platyognathus is an extinct genus of protosuchian crocodylomorph .

fossils are known from the early jurassic lower lufeng formation in yunnan , china and belong to the type and only species , p. hsui .

platyognathus

the dentition of the mesosuchian is similar to that of
platyognathus
and like
platyognathus
, the dentary is laterally expanded .
how can i put and write and define platyognathus in a sentence and how is the word platyognathus used in a sentence and examples ? 用platyognathus造句 , 用platyognathus造句 , 用platyognathus造句 , platyognathus meaning , definition , pronunciation , synonyms and example sentences are provided by ichacha . net .
because
platyognathus
is known from such incomplete material , there is yet to be an adequate cladistic analyses that can determine a phylogenetic position for
platyognathus
within protosuchia .
because
platyognathus
differed from any other known pseudosuchian , young suggested that it may belong to its own family .
other lufeng crocodylomorphs found in association with
strigosuchus
include the protosuchian
platyognathus
and the sphenosuchian
dibothrosuchus
.
the new skull revealed an additional diagnostic feature of
platyognathus
that distinguishes it from other protosuchians : the distinctive curvature of the jugal .
an unnamed mesosuchian from the salt wash member of the late jurassic morrison formation of western colorado in the united states possesses several features that are also seen in
platyognathus
.
because the infratemporal fenestra of
platyognathus
is very small and the mandibular symphysis extends posteriorly to the level of the seventh to ninth tooth , the genus is thought to be more derived than
orthosuchus
.
the authors of the description of the neotype suggested that
platyognathus
may be closely related to the clade containing protosuchidae and
shantungosuchus
because it possesses a dentary tooth that fits into a notch between the premaxilla and maxilla .
the jaw of
protosuchus richardsoni
, described by barnum brown in 1933 from arizona , is similar in size to that of
platyognathus
, although it lacks many of the distinguishing features of the chinese genus as described by young .
however , later studies revealed that the specimens had only two rows of osteoderms that more closely resembled those of the genus
platyognathus
, and that the presence of three rows of osteoderms in the holotype of
microchampsa
may have been an artifact of preservation .
the shortness of the snout relative to the rest of the skull and the presence of a laterally and ventrally open notch between the premaxilla and maxilla indicate that
platyognathus
is a member of protosuchia as defined by wu
et al .
( 1994 ) .
the classification of the genus continued to be debated , as descriptions of the fragmentary jaw were all that was available to study .
platyognathus
continued to be referred to pseudosuchia in the following years , and was suggested to have a close relationship with
sphenosuchus
.
based on these features , young found no close similarities between the jaw of
platyognathus
and any other known early crocodylomorph such as
notochampsa
,
sphenosuchus
,
erythrochampsa
, and
pedeticosaurus
( at the time , these crocodylomorphs were collectively known as the
pseudosuchia
) .
excluded taxa historically referred to euparkeriidae . dorosuchus neoetus sennikov , 1989 [ 63 ] , ‘ wangisuchus tzeyii ’ young , 1964 [ 64 ] , turfanosuchus dabanensis young , 1973 [ 65 ] , xilousuchus sapingensis wu , 1981 [ 66 ] , platyognathus hsui sennikov , 1989 [ 67 ] , dongusia colorata huene , 1940 [ 68 ] .
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full reference : c . - c . young . 1944 . on a supposed new pseudosuchian from upper triassic saurischian - bearing red beds of lufeng , yunnan , china . american museum novitates 1264 : 1 - 4
parent taxon : crocodyliformes according to r . b . irmis et al . 2013
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locality : huangchiaten , about 1 km north of sliawan , lufeng , yunnan province , china .
locality : about 10 m west of the intersection of the roads from zhangjiawa to dadi and from dawa elementary school to dahuangtian , yunnan province , china .
horizon : stratum 6 ( middle of dark red beds ) of lower lufeng formation .
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( meaning
chinese lizard
) was a tetanuran theropod dinosaur which lived during the early jurassic period . its fossils were found in the lufeng formation , in yunnan province , china . it was a bipedal carnivore . there is currently insufficient information available to reliably estimate its dimensions .
( σαυρος ) meaning
lizard
; thus
chinese lizard
. the specific name ,
, refers to the triassic , the period that the fossils were originally thought to date from .
was described and named by chung chien young , who is known as the ' father of chinese vertebrate paleontology ' , in 1948 .
, was found in the lower lufeng formation , and consists of a maxilla ( upper jaw ) , a lower jaw fragment and several teeth . the teeth are laterally compressed , and feature fine serrations both at their anterior and posterior edges . the teeth are also variable in size and are curved backwards . this material is too fragmentary to determine the length and weight of this dinosaur . over the years , other fossils were referred to
( 2013 ) examined the effect of the traumatic loss of teeth on the dental alveolus ( the socket in the jaw where the roots of teeth are held ) in dinosaurs .
is the first dinosaur where remodeling of the alveolus in the jaw was observed . the authors concluded that this finding
contributes to mounting evidence suggesting theropods were highly resilient to a broad spectrum of traumas and diseases .
