p o sto rb ital recessed (it is flat in o th er sy n ap sid s).
An tero v en tral p ro cess o f q u ad rato ju g al ab sen t (in o th er sy n ap sid s, th e q u ad rato ju g al is mu ch larg er in lateral v iew b ecau se it h as a lo n g an tero v en tral p ro cess) (Fig . 1 C, D).
Au tap o mo rp h ies o f E da pho sa urida e in clu d e:
Presen ce o f lateral tu b ercles o n n eu ral sp in es.
Neu ral sp in es lo n g an d circu lar in cro ss sectio n d istal to a laterally co mp ressed p ro x imal sectio n .
Cerv ical n eu ral sp in es lean an terio rly , wh ereas p o sterio r sp in es lean p o sterio rly .
Th ese th ree au tap o mo rp h ies co n trib u te to th e p ecu liar mo rp h o lo g y o f th e sail o f ed ap h o sau rs.
Au tap o mo rp h ies o f S phena co do ntia in clu d e:
Max illary su p racan in e b u ttress p resen t (th is stru ctu re is a th ick en in g o f th e max illa v isib le o n its in tern al su rface, d o rsal to th e can in ifo rm teeth ).
Premax illary teeth in d eep so ck ets. Th e teeth o f o th er sy n ap sid s are imp lan ted in sh allo w so ck ets.
Au tap o mo rp h ies o f Ha pto dus g a rnettensis in clu d e:
Th e p resen ce o f a small facial ex p o su re o f th e sep to max illa. In o th er sy n ap sid s, th e sep to max illa is restricted to th e ex tern al n aris, ex cep t in mo st th erap sid s, wh ere it h as a larg e facial ex p o su re.
Po stcan in ifo rm teeth ro b u st, ch isel- sh ap ed . Th e sh ap e o f marg in al teeth is q u ite v ariab le in early sy n ap sid s, b u t th eir mo rp h o lo g y in H. g arn etten sis is u n iq u e.
Teeth p resen t o n th e p alatal ramu s o f th e p remax illa. In o th er sy n ap sid s, th is b o n e o n ly b ears marg in al teeth .
Au tap o mo rp h ies o f u nna med cla de C (Palaeo h atteria, Pan telo sau ru s, Cu tleria, an d Sp h en aco d o n to id ea) in clu d e:
Presen ce o f a reflected lamin a o f th e an g u lar (Fig . 1 C, D). Th is v en tral flan g e o f th e an g u lar ev en tu ally b ecame p art o f th e ty mp an ic an n u lu s o f mammals (th e rin g th at su p p o rts th e ear d ru m).
Au tap o mo rp h ies o f P a la eo ha tteria lo ng ica uda ta in clu d e:
A co n tact b etween th e p o sto rb ital an d th e su p ratemp o ral.
Au tap o mo rp h ies o f u nna med cla de D (Pan telo sau ru s, Cu tleria, an d Sp h en aco d o n to id ea) in clu d e:
Ju g al an terio r p ro cess b ro ad an terio rly (it is n arro w in o th er sy n ap sid s) (Fig . 1 C).
On ly o n e au tap o mo rp h y was id en tified in P a ntelo sa urus sa x o nicus:
Po sterio r en d o f d en tary well b elo w th e p o sterio r ed g e o f th e jaw. In o th er sy n ap sid s, th e p o sterio r en d o f th e d en tary is n ear th e d o rsal ed g e o f th e jaw.
Au tap o mo rp h ies o f u nna med cla de E (Cu tleria an d Sp h en aco d o n to id ea) in clu d e:
Nasal lo n g er th an th e fro n tal (th e n asal is sh o rter in mo st o th er sy n ap sid s). A lo n g n asal ap p eared co n v erg en tly in o p h iaco d o n tid s (Fig . 2 B-D).
A d eep n o tch o n th e p arietal fo r th e an terio r tip o f th e su p ratemp o ral (th is n o tch is sh allo w in o th er sy n ap sid s) (Fig . 3 C).
