pamizinisaurus is a genus of sphenodontian reptile known from lower cretaceous ( albian ) tlayúa formation of mexico .

a crushed skeleon of a juvenile reptile was found in tlayua quarry , in central mexico .

it was named pamizinsaurus tlayuaensis by reynoso in 1997 , after the name of the quarry of which it was found .

its skull length is 16 millimetres ( 0.63 in ) .

the fossil was covered in small round osteoderms that could have protected it from predators .

pamizinsaurus

pamizinsaurus was a genus of the subfamily sphenodontinae ; grouping it with the modern sphenodon ( better known as the tuatara ) , zapatadon , cynosphenodon , homoeosaurus , sapheosaurus , and ankylosphenodon .
pamizinsaurus was a genus of the subfamily sphenodontinae ; grouping it with the modern sphenodon ( better known as the tuatara ) , zapatadon , cynosphenodon , homoeosaurus , sapheosaurus , and ankylosphenodon .
the taxon pamizinsaurus from the early cretaceous of mexico ( reynoso 1997 ) was identified as a wildcard taxon that strongly influenced the results . this may partially be due to high amounts of missing data ( 64 % ) in this taxon ; however , that several other taxa have even higher amounts of missing data ( eilenodon 68 % ; cynosphenodon : 73 % ; toxolophosaurus : 79 % ) but proved to be less problematic , indicates that the wildcard status of pamizinsaurus might also be due to some character conflict .
reduced strict consensus ( after removing pamizinsaurus ) of sphenodontian interrelationships from 24 most parsimonious trees of 186 steps showing the phylogenetic position of sphenocondor gracilis gen . et sp . nov . bremer support and frequency differences ( gc ) , calculated by symmetric resampling , are indicated above the nodes .
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in the strict consensus of the present analysis , the inter - relationships within recovered groupings of derived sphenodontians are poorly resolved , as in other analyses with somewhat similar taxon sampling ( e . g . reynoso , 2000 , 2005 ) . however , this is mainly because of the unstable position of pamizinsaurus , which might be a consequence of the immature condition of the sole known specimen and its peculiar combination of character states ( reynoso , 1997 ) . therefore , pamizinsaurus was excluded from the strict consensus ( but not from the analysis ) , rendering a more resolved topology in the reduced strict consensus ( fig . 5 ) .
in a next step , we thus removed pamizinsaurus from the matrix and ran a second analysis with the same settings as above . the analysis resulted in only two most parsimonious trees with a length of 184 steps ( ci 0 . 473 , ri 0 . 654 , rc 0 . 309 ) . the strict consensus tree of these two trees (
vs - m size . see clevosaurs entry for basic skull structure . $ teeth fused to jaw margin ( fully acrodont ) and not replaced ; less than 4 ( 7 ? ) premaxillary teeth ; $ marginal teeth added posteriorly as jaw grows ; dentary fits between maxillary teeth and parallel row of palatal teeth ; meckelian canal runs along midline of jaw & is at least partially open ; broad mandibular symphysis ; prominent coronoid process of dentary , posterior process of dentary ends posterior to coronoid ; with posterior surangular facet ; frontals not fused ? ? ) ; $ lacrimal absent ; large upper temporal fenestra ; complete lower temporal bar ; quadratojugal retained ; rigid quadrate ; well - developed posterior tubercle on posterior margin of ischium ; epiphyses present with determinate growth ; may have osteoderms ( pamizinsaurus ) ; food crushed thoroughly ( insects & birds ) .
in a next step , we thus removed pamizinsaurus from the matrix and ran a second analysis with the same settings as above . the analysis resulted in only two most parsimonious trees with a length of 184 steps ( ci 0 . 473 , ri 0 . 654 , rc 0 . 309 ) . the strict consensus tree of these two trees ( fig . 3 and information s1 ) is considerably better resolved than that resulting from the full analysis , but , importantly , it does not contradict the results of the latter , neither in the strict consensus , nor in any reduced consensus tree . thus , we chose these results to illustrate and further explore the relationships of oenosaurus . bootstrap ( 10000 replicates ) and bremer support analyses were carried out on the pruned matrix , and the following list of synapomorphies is based on the strict consensus tree resulting from the pruned analysis .
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full reference : v . h . reynoso . 1997 . a ' beaded ' sphenodontian ( diapsida : lepidosauria ) from the early cretaceous of central mexico . journal of vertebrate paleontology 17 : 52 - 59
type specimen : instituto de geologãa , universidad nacional autã³noma de mã©xico , a partial skeleton ( severaly crushed skeleton of a juvenile ) . its type locality is tlayua formation , which is in an albian lagoonal / restricted shallow subtidal lime mudstone in the tlayua formation of mexico .
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sphenodontia : diphydontosaurus + ( planocephalosaurus + * : clevosauridae + stem group sphenodontinae ) .
comments : cladistically , this taxon ( equivalent to the standard linnean taxonomic sphenodontidae ) should perhaps be called something like sphenodontoidea , as it includes clevosauridae , pleurosauridae , . crown group sphenodontidae , etc
introduction to the sphenodontidae ; turtle , tuatara , crocodile checklist - - 3 ; jurassic page ' s gallery : the terrestrial wildlife ( sapheosaurus ) ; lebende fossilien , intro , teil 2 ( homoeosaurus ) .
references : evans & sigogneau - russell ( 1997 ) ; reynoso 1996 ) ; reynoso ( 1997 ) ; reynoso & clark ( 1998 ) .
image : kallimodon among a very good collection of fossil photos ) from herbert krause . reproduced by permission .
note : the nomenclature of sphenodont taxa tends to be rather confused . the scheme used here is adopted from the sensible system of mikko haaramo , who credits wu , x - c ( 1994 ) , late triassic - early jurassic sphenodontians from china and the phylogeny of the sphenodontia , in fraser , nc & h - d sues ( eds . ) , in the shadow of the dinosaurs , cambridge univ . press . atw001117 .