, can be distinguished based on the fact that a vertical groove is present on the lateral premaxilla adjacent to contact with the maxilla .
, based on the fact that the anterior end of the jugal does not participate in the internal antorbital fenestra and that the maxillary tooth row is completely in front of the eye socket .
, features elsewhere in the skeleton suggest it is closer to tetanuran theropods . rauhut ( 2003 ) regarded
( zljt01 ) in 2007 from the lufeng basin . it consists of an incomplete skull and other postcranial fragments . phylogenetic analysis of this specimen , demonstrates that
ivpp v34 was recovered in the zhangjiawa member of the lufeng formation , in yunnan , china . these remains were discovered at the dark red beds that were deposited during the sinemurian stage of the jurassic period , approximately 196 - 183 million years ago . several other discoveries referred to
were made in the zhangjiawa member : specimens ivpp v97 ( postcrania ) , ivpp v36 ( teeth ) , ivpp 37 ( teeth ) , ivpp v88 ( ilium ) , ivpp v35 ( teeth and postcranial bones ) , ivpp v100 and ivpp v48 ( teeth and postcranial bones ) discovered in 1938 by m . bien & c . c . young , fmnh cup 2001 - 2003 discovered by e . oehler and hu . specimens fmnh cup 2097 , fmnh cup 2098 , fmnh cup 2004 , fmnh cup 2005 were discovered in 1948 by m . bien & c . c . young at zhangjiawa member , as well .
, a maxilla with four teeth , was collected by lee in the 1940s , in the dull purplish beds of shawan member of the lufeng formation , that were deposited during the hettangian stage of the jurassic period , approximately 201 - 199 million years ago . several other discoveries were made in the shawan member : parts of two skeletons attributed to
were discovered by sou in 1956 , specimen ivpp v279 ( tooth ) was discovered by c . c . young in 1938 , in dark red clayish sandstone , and specimen ivpp v381 ( several teeth ) was discovered by c . c . young , in blue mudstone . the
, were recovered in the shawan member of lufeng formation . this material was discovered in 1987 in the dull purplish beds that were deposited during the hettangian stage of the early jurassic , approximately 201 - 199 million years ago .
, and other early reptiles . other dinosaurs discovered included indeterminate remains of sauropods , and the early armored dinosaurs
sinosaurus had a deep notch at the suture between premaxillary and maxillary bones . this notch was very powerfully built , possibly to anchor muscles that would produce a powerful bite . sinosaurus might have been either a scavenger or a predator . the premaxilla was probably covered with the narrow , hooked beak used to tear skins and abdominal walls of poor victims . a crested crown , which is sometimes fan - shaped , may have been helpful to keep the abdominal wall open while devouring . the feet of sinosaurus were similar to those of himalayan vultures , and had become adapted to eating the corpses of large animals , such as prosauropods .
for the first time , a phylogenetic analysis including all putative euparkeriid taxa is conducted , using a large data matrix analysed with maximum parsimony and bayesian analysis . using parsimony , the putative euparkeriid dorosuchus neoetus from russia is the sister taxon to archosauria + phytosauria . euparkeria capensis is placed one node further from the crown , and forms a euparkeriid clade with the chinese taxa halazhaisuchus qiaoensis and ‘ turfanosuchus shageduensis ’ and the polish taxon osmolskina czatkowicensis . using bayesian methods , osmolskina and halazhaisuchus are sister taxa within euparkeriidae , in turn sister to ‘ turfanosuchus shageduensis ’ and then euparkeria capensis . dorosuchus is placed in a polytomy with euparkeriidae and archosauria + phytosauria . although conclusions remain tentative owing to low node support and incompleteness , a broad phylogenetic position close to the base of archosauria is confirmed for all putative euparkeriids , and the ancestor of archosauria + phytosauria is optimized as similar to euparkeriids in its morphology . ecomorphological characters and traits are optimized onto the maximum parsimony strict consensus phylogeny presented using squared change parsimony . this optimization indicates that the ancestral archosaur was probably similar in many respects to euparkeriids , being relatively small , terrestrial , carnivorous and showing relatively cursorial limb morphology ; this bauplan may have underlain the exceptional radiaton and success of crown archosauria .
archosauria , the diapsid clade represented today by birds and crocodilians and including the extinct dinosaurs , is highly speciose ( with over 9000 species of extant bird and crocodilian [ 1 ] ) . archosaurs filled most major terrestrial ecological niches for over 150 million years [ 2 – 7 ] , from the middle triassic to the end of the cretaceous . the ‘rise’ of the archosaurs to this position of ecological dominance took place coincident with the extinction or decline of many therapsid taxa at the end of the permian and during the course of the early–middle triassic , which had previously filled most large - bodied terrestrial niches [ 2 , 3 , 8 – 14 ] . the rise of the archosaurs is one of the landmark terrestrial faunal transitions and is an outstanding example of a large - scale adaptive radiation in the fossil record [ 2 , 15 , 16 ] , with archosaurs diversifying into carnivorous , herbivorous , aquatic , terrestrial and volant forms of greatly varying sizes [ 2 , 14 – 17 ] .