Cu ttin g ed g es o n marg in al teeth (th ese ed g es are ab sen t in mo st o th er sy n ap sid s).
Au tap o mo rp h ies o f C utleria wilma rthi in clu d e:
An an terio rly co n cav e, in d en ted ju g al-sq u amo sal su tu re (in mo st o th er sy n ap sid s, th is su tu re ex ten d s p o stero v en trally o r an tero v en trally ).
Au tap o mo rp h ies o f S phena co do nto idea in clu d e:
Fro n tal o rb ital p ro cess ex ten d s far laterally (it is p o o rly d ev elo p ed , if p resen t, in o th er sy n ap sid s) (Fig . 2 C).
Deep p refro n tal p o ck et (a co n cav ity n ear th e an tero d o rsal ed g e o f th e o rb it).
Vo merin e teeth ab sen t (th e v o mer b ears a sh ag reen o f d en ticles in o th er sy n ap sid s).
Au tap o mo rp h ies o f S phena co do ntida e in clu d e:
Presen ce o f a v en tral n arial p ro cess o f th e n asal (ju st p o sterio r to th e ex tern al n aris) (Fig . 1 C).
Fro n tal an terio r p ro cess n arro wer th an its p o sterio r p ro cess (th ese two p ro cesses are eq u ally wid e in o th er sy n ap sid s) (Fig . 2 C).
Po sto rb ital-sq u amo sal co n tact ex ten siv e (th is co n tact is n arro w in o th er sy n ap sid s, wh en p resen t) (Fig . 1 C).
Su p ratemp o ral co n tactin g p o sto rb ital (th is co n tact is ab sen t in mo st o th er eu p ely co sau rs) (Fig . 2 C).
Paro ccip ital p ro cess ex ten d s v en tro laterally an d p o sterio rly (it ex ten d s laterally in mo st o th er sy n ap sid s) (Fig . 3 C). Th is ch aracter is co n v erg en t in so me th erap sid s (Fig . 3 D).
Can in ifo rm ro o t b u lg es in to ch o an a (in o th er sy n ap sid s, th e can in ifo rm to o th d o es n o t co n strict th e ch o an a).
Au tap o mo rp h ies o f T hera psida in clu d e:
In terp tery g o id v acu ity sh o rt (less th an 2 0 p ercen t o f th e len g th o f th e p alate, measu red fro m its an terio r en d to th e p o sterio r ed g e o f th e tran sv erse flan g e o f th e p tery g o id ).
Brain case firmly su tu red to b ack o f d ermal sk u ll ro o f. Th e b rain case o f mo st o th er sy n ap sid s is attach ed fairly lo o sely to th e p o sterio r ed g e o f th e ch eek an d sk u ll tab le. In th ese tax a, th e p aro ccip ital p ro cess is h eld b etween relativ ely th in sh eets o f t
h e tab u lar an d sq u amo sal. Brain cases o f th ese early sy n ap sid s o ften b ecame d etach ed fro m th e rest o f th e sk u ll p rio r to fo ssilizatio n . In th erap sid s, th e b rain case remain s firmly su tu red to th e d ermal sk u ll elemen ts.
No mo re th an twelv e u p p er p o stcan in ifo rm teeth (Fig . 1 D). Oth er sy n ap sid s h av e at least th irteen p o stcan in ifo rm teeth (Fig . 1 A, B).
Ecto p tery g o id teeth ab sen t. Oth er sy n ap sid s h av e small teeth (d en ticles) o n th e ecto p tery g o id .
Th is list o f au tap o mo rp h ies tak es in to co n sid eratio n th e recen t d isco v ery o f Tetracerato p s, th e o ld est k n o wn th erap sid (Lau rin an d Reisz, 1 9 9 0 , 1 9 9 6 ). Tetracerato p s lack s man y d eriv ed ch aracters p resen t in o th er th erap sid s. Th erefo re, th is list d iffers fr
o m o th er p u b lish ed lists o f au tap o mo rp h ies o f th erap sid s (Ho p so n an d Barg h u sen , 1 9 8 6 ; Gau th ier et al., 1 9 8 8 ; Kemp , 1 9 8 8 ; Ho p so n , 1 9 9 1 ).