1 or more robust
incisor
teeth forming chisel edge ankylosed to premaxilla ; palatine dentition ; marginal dentition acrodont or absent ; shearing ( not crushing or piercing ) bite with simple orthal up - and - down ) jaw motion ; $ long thin posterior process of premaxilla excludes maxilla from margin of nares ; $ lacrimal absent ; may have dentine
lips
; $ antorbital skull very short ( < 25 % of total skull ) ; triradiate postfrontal broadly contacts frontal ; complete lower temporal bar formed by jugal loosely contacting slim quadratojugal ; $ dorsal process of jugal elongate ; $ large ( > 25 % skull length ) lower temporal fenestra ; gap between jugal and quadratojugal small ; proximal caudal vertebrae have large transverse processes ; caudal autotomy septa may be present distally ; humerus strongly bent and expanded at both ends ; femur gently s - shaped .
note : for reasons i have not been able to determine , no one seems to have gotten around to giving this group a latinized family name . atw . update : this has now been rectified . on the basis of new material arantes et al 2009 formalises the family clevosauridae , composed of clevosaurus , brachyrhinodon and polysphenodon . mak101016
links : diapsida ; clevosaurus bairdi ( but server is frequently reduced or absent ! ) ; autapomorphies of diapsid clades ; untitled document ; lecture 12 - early jurassic .
reynoso 1996 ) ; reynoso & clark 1998 ) ; sues et al . ( 1994 ) .
short snout like brachyrhinodon , multiple rows of teeth on the palate . it shows sphenodontian characters such as a groove between the maxilla and palatine for the dentary
comments : known from a partial skull and skeleton . the only known specimen went missing in the 1930s , but a number of casts and plaster moulds are available ( fraser & benton 1989 p . 416 . )
brachyrhinodon taylori , skull in ventral ( a ) , dorsal ( b ) and lateral ( c ) view . from fraser & benton 1989 p . 416 . scale bar 1 cm . note palatal teeth in ( a ) ; teeth on the upper palate ( roof of the mouth ) are a primitive feature found in most basal amniotes
small ( c . 15 cm long ) , short - snouted skull with premaxillary beak . benton 1985
kallimodon pulchellus ( zittel ) , 1887 . late jurassic ; kelheim , bavaria . ventral aspect . from zittel / eastman / woodward 1902 p . 151 . this represents a type similar to or intermediate between homoeosaurus and sapheosaurus
sphenodontoidea : clevosaurinae + * : homoeosaurus + pleurosauridae + ( sapheosaurus + sphenodontinae ) .
vertebrae amphicoelous , sometimes with persistent notochord ; intercentra present in cervical ( neck ) and caudal ( tail ) regions . external nares separated ; interclavicle t - shaped ; dermal scales subrectangular . greatiy enlarged postfrontal bone extending far posteriorly ( to the rear ) on parietal . except for a few teeth on the vomer in juveniles and the palatine row , all other palatal teeth absent ( i . e . teeth on the roof of the mouth , in contrast to primitive amniotes ) . premaxillae each with a small pointed tooth . a single series of enlarged , depressed , triangular , acrodont teeth present on maxillae , mandibles , and outer edge of the palatines ; vomer toothless . pattern of tooth wear facets , tooth ultrastructure , and anteroposterior length of the mandibular articulation , all of which indicate development of the propalinal masticatory movements ( front to rear chewing ) as with the recent sphenodon zittel / eastman / woodward 1902 , gauthier et al 1988 p . 25
kimmeridgian ( lithographic stone ) of bavaria , and cerin , france . kimmeridgian of hanover , and purbeckian ( tithonian ) of england .
from zittel / eastman / woodward 1902 ( copyright expired ) :
attaining a length of between 20 and 40 cm . , and differing from the recent sphenodon in that intercentra are absent between the dorsal vertebrae ribs without uncinate processes , and humerus not pierced by entepicondylar foramen . mandibular rami united at the symphysis by ligaments ; second sacral rib bifid distally .
applies also to sapheosaurus and kallimodon gauthier et al 1988 p . 26 : compared to sphenodontines and sapheosaurs , homoeosaurus is distinguished by smaller size , gracile and elongate limbs , and a broad parietal table ( skull roof above the orbits ( eye holes ) ) , all of which , with the possible exception of the limb proportions , are plesiomorphic ( shared ancestral ) features .
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early cretaceous lepidosaurs ( reptilia : d . . . - pdf document ( 15 m )
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today were then held by sphenodontians . there were even several successful groups of aquatic sphenodontians such as
( loss of the tail - tip when threatened ) , and have transverse cloacal slits .
r . l . ditmars , litt . d , says ;
the tuatara resembles in form stout - bodies modern lizards , which we might call iguanas ; this resemblance is further intensified by a row of spines upon the back . it is dark olive , the sides sprinkled with pale dots . the eye has a cat - like pupil . large specimens are two and a half feet long . while superficial resemblance might tend to group this reptile with lizards , its skeleton and anatomy show it to belong to a different part of a technical classification .
of the british museum noted features similar to birds , turtles , and crocodiles . he proposed the order rhynchocephalia ( meaning
beak head
) for the tuatara and its fossil relatives .
proposed the sphenodontia to include only tuatara and their closest fossil relatives in 1925 .