previous phylogenetic positions found for euparkeria capensis . ( a ) [ 20 ] ; ( b ) [ 21 ] ; ( c ) [ 22 ] and ( d ) [ 23 ] .
most recent work has placed euparkeria capensis and other euparkeriids close to the base of , but outside , archosauria ( see for example the summaries in [ 32 , 33 ] , the placement hypothesized by borsuk - białynicka & evans [ 30 ] , and the phylogenetic analyses of benton & clark [ 34 ] , sereno & arcucci [ 35 ] , sereno [ 36 ] , juul [ 37 ] , bennett [ 38 ] , benton [ 39 ] , parker & barton [ 40 ] , nesbitt et al . [ 41 ] , dilkes & sues [ 21 ] , nesbitt [ 23 ] , brusatte et al . [ 2 ] , ezcurra et al . [ 22 ] , desojo et al . [ 42 ] , dilkes & arcucci [ 43 ] , schoch & sues [ 44 ] , sookias et al . [ 28 , 31 ] , parrish [ 45 ] ; figure 1 b – d ) . this contrasts with the placement of euparkeria capensis within the crown , as the sister taxon to ornithosuchidae + ornithodira ( within ‘ornithosuchia’ ) , found in an early analysis by gauthier [ 20 ] ( figure 1 a ) .
less consensus has been reached regarding the relationships between euparkeriids and other stem archosaurs . several analyses have placed euparkeria capensis as the sister taxon to archosauria [ 34 , 38 , 41 , 45 ] or phytosauria + archosauria [ 23 , 28 , 31 ] . however , many previous analyses have also placed proterochampsids , or proterochampsids + doswelliids , closer to the crown than is euparkeria capensis [ 2 , 22 , 35 – 37 , 42 ] ( benton [ 39 ] presents this relationship , but the opposite topology is no less parsimonious ; figure 1 b , c ) .
all previous phylogenetic work has placed doswelliids more crownward than euparkeriids ( [ 21 , 22 , 40 , 42 , 44 ] ; figure 1 b , c ) , either as the sister taxon to ( [ 21 , 44 ] when excluding tarjadia ruthae ; figure 1 b ) , in a polytomy with ( [ 40 , 44 ] when including tarjadia ruthae ) , or more derived than [ 22 , 42 ] ( figure 1 d ) proterochampsids .
the analysis of dilkes & sues [ 21 ] ( figure 1 b ) was unique in placing erythrosuchus africanus closer to the crown than euparkeria capensis , whereas ezcurra et al . [ 22 ] placed erythrosuchids closer to the crown than osmolskina czatkowicensis . all other phylogenetic analyses to date [ 22 , 23 , 28 , 31 , 40 – 42 , 44 , 46 ] have placed euparkeria capensis and other euparkeriids or taxa until recently considered euparkeriids ( namely dorosuchus neoetus , halazhaisuchus qiaoensis , ‘ turfanosuchus shageduensis ’ ) crownwards of erythrosuchids , including those expanding the dataset of dilkes & sues [ 21 ] ( e . g . [ 22 , 42 ] ) , and several questions have been raised regarding the validity of some of the scorings in the matrix of dilkes & sues [ 21 ] , including by the first author [ 43 ] .
although the referral of specimens to the hypodigm of dorosuchus neoetus remains tentative , this analysis included all referred material as a single terminal taxon following sookias et al . [ 31 ] . similarly , osmolskina czatkowicensis was scored as a single terminal taxon based on all referred material ; the holotype consists only of an anterior maxilla [ 29 ] , meaning analysing it alone is unlikely to be informative , and there is no reason to doubt the referral of any of the referred elements more than that of any others , making analysing subcombinations of material similarly uninformative . it must be noted strongly , however , that referral of all material other than the maxilla to osmolskina czatkowicensis remains extremely uncertain and thus do conclusions based on the referred material of this taxon ; no autamorphic euparkeriid characters unite the referred material , and at least one other archosauriform is found in the same assemblage .
all characters of butler et al . [ 46 ] which ceased to be phylogenetically informative ( i . e . all taxa , or all taxa bar one , were scored identically ) following taxon pruning were removed , along with characters 100 , 156 and 157 of nesbitt [ 23 ] because it was considered that the variation referred to in their states could not be reliably observed ( see [ 31 ] ) . an additional 50 characters from other datasets were added , including characters from dilkes [ 49 ] , ezcurra [ 22 ] , desojo et al . [ 42 ] , dilkes & arcucci [ 43 ] and ezcurra et al . [ 50 ] ( see electronic supplementary material ) . five new characters were added based on personal observations , yielding a total of 405 characters . these new characters , their distribution and delineation are outlined in the electronic supplementary material , also archived on dryad .
the main analysis was however conducted excluding one of these characters ( character 93 in the dataset ) . this character was based on the observations of borsuk - białynicka & evans [ 30 ] for osmolskina czatkowicensis ( ‘pterygoid , ridge along posteromedial corner , separating palatal flange from neck’ ) . this character is potentially of particular relevance to euparkeriid phylogeny , as it was identified as a synapomorphy uniting euparkeria capensis , osmolskina czatkowicensis and other archosauromorph taxa ; however , the state delimitation and scoring for this character were found to be very challenging , so it was not included in the main phylogenetic analysis presented .