Pag e last u p d ated March 2 1 , 1 9 9 7 .
References
Berman D. S., R. R. Reisz, J. R. Bo lt, an d D. Sco tt. 1 9 9 5 . Th e cran ial an ato my an d relatio n sh ip s o f th e sy n ap sid Varan o sau ru s (Eu p ely co sau ria: Op h iaco d o n tid ae) fro m th e Early Permian o f Tex as an d Ok lah o ma. An n als o f Carn eg ie Mu seu m 6 4 : 9 9 -1 3 3 .
Gau th ier J., A. G. Klu g e, an d T. Ro we. 1 9 8 8 . Amn io te p h y lo g en y an d th e imp o rtan ce o f fo ssils. Clad istics 4 : 1 0 5 -2 0 9 .
Ho p so n J. A. 1 9 9 1 . Sy stematics o f th e n o n mammalian Sy n ap sid a an d imp licatio n s fo r p attern s o f ev o lu tio n in sy n ap sid s. In : H.-P. Sch u ltze an d L. Tru eb (ed .) Orig in s o f th e h ig h er g ro u p s o f tetrap o d s-Co n tro v ersy an d Co n sen su s (1 ): 6 3 5 -6 9 3 . Ith aca: Co rn ell U
n iv ersity Press.
Ho p so n J. A. an d H. R. Barg h u sen . 1 9 8 6 . An An aly sis o f Th erap sid Relatio n sh ip s. In : P. D. M. Nich o las Ho tto n III, Jan J. Ro th , an d E. Caro l Ro th (ed .) Th e Eco lo g y an d Bio lo g y o f Mammal-lik e Rep tiles 8 3 -1 0 6 . Wash in g to n : Smith so n ian In stitu tio n Press.
Kemp T. S. 1 9 8 8 . In terrelatio n sh ip s o f th e Sy n ap sid a. In : M. J. Ben to n (ed .) Th e Ph y lo g en y an d Classificatio n o f th e Tetrap o d s (2 ): 1 -2 2 . Ox fo rd : Claren d o n Press.
Lau rin M. an d R. R. Reisz. 1 9 9 0 . Tetracerato p s is th e o ld est k n o wn th erap sid . Natu re 3 4 5 : 2 4 9 -2 5 0 .
Lau rin M. an d R. R. Reisz. 1 9 9 6 . Th e o steo lo g y an d relatio n sh ip s o f Tetracerato p s in sig n is, th e o ld est k n o wn th erap sid . Jo u rn al o f Verteb rate Paleo n to lo g y 1 6 : 9 5 -1 0 2 .
Orlo v Y. A. 1 9 5 8 . Primitiv e d in o cep h alian s o f th e Ish eev a fau n a (Titan o su ch i). Tran sactio n s o f th e Palaeo n to lo g ical In stitu te o f th e Acad emy o f Scien ces o f th e USSR 7 2 : 1 -1 1 3 .
Reisz R. R. 1 9 8 6 . Pely co sau ria. En cy clo p ed ia o f Paleo h erp eto lo g y 1 7 A: 1 -1 0 2 .
Reisz R. R., D. S. Berman , an d D. Sco tt. 1 9 9 2 . Th e cran ial an ato my an d relatio n sh ip s o f Seco d o n to sau ru s, an u n u su al mammal- lik e rep tile (Pely co sau ria: Sp h en aco d o n tid ae) fro m th e early Permian o f Tex as. Zo o lo g ical Jo u rn al o f th e Lin n ean So ciety 1 0 4 : 1 2 7 -1
84.