ditmars , raymond l . ,
reptiles of the world
the macmillan co . , new york , 1936 , p . xii
v . h . reynoso . 2000 . an unusual aquatic sphenodontian ( reptilia : diapsida ) from the tlayua formation ( albian ) , central mexico . journal of paleontology 74 : 133 - 148
cree , alison . 2002 . tuatara . in : halliday , tim and adler , kraig ( eds . ) , the new encyclopedia of reptiles and amphibians , oxford university press , oxford , pp . 210–211 . isbn 0 - 19 - 852507 - 9
fraser , nicholas ; sues , hans - dieter ; ( eds ) ( 1994 ) .
evans , s . e . , prasad , g . v . r . & manhas , b . k . , 2001 : rhynchocephalians ( diapsida : lepidosauria ) from the jurassic kota formation of india . zoological journal of the linnean society : vol . 133 , # 3 , pp . 309 - 334
rauhut , o . w . m . ; heyng , a . m . ; lópez - arbarello , a . ; hecker , a . ( 2012 ) . farke , andrew a , ed .
a new rhynchocephalian from the late jurassic of germany with a dentition that is unique amongst tetrapods
.
daugherty , ch ; cree , a ; hay , jm ; thompson , mb ( 1990 ) .
neglected taxonomy and continuing extinctions of tuatara (
evans , se ( 2003 ) .
at the feet of the dinosaurs : the early history and radiation of lizards
.
jones meh . 2009 . dentary tooth shape in sphenodon and its fossil relatives ( diapsida : lepidosauria : rhynchocephalia ) . in koppe t , meyer g , alt kw , eds . interdisciplinary dental morphology , frontiers of oral biology ( vol 13 ) . greifswald , germany ; karger . 9–15 .
evans se , jones meh ( 2010 ) the origin , early history and diversification of lepidosauromorph reptiles . in bandyopadhyay s . ( ed . ) , new aspects of mesozoic biodiversity , 27 lecture notes in earth sciences 132 , 27 - 44 .
this article is issued from wikipedia - version of the 11 / 25 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .
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rhynchocephalians , the sister group of squamates ( lizards and snakes ) , are only represented by the single genus sphenodon today . this taxon is often considered to represent a very conservative lineage . however , rhynchocephalians were common during the late triassic to latest jurassic periods , but rapidly declined afterwards , which is generally attributed to their supposedly adaptive inferiority to squamates and / or mesozoic mammals , which radiated at that time . new finds of mesozoic rhynchocephalians can thus provide important new information on the evolutionary history of the group .
a new fossil relative of sphenodon from the latest jurassic of southern germany , oenosaurus muehlheimensis gen . et sp . nov . , presents a dentition that is unique amongst tetrapods . the dentition of this taxon consists of massive , continuously growing tooth plates , probably indicating a crushing dentition , thus representing a previously unknown trophic adaptation in rhynchocephalians .
the evolution of the extraordinary dentition of oenosaurus from the already highly specialized zahnanlage generally present in derived rhynchocephalians demonstrates an unexpected evolutionary plasticity of these animals . together with other lines of evidence , this seriously casts doubts on the assumption that rhynchocephalians are a conservative and adaptively inferior lineage . furthermore , the new taxon underlines the high morphological and ecological diversity of rhynchocephalians in the latest jurassic of europe , just before the decline of this lineage on this continent . thus , selection pressure by radiating squamates or mesozoic mammals alone might not be sufficient to explain the demise of the clade in the late mesozoic , and climate change in the course of the fragmentation of the supercontinent of pangaea might have played a major role .
citation : rauhut owm , heyng am , lópez - arbarello a , hecker a ( 2012 ) a new rhynchocephalian from the late jurassic of germany with a dentition that is unique amongst tetrapods . plos one 7 ( 10 ) : e46839 . urltoken
editor : andrew a . farke , raymond m . alf museum of paleontology , united states of america
copyright : © 2012 rauhut et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
the recent genus sphenodon is the only living representative of the rhynchocephalia , the sister taxon of the squamata ( lizards and snakes ) within the lepidosauria [ 1 ] . due to the long history of the group , reaching back to at least the middle triassic ( ∼235 million years ) , and the supposedly primitive morphology of the genus , sphenodon is often regarded as a “living fossil” . thus , this taxon is frequently used as a model organism for a basal diapsid in investigations dealing with the evolution of reptiles , even in recent studies ranging from the structure of the pineal organ [ 2 ] to tetrapod locomotion [ 3 ] .
rhynchocephalians were a common component of mesozoic small vertebrate faunas [ 4 ] , [ 5 ] , and research in the past thirty years has shown that they were not only taxonomically and ecologically diverse during this time , but also showed a remarkable morphological variability [ 6 ] . indeed , several of the features that were thought to represent the plesiomorphic condition in sphenodon , such as the closed lower temporal arcade , were recently demonstrated to be apomorphic reversals to a condition resembling the ancestral morphology instead ( e . g . [ 7 ] , [ 8 ] ) . nevertheless , all rhynchocephalians but the most basal forms are characterized by a very specialized dentition pattern , in which the teeth are fully acrodont , no tooth replacement is present in post - hatchling individuals , and additional teeth are added at the posterior end of the tooth row during ontogeny [ 9 ] . although tooth shape itself is quite variable in rhynchocephalians [ 10 ] , this special type of tooth development was thought to limit their adaptive potential [ 11 ] . consequently , the demise of the group in the later mesozoic has been linked to the adaptive radiation of squamates and / or mammals ( see [ 5 ] ) .