] , which were not included in the dataset . the analysis used four chains sampled every 100 generations , with the analysis set to stop once it reached a standard deviation of split frequencies of 0 . 01 . the first quarter of the generations were discarded as burn in .
a reduced consensus analysis failed to improve resolution of the tree , and is thus not presented . although majority rule consensus trees did yield fewer polytomies than a strict consensus analysis , given that the strict consensus was relatively well resolved , that a strict consensus is the most conservative approach , and that the cut - off point in a majority rule consensus is somewhat arbitrary , and given the fact that a majority rule excludes a number of relationships which are equally parsimonious to those seen in the tree , these were not presented . the number of phylogenetic characters in the dataset characterizing fully quantitative variation was deemed insufficient to warrant quantitative delineation of character states using methods such as gap weighting [ 55 ] .
to examine the likely morphology of the common ancestor of archosauria and phytosauria , character states were optimized using squared change parsimony onto the parsimony phylogeny presented here , including two additional ecological descriptors—femoral length ( as a proxy for body size ) , and terrestriality ( whether generally considered to have been terrestrial , aquatic or volant ) .
bp , evolutionary studies institute ( formerly bernard price institute ) , university of the witwatersrand , johannesburg , south africa .
ifgt , institute for geosciences , eberhard - karls - universität tübingen , tübingen , germany ( formerly geologisch - paläontologisches institut tübingen , gpit ) .
macn - pv , museo argentino de ciencias naturales ‘bernardino rivadavia’ , paleontología de vertebrados , buenos aires , argentina .
pin , paleontological institute of the russian academy of sciences , moscow , russia .
pulr , universidad nacional de la rioja , paleontología , la rioja , argentina .
pvl , instituto miguel lillo , universidad nacional de tucumán , tucumán , argentina .
pvsj , división de paleontología , museo de ciencias naturales de la universidad nacional de san juan , argentina .
umzc , university museum of zoology , university of cambridge , cambridge , uk .
zpal , institute of paleobiology , polish academy of sciences , warsaw , poland .
the parsimony analysis yielded 16 most parsimonious trees of 1330 steps , with a consistency index of 0 . 358 and a retention index of 0 . 688 . dorosuchus neoetus was the sister taxon to archosauria + phytosauria , and all other euparkeriids formed a clade which was the sister taxon to dorosuchus neoetus + ( archosauria + phytosauria ) . within the euparkeriid clade , halazhaisuchus qiaoensis , ‘ turfanosuchus shageduensis ’ and osmolskina czatkowicensis formed a subclade to the exclusion of euparkeria capensis . one additional step was required to place dorosuchus neoetus as the sister taxon to the other euparkeriids . character states optimizing as autapomorphies for each taxon and synapomorphies supporting euparkeriidae , the subclade within euparkeriidae , and the position of dorosuchus neoetus are listed in table 1 , and a full synapomorphy list for all clades is given in the electronic supplementary material .
synapomorphies supporting key clades and autapormorphies of euparkeriid taxa . numbers refer to character numbers and state changes .
strict consensus tree of 16 most parsimonious trees , showing the position of all putative euparkeriid taxa ( bold ) . numbers at nodes are standard bootstrap values ( before first slash ) , gc bootstrap values ( after first slash ; only given if different from standard bootstrap value ) and decay ( = bremer ) indices ( after second slash ; only given if greatet than 1 ) .
the bayesian analysis yielded a topology very similar to the strict consensus tree of the parsimony analysis , with the positions of and relationships among euparkeriids differing only in that halazhaisuchus qiaoensis and osmolskina czatkowicensis were found to be sister taxa to the exclusion of ‘ turfanosuchus shageduensis ’ , and that dorosuchus neoetus was placed in a polytomy with euparkeriidae and phytosauria + archosauria ( figure 3 ) rather than as the sister taxon to the latter clade . the bayesian analysis showed two more polytomies than the parsimony analysis , with the polytomies being larger , including several clades ( table 2 ) .
bayesian consensus tree , showing the position of all putative euparkeriid taxa ( bold ) . numbers at nodes are posterior probabilities .
a total of 575 000 generations were required to reach a standard deviation of split frequencies of 0 . 01 . the posterior probability of a monophyletic euparkeriidae was low ( 0 . 61 ) , as were the posterior probabilities of the relationships within euparkeriidae ( less than or equal to 0 . 60 ) , of a monophyletic phytosauria + archosauria to the exclusion of dorosuchus neoetus ( 0 . 67 ) , and of the clade ( euparkeriidae , dorosuchus neoetus ( phytosauria + archosauria ) ) to the exclusion of proterochampsids and other stem taxa ( 0 . 62 ) .