Ro mer A. S. an d L. I. Price. 1 9 4 0 . Rev iew o f th e Pely co sau ria. Vo l. 2 8 . Geo lo g ical So ciety o f America, Sp ecial Pap ers, New Yo rk : Arn o Press.
Abo ut This Pa g e
Mich el Lau rin
Un iv ersité Paris, Fran ce
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Ro b ert R. Reisz
Un iv ersity o f To ro n to , Mississau g a, On tario , Can ad a
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Pag e co p y rig h t © 1 9 9 7 M ich el Lau rin an d Ro b ert
R. Reisz
Caseasauria
Eothyrididae
Caseidae
Synapsid skulls in lateral view. A, Cotylorhynchus, a Lower Permian caseid. B, Varanosaurus, a Lower Permian
ophiacodontid. C, Dimetrodon, a Lower Permian sphenacodontid. D, Titanophoneus, an Upper Permian therapsid.
Redrawn from B, Berman et al. (1995); C, Romer and Price (1940), and D, Orlov (1958). Abbreviations: An,
angular; Ar, articular; Bc, braincase; D, dentary; E, epipterygoid; F, frontal; J, jugal; L, lacrimal; M, maxilla; N,
nasal; P, parietal; Pm, premaxilla; Po, postorbital; Pof, postfrontal; Prf, prefrontal; Pt, pterygoid; Q, quadrate; Qj,
quadratojugal; Sa, surangular; Sp, splenial; Sq, squamosal; St, supratemporal; T, tabular. Scale bars equal 1 cm.
2
Pelycosauria
Pelycosauria: Varanopseidae
Pe1
• Ophiacodontia : nejprimitivnější
Pelycosauria, 1-2 m, velké rostrum přímá
čelist, spánková jáma malá, tropibasická l.
Pelycosauria: Edaphosauria Pe1 (Am,Eu,As) 1-3 m
Pelycosauria:
Sphenocodontia
Dimetrodon
Pe1
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Pelycosauria:
Sphenocodontia
Dimetrodon
Pe1
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Therapsida
• Extrémně diversifikovaná skupina Pe2- Tr2,
herbivoři i karnivoři , lebka 3 - 100 cm
• Afrika (Karoo formation), J Rusko, Čína
Autapomorphies of Therapsida include:
Interpterygoid vacuity short (less than 20 percent of the length of the palate,
measured from its anterior end to the posterior edge of the transverse flange of the
pterygoid).
. In therapsids, the braincase remains firmly sutured to the dermal skull elements.
No more than twelve upper postcaniniform teeth). Other synapsids have at least
thirteen postcaniniform teeth.
Ectopterygoid teeth absent. Other synapsids have small teeth (denticles) on the
ectopterygoid.
Tetraceratops
Jediný therapsid z Pe1
This list of autapomorphies takes into consideration the recent discovery of Tetraceratops, the oldest known
therapsid (Laurin and Reisz, 1990, 1996). Tetraceratops lacks many derived characters present in other
therapsids. Therefore, this list differs from other published lists of autapomorphies of therapsids (Hopson and
Barghusen, 1986; Gauthier et al., 1988; Kemp, 1988; Hopson, 1991).
Tetraceratops
Therapsida: Dinocephalia
Velká mozkovna (squamosum,
masivní artic/sq/quad kloub
etc., vzpřímená posice
Biarmosuchia: basální
Therapsida
Proburnetia
Deuterosauridae
(dravci),
Moschopidae (býl.
Karoo),
Tapinocephalidae
5
Therapsida: Titanosuchia
• Dinocephalia
• Značný rozvoj dentale - Heterodontní
unicuspidní zuby C
• Druhotné patro dobře vyvinuto (patr. zuby)
• Sklerotikální prstenec oka
• Moderní autopodium (2-3-3-3-3 články)
• Velcí až 4-5 m
• Anomodontia
Vysoce
specialisovaní
býložravci,
40 rodů nejúspěšnější
therapsidi (zejm.
skupina
Dicynodontia) -
Therapsida: Dicynodontia
Therapsida: Dicynodontia
Absence zubů, zobáky,
monokondylní
tropibasická lebka,
masivní pletence,
lumb.žebra etc.