here we report on a new taxon of rhynchocephalian from the late jurassic of southern germany that further underlines the morphological and ecological diversity of rhynchocephalians and indicates high adaptive plasticity even in the dentitions of these animals . the specimen was found in the middle parts of the moernsheim formation exposed at mühlheim ( fig . 1 ) , in a section of platy , siliceous limestones overlying lime bank b - h - 5 ( “krebs - bank” ) . the marine tithonian ( upper jurassic ) moernsheim formation covers the famous solnhofen lithographic limestones in the western region of the franconian alb in northern bavaria . in contrast to the underlying solnhofen formation , the sediments of the moernsheim formation are more heterogeneous , consisting of mainly siliceous limestones in alternate bedding with marly and limey sections , and with intercalated banks particularly in the lower parts of the formation .
the data matrix was analysed with paup 4 . 0b10 ( swofford 2003 ) , using a branch and bound search . the initial analyses resulted in 119 trees with a length of 187 steps ( ci 0 . 465 , ri 0 . 649 , rc 0 . 302 ) . the strict consensus tree of these trees showed poor resolution , with oenosaurus representing a member of a polytomy of rhynchocephalians more derived than planocephalosaurus ( information s1 ) ; only pleurosaurs and eilenodontines showed up as monophyletic clades within this polytomy . however , reduced consensus trees of these 119 trees recovered a monophyletic sphenodontinae , including oenosaurus , but without further resolution within this clade ( information s1 ) .
the electronic edition of this article conforms to the requirements of the amended international code of zoological nomenclature , and hence the new names contained herein are available under that code from the electronic edition of this article . this published work and the nomenclatural acts it contains have been registered in zoobank , the online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix “ urltoken ” . the lsid for this publication is : urn : lsid : zoobank . org : pub : 4a00d064 - a136 - 4944 - af8e - 427b941fb38c . the electronic edition of this work was published in a journal with an issn , and has been archived and is available from the following digital repositories : pubmed central , lockss .
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oenos , [ oinos ] greek , wine , referring to the franconian alb , a famous wine area , where the specimen was found , and saurus , [ sauros ] greek , lizard . species name refers to the village of mühlheim , close to mörnsheim , at the rim of the valley of the altmühl in bavaria .
bayerische staatssammlung für paläontologie und geologie , munich , germany , bspg 2009 i 23 ; partial skull and complete mandibles ( figs . 2 , 3 , 4 ) .
( a ) –skull in dorsal view . ( b ) –skull in ventral view . ( c–d ) –right mandible in lateral ( c ) and medial ( d ) views . ( e ) –left mandible in lateral view . ( f ) –stereophotographs of the skull in ventral view . abbreviations : a , angular ; ar , articular ; bsp , basisphenoid ; bt , basal tubera ; co , coronoid ; cos , coronoid shelf ; d , dentary ; ect , ectopterygoid ; eo , exoccipital / opisthotic ; f , frontal ; fo , foramen ; j , jugal ; m , maxilla ; mf , mandibular foramen ; mg , meckelian groove ; oc , occipital condyle ; pa , parietal ; pal , palatine ; po , postorbital ; pof , postfrontal ; prf , prefrontal ; pt , pterygoid ; sa , surangular ; sy , symphysis ; tp , tooth plate . scale bars are 10 mm .
( a ) –left maxillary tooth plate in ventral view ( anterior is to the left ) . ( b ) –enlargement of posterior part of left maxillary tooth plate , showing compound structure of the plate . ( c ) –further enlargement of left maxillary tooth plate , showing closely packed , worn teeth with central pulp cavities . scale bars are 1 mm ( a , b ) and 0 . 5 mm ( c ) .
posterior is to the right . abbreviations : bc , basal cavity ; f , foramen . small arrows point to bifurcating dentine channels . scale bar is 5 mm .
mörnsheim formation ( lower tithonian ) [ 16 ] , “krautworst naturstein” quarry in mühlheim near mörnsheim , central bavaria , germany ( fig . 1 ) .
small rhynchocephalian with the following autapomorphic characters : maxilla with a medial process at the posterior end ; ectopterygoid with a secondary lateral process that contacts the medial side of the maxilla ; palatines with a midline contact in ventral view ; strongly pronounced lateral longitudinal groove on the dentaries , housing several large foramina ; very high coronoid process , the anterior border of which forms an angle of approximately 90° with the tooth row ; coronoid with a pronounced shoulder medially ; dentition composed of extensive tooth plates formed by a multitude of fused , small , pencil - like teeth .
the skull of oenosaurus was preserved in ventral view , with the mandibles slightly displaced from the upper jaws . the skull is diagenetically dorsoventrally compressed and much of the dorsal skull roof is missing , as is the roof of the braincase . however , enough is preserved to record the general shape of the skull and the position and size of the orbitae ( fig . 2a ) .
the ectopterygoid attaches anterolaterally to the pterygoid wing . it is unusual in that it has two lateral processes , one that contacts the posterior end of the maxilla , as in other rhynchocephalians , and a more anterior one that extends anterolaterally lateral to the palatine and contacts the medial side of the maxilla ( fig . 2b , f ) . this process forms the posterior border of the suborbital fenestra , which is surrounded by the palatine , maxilla , and ectopterygoid . thus , the pterygoid is excluded from the rim of this fenestra , as in many other sphenodontids , but not more basal rhynchocephalians . in contrast to clevosaurs [ 21 ] , [ 22 ] , [ 23 ] there is no contact between the palatine and the ectopterygoid lateral to the fenestra . together with the medial process of the maxilla , the secondary anterolateral process frames a large foramen lateral to the palatine ridge .
only the floor of the braincase and the ventral parts of the occiput are preserved . the basisphenoid is narrow at the basicranial articulation , but rapidly widens posteriorly . the basipterygoid processes seem to be stout and short , but are mostly hidden by the pterygoids . the basisphenoid has a broad , shallow depression on the ventral side between the basipterygoid processes and the low , widely separated basal tubera . the occipital condyle is broad and not separated from the basioccipital body by a constricted neck , similar to the situation in the modern sphenodon .