based on the results of the optimization , the ancestor of crown archosauria and phytosauria was a relatively small ( 215 mm estimated femoral length ) , cursorial and gracile ( e . g . fourth trochanter present—character 310 , state 1—indicating relatively upright locomotion ; femoral distal condyles not projecting markedly beyond shaft—character 313 , state 1—contrasting with the robust femora of erythrosuchids and proterosuchids ) , terrestrial ( optimized as terrestrial , and lacking pit and ridge dermal ornamentation—character 33 , state 0—which is typical of aquatic taxa ) and carnivorous ( possessing serrated , bladelike , mediolaterally compressed marginal teeth—character 160 , state 1 , character 164 , state 0 and character 165 , state 1 , respectively ) animal ( see electronic supplementary material , figures s3–10 showing optimizations ) .
archosauromorpha huene , 1946 [ 57 ] sensu gauthier et al . , 1988 [ 58 ] .
archosauriformes gauthier et al . , 1988 [ 58 ] sensu nesbitt , 2011 [ 23 ] .
euparkeriidae huene , 1920 [ 59 ] sensu sookias & butler , 2013 [ 19 ] .
phylogenetic definition . stem - based definition—the most inclusive clade containing euparkeria capensis broom , 1913 [ 26 ] but not crocodylus niloticus laurenti , 1768 [ 60 ] or passer domesticus linnaeus , 1758 [ 61 ] ( from sookias & butler [ 19 ] ) .
included taxa and specimens . euparkeria capensis broom , 1913 [ 26 ] ( = browniella africana [ 27 ] ) , halazhaisuchus qiaoensis wu , 1982 [ 62 ] , osmolskina czatkowicensis borsuk - białynicka & evans , 2003 [ 29 ] , holotype of ‘ turfanosuchus shageduensis ’ wu , 1982 [ 62 ] .
distribution . late early triassic ( late olenekian ) of poland , early middle triassic ( early anisian ) of south africa , early or middle triassic ( late olenekian or early anisian ) of china .
diagnostic synapomorphies . non - crown archosauriforms with the following local synapomorphies : four premaxillary teeth ; two foramina for hypoglossal nerve on exoccipital ; vertically orientated parabasisphenoid ; discrete interparietals ( = postparietals ) ; two rows of keeled dorsal paramedian osteoderms with distinct anterior point / process ( all differentiating euparkeriids from many stem taxa and some crown taxa ) .
further differential diagnosis . absence of ossified astragalocalcaneal canal ( differentiating euparkeriids from non - archosauriform archosauromorphs and proterosuchids ) ; large anterolaterally opening foramen on anterolateral surface of maxilla at base of anterodorsal process ; calcaneal tuber deflected approximately between 20° and 50° posteriorly ; incompletely ossified medial wall of vestibule ( = otic capsule ) ; field of pointed , recurved teeth on pterygoid ( all differentiating euparkeriids primarily from crown taxa ) .
comments . osmolskina czatkowicensis has four premaxillary teeth [ 29 , 30 ] and , contra nesbitt [ 23 ] , euparkeria capensis has four , not three , premaxillary teeth ( figure 4 ) . five premaxillary teeth are seen in many stem archosaurs including erythrosuchids [ 69 ] , vancleavea campi [ 41 ] , proterochampsids [ 23 , 70 ] and in several pseudosuchian taxa ( e . g . turfanosuchus dabanensis [ 71 ] , xilousuchus sapingensis [ 66 ] ) . proterosuchids ( e . g . proterosuchus goweri —nmqr 880 ) , phytosaurs ( e . g . smilosuchus gregorii —ucmp 27200 ) , and other taxa with elongated premaxillae ( e . g . qianosuchus mixtus —ivpp v 13899 [ 23 ] ) have more than five premaxillary teeth , whereas some taxa lack premaxillary teeth entirely ( effigia okeeffeae [ 72 ] ) . a large number of crown archosaurs show four premaxillary teeth [ 23 ] ( e . g . batrachotomus kupferzellensis [ 73 ] , herrerasaurus ischigualastensis [ 74 ] ) .
euparkeria capensis sam - pk - 6047a premaxillary teeth as ( a ) ct slice and ( b ) volume reconstruction from ct data . t , tooth .
presence of two foramina for the hypoglossal nerve differentiates euparkeriids from erythrosuchids [ 75 ] , proterochampsids [ 43 ] , doswelliids [ 42 ] and many crown taxa ( e . g . batrachotomus kupferzellensis [ 76 ] , arizonasaurus babbitti [ 77 ] ) , all of which possess one . a vertically orientated parabasisphenoid is seen in most crown archosaurs [ 23 , 78 ] , erythrosuchids [ 72 , 75 ] and proterochampsids [ 23 , 43 ] , whereas that of mesosuchus browni , prolacerta broomi , proterosuchids [ 23 , 78 ] , doswelliids [ 21 , 42 ] , trilophosaurus buettneri [ 79 ] and youngina capensis [ 80 ] is more horizontally orientated .