Vesměs velcí býložravci, vč.
vodní (jako želvy)
6
Therapsida: Theriodontia
• Gorgonopsia
• Therocephalia
• Cynodontia:
– Cynognathidae, Diademodontoidea,
Probainognathus, Trithelodontia,
Tritylodontia, Mammala
Therapsida: Theriodonta - Gorgonopsia
• Vysoké končetiny
• Caninisovaná dentice (u některých extrémně
vč. redukce postcaninů - Smilesaurus,
Scymnognathus), vysoké rostrum, prostorná
nosní dutina
• Pohybově aktivní dravci - cf. psi, Ca 1-2 m ,
Gorgonopsia
7
Therapsida: Theriodonta - Cynodontia
• Drobní, výrazně heterodontní dentice, I, C,
dominují postcaniny - vícehroté, dokonalé
druh.patro, uzavřená štěrbina, for. incisivum
prox.,
• Terminální posice nozder
• Efektivní čelistní kloub - válcovité quadratum,
masivní osvalení,
• Značný rozvoj dentale - proc.coronoideus,
• Redukce lumb. žeber (?živorodost)
Nejstarší Cynodontia
(Pe3): Procynosuchus
8
Therapsida: Theriodonta - Cynodontia
Diademontoidea:
Větší herbivorní
formy
Diademodon
Ca 25 cm
Morganucodon
Cynodontia: Probainognathus
Probainognathus
Cynodontia:
Tritylodonta
= Tritylodontidae
Vysoce
specialisovaní
herbivoři
9
Therapsida: Theriodonta - Tritylodontia
Tritylodonta:
Tritylodonta
Ictidosauria
Oligokyphus squam-dent
kloub, zuby!
Srv.: Tritylodonta
Cynodontia:
Trithelodontia
Diarthrognatha:
Diarthrognathus - oba
klouby
10
dtto
Diarthrognathus
MAMMALIA !
Adelobasileus
Tr3 Texas
Sinoconodon
Docodonta / Morganucodonta:
MAMMALIA !
•
•
•
•
•
Morganucodon watsoni Tr3 Rhaet (lom Duchy, GB)
drobný
Postcaniny s 3 seriálními hroty
5I, 5 molariformů, premoláry
Mandibulární zářez s foramen mandibulae
(chrupavčité articulare)
• Končetiny savčí, pletence pokročilé
Morganucodon
Celkem ca 8 lokalit a rodů
UK, EU(Portugalsko)
USA, SAf., SAm
11
Docodonta
Middle Jurassic
Jiulongshan Formation of
the Inner Mongolia
Region, dated
approximately 164 million
years ago, Castorocauda is
the earliest-known
mammal that had
specialized skeletal and
soft-tissue features for
swimming and teeth for
eating fish.
Triconodonta (Jura)
• Heterodontní chrup: trituberkulární
postcaniny, velký P4/4, : střižný efekt
(alterace max/md. cuspů)
• fosa temporalis na ramus mand.,
• Mastikace moderní: m.temporalis,
m.masseter, m.pterygoideus
• Vnitřní mandibuální zářez jen nepatrný
• Etmoturbinalia, cribriformní destička
12
Pantotheria:
Symmetrodonta (např. Amphidontidae, Spalacotheriidae)
Eupantotheria: Amphitheriidae, Dryolestidae,
Aegialodontidae etc.
•
savci s triangulárním
uspořádáním cuspů na
postcaninech,
•
První již s
Morganucodonem
Jura 1: Kuehnotheriidae
(Symmetrodonta s.str.
nebo předkovská skupina
Pantotheria, sesterská
Morganucodon?)