the lower jaw is 33 mm long and is especially notable for its high coronoid process , which exceeds the tooth row by approximately 8 . 7 mm in the right mandible and is thus almost 1 . 5 times higher than the body of the dentary ( fig . 2c , d ) . the dentary accounts for most of the length of the mandible , and , as in all rhynchocephalians , a posteroventral process of the dentary reaches posteriorly beyond the coronoid process to the level of the anterior margin of the mandibular articulation . a large incision in the posterodorsal margin of the dentary marks the enlarged mandibular foramen ( fig . 2c , e ) . a pronounced groove with several large foramina is present on the lateral side of the dentary , and its dorsal margin seems to be formed by a secondary bone skirt [ 5 ] , [ 23 ] , which probably accounts at least partially for the strong transverse thickening of the tooth - bearing part of the dentary . anteriorly , the ventral margin of the bone extends ventrally to form a small , triangular “chin” , as it is present in many rhynchocephalians ( e . g . [ 20 ] , [ 24 ] – [ 26 ] ) . there is no splenial .
the postdentary bones are entirely restricted to the posterior half of the mandible . the surangular makes up the posterior margin of the ventral part of the coronoid process and continues posteriorly to the end of the mandible . the mandibular articulation is developed as a longitudinal dorsal ridge on the articular . the ridge is sharp - edged dorsally and slightly displaced to the lateral side . medially , it is flanked by a slightly anteroposteriorly concave medial bulge that becomes more pronounced posteriorly . this morphology is similar to the condition in derived rhynchocephalians , possibly indicating a propalinal movement of the lower jaw . the retroarticular process is short and stout . the medial side of the coronoid is thickened and forms a pronounced shelf below the tip of the coronoid process .
the most unusual character of the new taxon is the dentition . unlike the situation in any other rhynchocephalian , broad tooth plates are present in both the maxillaries and dentaries . the maxillary tooth plates ( fig . 2b , f , 3a ) are broad posteriorly and narrow in their anterior half . they are 13 mm long and 4 mm wide at their widest part . the lateral edge of the tooth plates is slightly raised and shows a few small bumps . the dentary tooth plates are elongate and oval to subrectangular in outline , with a length of 13 mm and a maximum width of 3 . 5 mm . ct data shows that the central part of the tooth plate is placed in a broad depression on the dorsal surface of the mandible , though the tooth implantation can still be regarded as acrodont rather than protothecodont , as it is the case in some basal rhynchocephalians [ 27 ] .
under closer inspection , the surfaces of the tooth plates show small , but clearly defined , round , oval , or angular subunits , made up of concentric dentine layers with a small central cavity ( fig . 3 ) . fine striations radiate outwards from the central cavity . micro ct - images of the dentary tooth plate show that these dentine tubes extend over the entire height of the plate ( fig . 4 ) . in the basal part , the central cavities become larger and form large , elongate cavities at the base of the tooth plate . in some instances , the central cavities of the dentine tubes bifurcate in their course from the base to the occlusal surface . tooth enamel seems to be absent , but the dentine tubes are tightly cemented together towards the occlusal surface . at the occlusal surface , the central cavities of the tubes also seem to be filled with cement . the dentine tubes are generally somewhat larger in the central parts of the tooth plate , but become smaller towards the margins ( fig . 3a ) . the dentine tubes reach further ventrally in the anterior part of the dentary than in the posterior part , which might indicate a later addition of posterior dentine tubes during ontogeny , as it is the case with individual teeth in other rhynchocephalians [ 9 ] .
despite its apomorphic cranial morphology and highly unusual dentition , cladistic analysis ( see information s1 ) places oenosaurus well within the rhynchocephalia ( fig . 5 ) , because the taxon shows numerous synapomorphies of this clade and more exclusive ingroups , such s sphenodontia and sphenodontidae . these characters include a pronounced ridge or tooth row laterally on the palatine , a high coronoid process in the mandible , a posterior process of the dentary that extends posterior to the coronoid process , an enlarged mandibular foramen , the lack of a splenial , and an acrodont dentition [ 18 ] , [ 22 ] , [ 26 ] , [ 28 ] – [ 30 ] . within rhynchocephalians , oenosaurus even falls within the clade that includes the recent form , the sphenodontinae [ 5 ] , [ 24 ] , though support for this placement is weak ( see information s1 ) , and further material of this taxon ( and others , which , with the exception of sphenodon , are generally poorly known [ 5 ] ) might lead to changes in this part of the tree . nevertheless , the phylogenetic position of the new taxon within sphenodontidae ( sensu [ 24 ] ) is well supported .
cladistic analysis of 70 osteological characters in 19 rhynchocephalian and two outgroup taxa resulted in the recovery of 2 trees with 184 steps ( see information s1 ) . a strict consensus tree shows oenosaurus well nested within sphenodontine rhynchocephalians , as a close relative to the recent sphenodon . 1 , rhynchocephalia ; 2 , sphenodontinae .
an interesting problem is the evolution of these tooth plates . the phylogenetic position of oenosaurus shows that this taxon is well nested within taxa that have the specialized acrodont dentition typical for sphenodontids . the very peculiar plates are therefore derived from such a dentition and demonstrate a surprising evolutionary plasticity of such a seemingly highly specialized tooth anlage [ 9 ] . a possible , but currently untestable , hypothesis of the evolution of these tooth plates might be that they formed by fusion and modification of the hatchling dentition , which consists of small , peg - like teeth with active tooth replacement in several rhynchocephalians [ 41 ] , including the modern sphenodon [ 42 ] , [ 43 ] . however , further studies and , possibly , further discoveries of this or other taxa with similar dentitions are necessary to test this hypothesis .