presence of a separately ossified interparietal in euparkeria capensis ( sam - pk - 5867 ) and osmolskina czatkowicensis ( inferred based on the form of the parietals [ 30 ] ) differentiates euparkeriids from crown taxa ( e . g . batrachotomus kupferzellensis [ 73 ] , herrerasaurus ischigualastensis [ 74 ] ) , proterochampsids ( e . g . chanaresuchus bonapartei [ 70 ] ) and doswelliids ( doswellia kaltenbachi [ 21 ] ) , but is the same condition seen in many stem taxa including erythrosuchids ( e . g . erythrosuchus africanus [ 69 ] ) and proterosuchids ( e . g . proterosuchus fergusi [ 81 ] ) [ 23 ] .
two rows of anteriorly pointed paramedian osteoderms are seen in many crown pseudosuchians ( e . g . ticinosuchus ferox [ 82 ] , rauisuchus tiradentes [ 83 ] ; figure 5 ) . most ornithodirans ( e . g . herrerasaurus ischigualastensis [ 74 ] , silesaurus opolensis [ 84 ] ) and many stem archosaurs ( e . g . erythrosuchids and proterosuchids [ 23 ] —although some controversy remains regarding erythrosuchids as discussed below ; mesosuchus browni [ 49 ] ) lack dorsal osteoderms entirely . the osteoderms of proterochampsids differ in that they are rounded anteriorly , and form a single paramedian row [ 23 , 70 ] , and those of phytosaurs differ in that they are rounded anteriorly and rugose [ 23 ] . the osteoderms of vancleavea campi are similar in shape to those of euparkeriids , but cover much of the body rather than forming only paramedian rows [ 41 ] . a single osteoderm pertaining to koilamasuchus gonzalezdiazi [ 22 ] is broadly similar to those of euparkeriids , but is more rounded anteriorly than those of euparkeria capensis .
osteoderms of euparkeriid taxa compared to other basal archosauriforms . euparkeria capensis umzc t . 692 in ( a ) dorsal and ( b ) ventral view ; euparkeria capensis sam - pk - 6049a in ( c ) dorsal view ; halazhaisuchus qiaoensis ivpp v6027 - 9 in ( d ) dorsal and ( e ) ventral view ; halazhaisuchus qiaoensis ivpp v6027 - 2 in ( f ) dorsal view ; osmolskina czatkowicensis zpal rv / 1339 in ( g ) dorsal and ( h ) ventral views ; osmolskina czatkowicensis zpal rv / 1335 in ( i ) dorsal and ( j ) ventral views ; osmolskina czatkowicensis zpal rv / 1338 in ( k ) dorsal view ; archosauriformes indet . zpal rv / 1337 from czatkowice 1 in ( l ) dorsal and ( m ) ventral views ; different koilamasuchus gonzalezdiazi macn - pv 18119 paramedian osteoderms in ( n , o ) dorsal , and ( p ) anterior or posterior view ; putative osteoderm of erythrosuchus africanus nhmuk pv r3592 in ( q , r ) dorsal or ventral and ( s ) medial / lateral / posterior / anterior view ; rauisuchus tiradentes osteoderms in dorsal view : bspg as xxv cervical osteoderm ( t ) ; bspg as xxv 97 anterior dorsal osteoderm ( u ) ; bspg as xxv 94 posterior dorsal osteoderm ( v ) ; bspg as xxv 121 caudal osteoderms ( w , x ) . image ( k ) courtesy of m . borsuk - białynicka and zpal ; images ( n – p ) courtesy of m . d . ezcurra ; images ( t – x ) courtesy of s . lautenschlager / o . rauhut and the linnean society of london . k , median keel .
non - archosauriform archosauromorphs and proterosuchids possess an ossified astragalocalcaneal canal , which is absent in all other archosauriforms [ 23 , 78 , 85 ] . a large anteriorly opening foramen ( = anterior maxillary foramen of modesto & sues [ 86 ] ) on the lateral surface of the maxilla at the base of the anterodorsal process ( which is definitively homologous with that of euparkeria capensis —see below and [ 23 ] pp . 66–67 ) is absent in all crown taxa except lotosaurus adentus but is present in stem taxa excluding proterochampsids , some erythrosuchids , vancleavea campi [ 23 ] and youngina capensis [ 87 ] . the calcaneal tuber of stem archosaurs except proterochampsids is deflected by less than 20° , whereas that of crown archosaurs is deflected more than 50° posteriorly ; like that of phytosaurs and proterochampsids , the calcaneal tuber of euparkeriids is deflected between 20° and 50° posteriorly ( figure 6 ) .
articulated astragalus and calcaneum of ( a ) euparkeria capensis umzc t . 692 and ( b ) osmolskina czatkowicensis ( astragalus zpal rv / 811 , calcaenum zpal rv / 1253 ) in proximal view .
the medial wall of the vestibule in stem archosaurs is incompletely ossified , whereas that in many ( but not all— silesaurus opolensis and plateosaurus engelhardti show partial ossification [ 23 ] ) crown archosaurs is fully ossified [ 23 , 76 ] . all crown taxa lack pointed , recurved pterygoid teeth , though some crown taxa ( e . g . eoraptor lunensis [ 88 ] ) do show blunt pterygoid teeth . erythrosuchus africanus [ 69 ] and potentially herbivorous taxa such as trilophosaurus buettneri [ 89 ] and derived rhynchosaurs such as hyperodapendon gordoni [ 90 ] are the only stem taxa to lack pterygoid teeth entirely , although those of mesosuchus browni are blunt ( sam - pk - 6536 ) .