Haramyidae - Jura 1
• Vícehrbolkaté stoličky s
centrální fossou - pouze
isol. zuby
Haramyidae=Microlestidae
=Microleptidae (G, CH,
GB)
Multituberculata
•
•
•
•
•
Vysoce specialisovaná dentice a čelistní aparát:
Veliké premaxily s 1-3 I (I2/ největší), diastema
Multituberkulátní zuby (6-3/5-3)
Chybí septomaxila
Distální extense maxil, palatinum vytěsněno do
středu patra, nevstupuje do orbity
• Převaha plasiomorfií (Mammalia i Pelycosauria)
např. tyčkovitá cochlea, ?absence vnějšího
zvukovodu, ale
• 3 ušní kůstky
• Ptilodus
13
• Multitubercuata
• vs.
• Gondwanatheria
• Pal/Cr Patagonie,
Indie, Madagascar,
Tanzanie
Eupantotheria
• Dryolestidae - drobní
omnivoři J-Cr
• Peramuridae
– Cr
14
C Morganucodon, A Monotremata, B Theria
Didelphis
Pila praeotica
a volné cavum epiptericum
Ornithorhynchus
Mammalia: pila preaotica chybí (dtto u Morganucodon),
cavum epiptericum překryto ectotympanicum
U Monotremata - embryonálně
Trithelodonta
Morganucodon
15
Yanoconodon allini
sheds new light on
the
evolution of the
mammalian ear.
March 15, 2007
A new eutriconodont mammal and evolutionary development in early mammals.
Luo Z-X (1,2), Chen P. (3), Li G. (3) and Chen M. (2), (2007) Nature 446:288-293.
Abstract: Detachment of the three tiny middle ear bones from the reptilian mandible is an important innovation of modern mammals. Here we
describe a Mesozoic eutriconodont nested within crown mammals that clearly illustrates this transition: the middle ear bones are connected to the
mandible via an ossified Meckel's cartilage. The connected ear and jaw structure is similar to the embryonic pattern in modern monotremes (egglaying mammals) and placental mammals, but is a paedomorphic feature retained in the adult, unlike in monotreme and placental adults. This
suggests that reversal to (or retention of) this premammalian ancestral condition is correlated with different developmental timing (heterochrony)
in eutriconodonts. This new eutriconodont adds to the evidence of homoplasy of vertebral characters in the thoraco-lumbar transition and unfused
lumbar ribs among early mammals. This is similar to the effect of homeobox gene patterning of vertebrae in modern mammals, making it plausible
to extrapolate the effects of Hox gene patterning to account for homoplastic evolution of vertebral characters in early mammals.
Published in Nature
16
Závěr
• Jednotlivé savčí znaky se objevují ve
fylogenesi nezávisle
• Typická je mozaika odvozeného stavu
některých znaků a ancestrálního stavu u
znaků jiných
• Srv. Prototheria
Změny mastikace
17
Cope-Osborn
tuberculosectoriální +
trituberkulátní
= tribosfénický molár
Simpson (1936)
Klasický obraz časné evoluce savců
savců:
e.g. Crompton 1971
• Tr3/Jura1: Non-mammalian Pantotheria
(Morganucodon) vs.
• Direct mammalian ancestors: Kuehneotherium
• Jura 3: pretribosphenic Peramus
• Cr 1: tribosphenic Aegialodon
• Masivní žebra a obratle - savčí pohyb. modus
• Cr 2/3: nejstarší doklady Theria s.str.:
Deltatheridium , Holoclemensia, Prokennalestes
• souběžně: Multituberculata a Monotremata
• Takřka všechny doklady ze severních kontinentů:
evoluce savců na severu
18
Obdurodon discksoni
Mio, Riversleigh Formation
Od konce 80.let – explose nových nálezů
a názorových přestaveb
• Zuby,
• Velký, přímé
rostrum,
• Mohutné
septomaxily,
oddělené od
premaxil (srostlé u
Ornithorhynchus
• Shuotherium Chow et Rich, 1982 (Jura3, Čína a Anglie:
Sogogneau-Russell 1998): alternativní modelace
tribosfenického vzoru, resp. tribosfenické okluse:
„talonid“ před trigonidem
• Steropodon Archer et al., 1985 (Cr3: Australia)
• ------------------------------ pod čarou:
•
Ornithorhynchus
anatinus,
Souběžně: změna taxonomické dikce (i klasifikačního usu cf.