despite their specialized dentition , rhynchocephalians are quite variable in their tooth morphology . rhynchocephalian teeth can be referred to three basic functional types [ 10 ] : basal forms usually have small , conical teeth that are mainly used for piercing , most sphenodontids have anteroposteriorly elongate teeth that are used for cutting and slicing , and opisthodontians [ 18 ] have transversely expanded teeth capable of grinding and shredding . these different tooth types can roughly be equated with insectivory , generalized carnivory ( or piscivory , in the marine pleurosaurs ) , and herbivory , respectively [ 10 ] . the tooth plates of oenosaurus now demonstrate a further type of dentition in rhynchocephalians , a crushing dentition , indicating a durophagous diet . therefore , the new taxon adds a further trophic adaptation and a previously unknown ecotype to the already recognized ecological diversity of mesozoic rhynchocephalians [ 10 ] .
character list , data matrix , analytical procedures , consensus trees , and list of synapomorphies at internal nodes for the cladistic analysis mentioned in the text .
we thank roland pöschl , who found the fossil , and the owners and operators of the krautworst naturstein quarry , ulrich leonhardt , roland pöschl and uwe krautworst , for donating the specimen to the bspg . ulrich leonhardt , stefan sónyi , and renate liebreich are thanked for preparation of the fossil . the paper benefited from discussions with marc jones and alexander nützel . furthermore , critical comments by marc jones and nick fraser considerably helped to improve the paper . ct scans were carried out at the steinmann institute of the university of bonn with the help of irina ruf , which is greatly appreciated .
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oliver w . m . rauhut , 1 , 2 , * alexander m . heyng , 2 adriana lópez - arbarello , 1 and andreas hecker 3
* e - mail : ed . nehcneum - inu . zrl @ tuhuar . o
this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are properly credited .
here we report on a new taxon of rhynchocephalian from the late jurassic of southern germany that further underlines the morphological and ecological diversity of rhynchocephalians and indicates high adaptive plasticity even in the dentitions of these animals . the specimen was found in the middle parts of the moernsheim formation exposed at mühlheim (
) , in a section of platy , siliceous limestones overlying lime bank b - h - 5 ( “krebs - bank” ) . the marine tithonian ( upper jurassic ) moernsheim formation covers the famous solnhofen lithographic limestones in the western region of the franconian alb in northern bavaria . in contrast to the underlying solnhofen formation , the sediments of the moernsheim formation are more heterogeneous , consisting of mainly siliceous limestones in alternate bedding with marly and limey sections , and with intercalated banks particularly in the lower parts of the formation .
) is considerably better resolved than that resulting from the full analysis , but , importantly , it does not contradict the results of the latter , neither in the strict consensus , nor in any reduced consensus tree . thus , we chose these results to illustrate and further explore the relationships of oenosaurus . bootstrap ( 10000 replicates ) and bremer support analyses were carried out on the pruned matrix , and the following list of synapomorphies is based on the strict consensus tree resulting from the pruned analysis .
was preserved in ventral view , with the mandibles slightly displaced from the upper jaws . the skull is diagenetically dorsoventrally compressed and much of the dorsal skull roof is missing , as is the roof of the braincase . however , enough is preserved to record the general shape of the skull and the position and size of the orbitae (
the skull seems to have been rather robust . as preserved , it is 24 . 3 mm long . unfortunately , the exact skull length cannot be determined , since the premaxillae are missing , but these bones probably did not account for more than an additional 2–3 mm . thus , the skull was slightly broader ( 28 mm ) than long , as in many rhynchocephalians , but unlike the slender , elongate skull of the marine pleurosaurs
. premaxillae and nasals are missing . the frontals are elongate and narrow , and flanked by the prefrontals anterolaterally (
) . there is no trace of a lacrimal , though it cannot be ruled out completely that this might be due to preservation . the posterior part of the skull roof is poorly preserved and nothing can be said about the shape and size of the temporal fenestrae . a small fragment of the parietals shows that these bones were fused and formed a narrow sagittal crest medially . the postfrontals and postorbitals formed a broad shelf anterior to the dorsal temporal fenestra . the jugal was broad and reached anteriorly to almost half - length of the orbit . the maxillae are massive and broad , but rapidly narrow anteriorly . posteriorly , the maxilla has a stout lateral process for the contact with the jugal and a smaller medial process that slots into the forked lateral process of the ectopterygoid .
) . the vomers were small . their posterior ends separate the anterior ends of the palatines , but do not reach the pterygoids posteriorly , so that the palatines meet at the midline . such a midline contact of the palatines was illustrated for the marine pleurosaurs by carroll and wild
, but not described in the text , and the line of contact is dotted in their reconstruction , so that some uncertainty of the condition in these animals remains . the palatines contact the maxillae laterally in a broad suture at about one third of the length of the latter bone , and form the posterior border of the internal choanae . the bones flank the pterygoids laterally over most of the length of the anterior process of the latter bone , taper posteriorly , and end a short way posterior to the posterior end of the maxillae . as in all sphenodontids , there is a raised ridge along the lateral edge of the posterior half of the palatines , but , due to preservation , it is uncertain if this ridge bore teeth , as it is the case in most taxa
, the ridge diverges from the maxillary posteriorly and is more or less parallel to the midline of the skull . the pterygoids taper anteriorly and broaden abruptly posteriorly towards the ventrolaterally extending pterygoid wings . there is no interpterygoid vacuity , but the pterygoids form a midline suture up to the basicranial articulation . posteriorly , the slender , but well - developed , quadrate wings of the pterygoids diverge away from the ventral side of the braincase . a small fragment of bone lateral to the remnants of the left quadrate wing of the pterygoid might represent a remain of the pterygoid wing of the quadrate . unlike basal rhynchocephalians , there are no teeth on the pterygoid . no suture between the quadrate wing of the pterygoid and the pterygoid wing of the quadrate is visible in the preserved portions , so the latter bone , which is not preserved , might have had a rather short pterygoid wing .