comments . euparkeriidae was divided into the subfamilies euparkeriinae [ 59 ] and dorosuchinae by sennikov [ 63 , 91 ] . euparkeriinae was composed of euparkeria capensis and browniella africana , but given that browniella africana is considered synonymous with euparkeria capensis this would render euparkeriinae monospecific . following the results of the phylogenetic analysis presented here , dorosuchinae as conceived by sennikov [ 63 , 91 ] is not monophyletic , as halazhaisuchus qiaoensis and ‘ turfanosuchus shageduensis ’ are recovered within euparkeriidae whereas dorosuchus neoetus is not , while the other two proposed members of this subfamily—‘ wangisuchus tzeyii ’ and turfanosuchus dabanensis —are considered a nomen dubium [ 28 ] and a pseudosuchian [ 23 , 46 ] , respectively . thus , employment of these subfamilies is not considered to be useful . a subclade is found within euparkeriidae by the current analysis ( grouping halazhaisuchus qiaoensis and osmolskina czatkowicensis to the exclusion of euparkeria capensis ) , but given the low support and paucity of diagnostic characters for this clade , a clade name is not erected .
occurrence . single accumulation close to aliwal north , eastern cape , south africa , within the burgersdorp formation , beaufort group , within subzone b of the cynognathus assemblage zone ( early middle triassic : early anisian ) .
diagnosis . euparkeriid distinguished from other taxa by the following autapomorphies : low peak on premaxilla projecting into external naris roughly at anteroposterior midpoint of naris , posterodorsal process ( = postnarial process ) of premaxilla primarily vertical , and rounded at its posterodorsal tip . distinguished from osmolskina czatkowicensis by possessing exoccipitals discrete from the opisthotics , a premaxilla lacking any slight overhang / downturn and which has a more vertical posterodorsal process ( approx . 90° versus 50° in osmolskina czatkowicensis ) and is clearly attached by a facetted articulation to the maxilla ( unlike the weak attachment in osmolskina czatkowicensis ) and not separated from it by an additional antorbital space ( unlike in osmolskina czatkowicensis ) , a tapered and posteriorly curved nasal process of the maxilla ( contrasting with the subquadrangular process of osmolskina czatkowicensis ) , and distinguished from osmolskina czatkowicensis and halazhaisuchus qiaoensis in lacking a tuber for the attachment of the m . triceps on the scapula .
synonymy . browniella africana [ 27 ] is considered a junior subjective synonym of euparkeria capensis , following ewer [ 93 ] . no substantial morphological differences can be identified to distinguish the taxa , with the second pubic foramen and thinner , narrower pubis identified in euparkeria capensis by broom [ 27 ] being artefacts of damage ( sam - pk - 5867 ) and also reflecting overall size difference between the specimens . the ventral part of the ischium of the holotype of ‘ browniella africana ’ ( sam - pk - 6047a ) is damaged , and the greater constriction of the ilium near its middle in this taxon than in euparkeria capensis identified by broom [ 27 ] is thus also seemingly due to damage and / or unassessable .
type species . osmolskina czatkowicensis borsuk - białynicka and evans , 2003 [ 29 ] .
holotype . zpal r - i / 77 , anterior maxilla including anterodorsal process .
referred material . catalogued material : 200 cranial and mandibular bones including braincase ; 30 vertebrae from cervical , dorsal , sacral and caudal regions ; five ilia ; 30 limb bones . other referred material : several hundred incomplete bones and teeth .
occurrence . karstic deposits of czatkowice locality , małopolska , poland ( late early triassic : late olenekian ) .
diagnosis ( based on holotype only ) . the holotype maxilla is distinguished from that of euparkeria capensis by possession of a subquadrangular nasal process ( contrasting with the tapered process of euparkeria capensis ) which reaches less far posteriorly and is less posteriorly curved , and a relatively larger anterior maxillary foramen .
further diagnosis based on referred material . distinguished from euparkeria capensis by the features listed in the diagnosis of euparkeria capensis . distinguished from halazhaisuchus qiaoensis and euparkeria capensis by the possession of a relatively less elongated humerus with larger offset between the angles of its proximal and distal ends , and a strongly anterior position of the coracoid foramen . distinguished from halazhaisuchus qiaoensis by possession of a relatively mediolaterally thicker scapular blade and a less pronounced and less regularly circular attachment scar for the m . triceps on the scapula , lacking a clear central depression .
comments . borsuk - białynicka & evans [ 30 ] additionally diagnosed osmolskina czatkowicensis as differing from euparkeria capensis in possessing a mandible that does not increase in depth posteriorly , and a more twisted femur ( 55° versus 32° offset between proximal and distal ends ) . it is considered here however that the mandible of osmolskina czatkowicensis is not sufficiently well known to be certain as to whether it increased in depth posteriorly or not , and the offset between proximal and distal ends of the femur in euparkeria capensis is relatively variable ( e . g . approx . 45° in sam - pk - 6047b and 30° in sam - pk - 5883 ) , whereas that in osmolskina czatkowicensis is often not easily characterized owing to most femora being only partially preserved . thus , these characteristics are excluded from the current diagnosis .