kladistické sekvencování apod.): vyprazdňování náplně
nominálních vyšších taxonů – konkretisace profesionálních
diskusí důslednou citací jednotlivých forem (druhů)
Recent
●
;
Od přelomu 80. a 90. let - doklady savců z jižních
kontinentů
Vincelestes Bonaparte, 1986 (Cr1, Patagonie)
Ausktribosphenos, Bishops Rich et al. 1997,1999 (Cr1
Australia)
1999: Ambodro mahabo Flynn et al. - Jura2, Madagascar
Ambodro – dokládá velmi
časný vznik tribosfenické
organisace, posouvá
počátek vlastních savců
„Tribosphenida“ do Jury – a
do Gondwany !
19
Luo, Cifelli et Kielan-Jaworowska 2001:
Dual origin of tribosphenic mammals
• Australosphenida: Monotremata (incl. Steropodon),
Ausktribospehnida, Ambodro
- apomorfie: spojitý
cingulární záhyb na linguální straně spodních stoliček,
poslední premolár s trigonidem (tj. 5 M!?) - dtto u
Obdurodon, plesiomorf: postdentární zářez (u
Ausktribosphenos)
• Boreosphenida (incl. Tribosphenida McKenna, 1975):
nemají mesiální ani linguální cingulum na M/x, ale
cingulární cuspidy, etc. Biogrogr.: Cr1-N-hem., Cr3migrace do S Am, Indie a Af
Asfaltomylos Rauhut et al. 2002 (Jura, Patagonie)
Australosphenida : monophylum +
Shuotherium
Cr 1 Yixian
Formation,
China, 125
My BP
Spodní Jura 195 My (Lufeng Prov,
SW China): nejmenší druhohorní
savec, extrémně velký mozek
Hadroconium wui Luo,
Crompton, Sue, 2001
• 1999: Jeholodens
• (Eutriconodonta)
http://www.digimorph.org/specimens/Hadro
codium_wui/
20
Cr 1 Yixian Formation, China, 125 My BP
• Nejstarší doklad
EUTHERIA:
• Eomaia scansoria Ji et
al., 2002
Cr1 Čína 125 My ago (tj.
50 My před jinými Euth.
Úprava autopodia ukazuje na šplhání a stromový způsob života
..nejstarší vačnatec
An Early Cretaceous Tribosphenic Mammal and Metatherian Evolution :Yixian
• Yixiang 2004: Repenomamus
1 m - predátor juv. dinosaurů
(Psittacosaurus)
Derived features of a new boreosphenidan mammal from the Lower Cretaceous Yixian Formation of China suggest that it
has a closer relationship to metatherians (including extant marsupials) than to eutherians (including extant placentals). This
fossil dates to 125 million years ago and extends the record of marsupial relatives with skeletal remains by 50 million years.
It also has many foot structures known only from climbing and tree-living extant mammals, suggesting that early crown
therians exploited diverse niches. New data from this fossil support the view that Asia was likely the center for the
diversification of the earliest metatherians and eutherians during the Early Cretaceous.
Sinodelphys szalayi is more closely related to extant marsupials than to extant placentals
and stem taxa of boreosphenidans in its many marsupial-like apomorphies in the skeleton
and anterior dentition
21
Earliest Known Gliding Mammal,Volaticotherium antiqus
2006 China: Inner
Mongolia (Cr ?)
PROTOTHERIA:
Monotremata
3 sluchové
kůstky,
tribosfénický
molár
Cochlea spiralní
Ukončený růst
lebky
22