the ectopterygoid attaches anterolaterally to the pterygoid wing . it is unusual in that it has two lateral processes , one that contacts the posterior end of the maxilla , as in other rhynchocephalians , and a more anterior one that extends anterolaterally lateral to the palatine and contacts the medial side of the maxilla (
) . this process forms the posterior border of the suborbital fenestra , which is surrounded by the palatine , maxilla , and ectopterygoid . thus , the pterygoid is excluded from the rim of this fenestra , as in many other sphenodontids , but not more basal rhynchocephalians . in contrast to clevosaurs
there is no contact between the palatine and the ectopterygoid lateral to the fenestra . together with the medial process of the maxilla , the secondary anterolateral process frames a large foramen lateral to the palatine ridge .
the lower jaw is 33 mm long and is especially notable for its high coronoid process , which exceeds the tooth row by approximately 8 . 7 mm in the right mandible and is thus almost 1 . 5 times higher than the body of the dentary (
) . the dentary accounts for most of the length of the mandible , and , as in all rhynchocephalians , a posteroventral process of the dentary reaches posteriorly beyond the coronoid process to the level of the anterior margin of the mandibular articulation . a large incision in the posterodorsal margin of the dentary marks the enlarged mandibular foramen (


macrobathra mesopora is a moth in the cosmopterigidae family .

it was described by meyrick in 1886 .

the adult moths have off-white forewings with bold dark brown bands and patches .

the hindwings are dark brown .

the wingspan is about 1.5 cm .

it is found in australia , where it has been recorded from new south wales .

macrobathra mesopora

macrobathra mesopora meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 804 ; tl : blackheath , new south wales
macrobathra crococephala meyrick , 1936 ; exotic microlep . 5 ( 2 ) : 50
macrobathra auratella viette , 1958 ; rev . franc . ent . 25 ( 2 ) : 119
macrobathra is a genus of moth in the family cosmopterigidae . most species are endemic to australia .
macrobathra cineralella viette , 1957 ; mém . inst . sci . madagascar ( e ) 8 : 162
macrobathra vexillariata lucas , 1901 ; proc . r . soc . qd 16 : 90 ; tl : brisbane
macrobathra crococosma meyrick , 1922 ; exotic microlep . 2 ( 16 ) : 510 ; tl : queensland , cairns
macrobathra gentilis meyrick , 1918 ; exotic microlep . 2 ( 7 ) : 212 ; tl : kanara , dharwar
macrobathra pyrodoxa meyrick , 1926 ; exot . microlep . 3 ( 10 ) : 310 ; tl : new ireland
macrobathra arneutis meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 218 ; tl : assam , margherita
macrobathra astrota meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 216 ; tl : queensland , herberton
macrobathra decataea meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 216 ; tl : queensland , townsville
macrobathra hedrastis meyrick , 1915 ; exot . microlep . 1 ( 10 ) : 295 ; tl : tenasserim , mergui
macrobathra myrocoma meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 218 ; tl : assam , khasis
macrobathra notomitra meyrick , 1924 ; exot . microlep . 3 ( 4 ) : 98 ; tl : bengal , pusa
macrobathra notozyga meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 217 ; tl : queensland , herberton
macrobathra recrepans meyrick , 1926 ; exot . microlep . 3 ( 10 ) : 310 ; tl : rhodesia , umvuma
macrobathra synacta meyrick , 1920 ; exotic microlep . 2 ( 12 ) : 364 ; tl : south australia , yutala
macrobathra proxena meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 216 ; tl : nyassaland , mt mlanje
macrobathra anacampta meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 217 ; tl : queensland , herberton , brisbane
macrobathra antimeloda meyrick , 1924 ; exot . microlep . 3 ( 18 - 20 ) : 614 ; tl : madagascar , imerina
macrobathra neurocoma meyrick , 1930 ; exot . microlep . 3 ( 18 - 20 ) : 614 ; tl : cameroons , lolodorf
macrobathra nomaea meyrick , 1914 ; exot . microlep . 1 ( 7 ) : 217 ; tl : ceylon , colombo , peradeniya
macrobathra ochanota meyrick , 1915 ; exot . microlep . 1 ( 10 ) : 295 ; tl : coorg , dibidi , 3500ft
macrobathra petalitis meyrick , 1915 ; exot . microlep . 1 ( 10 ) : 295 ; tl : coorg , dibidi , 3500ft
macrobathra anisodora meyrick , 1924 ; exot . microlep . 3 ( 4 ) : 98 ; tl : cape colony , kowie r .