holotype ( from [ 28 ] ) . ivpp v6027 , posterior three cervical and anterior three dorsal vertebrae in articulation with osteoderms and incomplete ribs ( v6027 - 1 ) , seven dorsal vertebrae in articulation with osteoderms ( v6027 - 2 ) , left ( v6027 - 3 ) and right ( v6027 - 4 ) scapulae , left ( v6027 - 3 ) and partial right ( v6027 - 4 ) coracoids , right humerus ( v6027 - 5 ) , ulna ( v6027 - 6 ) and radius ( v6027 - 7 ) , an isolated left cervical rib ( v6027 - 8 ) , and an isolated median osteoderm ( v6027 - 9 ) . all material probably pertains to a single individual .
occurrence . fugu county , shaanxi province , china , within the lower ermaying formation ( lower or middle triassic : late olenekian to early anisian ) .
diagnosis . euparkeriid distinguished from other taxa by the following autapomorphies : ( i ) strongly pronounced tuber on the scapula , for attachment of the scapular head of the m . triceps , that is circular in outline when the scapula is in lateral view , with the apex of the tuber slightly depressed ; ( ii ) pronounced muscle attachment scar on the scapula in the form of a depressed strip on the lateral surface of the blade running from anterodorsal to posteroventral , beginning at an abrupt kink in the anterior margin at around midlength of the blade ( from sookias et al . [ 28 ] ) . similar m . triceps tubera in other taxa differ in shape , with that in osmolskina czatkowicensis being less regularly circular and lacking a central depression , and those in crown archosaurs being teardrop shaped and lacking a depression ( e . g . batrachotomus kupferzellensis [ 92 ] ) . no similar muscle scar is seen in other euparkeriids or early archosauromorphs . also differentiated from other euparkeriids in possessing epipophyses on the cervical vertebrae .
comments . sookias et al . [ 28 , p . 9 ] , further diagnose halazhaisuchus qiaoensis from other stem and crown archosaurs by a unique combination of characters ‘two rows of paramedian scutes with exposed surfaces at least twice as long as wide when articulated , tapering anterior processes and broad , rounded posterior margins , each having a longitudinal keel closer to the medial margin than the lateral one ; large flattened flange projecting from the proximal part of the anterior margin of each cervical rib ; presence of a tuber on the scapula for attachment of the scapular head of the m . triceps ; presence of dorsal intercentra ; epipophyses on cervical vertebrae’ .
holotype ( from [ 28 ] ) . ivpp v6028 , mostly complete right mandible ( v6028 - 1 ) , six cervical vertebrae missing upper neural arches and neural spines ( v6028 - 2 ) , right scapula ( v6028 - 3 ) , coracoid ( v6028 - 3 ) , humerus ( v6028 - 4 ) , radius ( v6028 - 7 / 8 / 9 ; note that the correct subnumbers for the radius , ulna and fibula are uncertain ) , ulna ( v6028 - 7 / 8 / 9 ) , femur ( v6028 - 5 ) , tibia ( v6028 - 6 ) and fibula ( v6028 - 7 / 8 / 9 ) . all material probably pertains to a single individual .
occurrence . jungar banner , nei mongol autonomous region , china , from the lower ermaying formation ( lower or middle triassic : late olenekian or early anisian ) .
comments . found to be undiagnostic beyond archosauriformes indet . [ 28 ] . placed within euparkeriidae in the current phylogenetic analysis and thus potentially remains referable to the clade . shows no scoring differences with halazhaisuchus qiaoensis , but lack of autapomorphies uniting the taxa mean that they cannot be synonymized ( see [ 28 ] ) .
the results of the current analysis support recent work examining euparkeriid phylogeny ( see [ 28 , 31 ] ) , with halazhaisuchus qiaoensis and ‘ turfanosuchus shageduensis ’ forming a clade with euparkeria capensis and osmolskina czatkowicensis , whereas dorosuchus neoetus is alternatively placed outside euparkeriidae as the sister taxon to phytosauria + archosauria or in a polytomy with euparkeriidae and phytosauria + archosauria ( see [ 31 ] ; figures 2 and 3 ) . five unambiguous local synapomorphies support euparkeriidae ( table 1 ) , but no unique apomorphies could be identified for the clade ( see section systematic palaeontology ) .
the other four synapomorphies supporting euparkeriidae can only be scored for euparkeria capensis and osmolskina czatkowicensis —the two most completely known euparkeriid taxa : presence of large anterolaterally opening foramen on anterolateral surface of maxilla ( character 29 , state 1 ) ; presence of four premaxillary teeth ( character 6 , state 1 ) ; presence of two foramina for cranial nerve xii in the exoccipital ( character 115 , state 0 ) ; and separately ossified interparietal ( character 139 , state 0 ) . the distribution of these characters is discussed further in the diagnosis .