macrobathra equestris meyrick , 1910 ; j . bombay nat . hist . soc . 20 ( 1 ) : 143 ; tl : khasis
macrobathra hexadyas meyrick , 1906 ; trans . r . soc . s . aust . 30 : 35 ; tl : rosewood , queensland
macrobathra homocosma meyrick , 1902 ; trans . r . soc . s . aust . 26 : 167 ; tl : duaringa , queensland
macrobathra quercea moriuti , 1973 ; tyô to ga 23 ( 2 ) : 35 ; tl : kuragaritoge , osaka pref . , honshu
macrobathra xanthoplaca meyrick , 1902 ; trans . r . soc . s . aust . 26 : 167 ; tl : melbourne , victoria
macrobathra monoclina meyrick , 1915 ; exot . microlep . 1 ( 10 ) : 294 ; tl : ceylon , colombo ; coorg , dibidi
macrobathra peraeota meyrick , 1921 ; ann . transv . mus . 8 ( 2 ) : 98 ; tl : portuguese east africa , magude
macrobathra aneurae turner , 1932 ; proc . linn . soc . n . s . w . 57 : 272 ; tl : queensland , charleville
macrobathra asemanta lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 103 ; tl : windermere , tasmania
macrobathra chryseostola turner , 1932 ; proc . linn . soc . n . s . w . 57 : 272 ; tl : queensland , rockhampton
macrobathra dasyplaca lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 103 ; tl : windermere , tasmania
macrobathra diplochrysa lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 104 ; tl : brisbane , queensland
macrobathra embroneta turner , 1932 ; proc . linn . soc . n . s . w . 57 : 274 ; tl : queensland , brisbane
macrobathra galenaea meyrick , 1902 ; trans . r . soc . s . aust . 26 : 167 ; tl : sydney , new south wales
macrobathra phryganina turner , 1932 ; proc . linn . soc . n . s . w . 57 : 268 ; tl : queensland , toowoomba
macrobathra platyzona turner , 1932 ; proc . linn . soc . n . s . w . 57 : 273 ; tl : queensland , toowoomba
macrobathra polypasta turner , 1932 ; proc . linn . soc . n . s . w . 57 : 272 ; tl : queensland , charleville
macrobathra fasciata ; meyrick , 1921 , ann . transv . mus . 8 ( 2 ) : 98 ; [ nhm card ] ; [ afromoths ]
macrobathra gastroleuca lower , 1905 ; trans . r . soc . s . austr . 29 : 109 ; tl : broken hill , new south wales
macrobathra isoscelana lower , 1893 ; trans . proc . r . soc . s . austr . 17 ( 1 ) : 182 ; tl : blackwood
macrobathra nimbifera turner , 1932 ; proc . linn . soc . n . s . w . 57 : 274 ; tl : queensland , national park
macrobathra psathyrodes turner , 1932 ; proc . linn . soc . n . s . w . 57 : 270 ; tl : queensland , bunya mts
macrobathra stenosema turner , 1932 ; proc . linn . soc . n . s . w . 57 : 271 ; tl : northern territory , darwin
macrobathra baliomitra turner , 1932 ; proc . linn . soc . n . s . w . 57 : 270 ; tl : new south wales , sydney
macrobathra heterocera lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 102 ; tl :
billopp
, tasmania
macrobathra anemarcha meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 805 ; tl : launceston , tasmania
macrobathra brontodes meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 821 ; tl : rosewood , queensland
macrobathra dispila turner , 1932 ; proc . linn . soc . n . s . w . 57 : 273 ; tl : n . queensland , cape york
macrobathra euryxantha meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 803 ; tl : duaringa , queensland
macrobathra leucozancla turner , 1932 ; proc . linn . soc . n . s . w . 57 : 270 ; tl : n . queensland , cape york
macrobathra melanargyra meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 817 ; tl : brisbane , queensland
macrobathra melanota meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 809 ; tl : oowoomba , queensland
macrobathra micropis lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 102 ; tl : coomooboolaroo range , duaringa , queensland
macrobathra microspora lower , 1900 ; proc . linn . soc . n . s . w . 25 ( 3 ) : 415 ; tl : mackay , queensland
macrobathra niphadobola meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 810 ; tl : rosewood , queensland
macrobathra paracentra lower , 1893 ; trans . proc . r . soc . s . austr . 17 ( 1 ) : 182 ; tl : gisborne , victoria
macrobathra platychroa lower , 1897 ; proc . linn . soc . n . s . w . 22 ( 2 ) : 270 ; tl : gisborne , victoria
macrobathra anemodes meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 819 ; tl : wirrabarra , south australia
macrobathra drosera lower , 1901 ; trans . r . soc . s . aust . 25 ( 2 ) : 96 ; tl : broken hill , new south wales
macrobathra harmostis meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1678 ; tl : geraldton , west australia
macrobathra hemitropa meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 817 ; tl : wirrabara , south australia
macrobathra heterozona meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1676 ; tl : northampton , west australia
macrobathra hyalistis meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1679 ; tl : york , west australia
macrobathra parthenistis meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1678 ; tl : carnarvon , west australia
macrobathra philopsamma lower , 1900 ; proc . linn . soc . n . s . w . 25 ( 1 ) : 47 ; tl : semaphore , south australia
macrobathra pompholyctis meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1677 ; tl : york , west australia
macrobathra synastra meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 815 ; tl : geroge´s bay , tasmania
macrobathra allocrana turner , 1916 ; proc . linn . soc . n . s . w . 41 ( 2 ) : 336 ; tl : n . queensland , innisfail
macrobathra callispila turner , 1916 ; proc . linn . soc . n . s . w . 41 ( 2 ) : 334 ; tl : north australia , port darwin
macrobathra heminephela meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 806 ; tl : launceston and hobart , tasmania
macrobathra porphyrea meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 820 ; tl : sydney , new south wales
macrobathra rhodospila meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 814 ; tl : sydney , new south wales
macrobathra xuthocoma meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 813 ; tl : sydney , new south wales
macrobathra aphristis meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1677 ; tl : carnarvon and northampton , west australia
macrobathra euspila turner , 1932 ; proc . linn . soc . n . s . w . 57 : 269 ; tl : new south wales , sydney ( woy woy )
macrobathra phernaea lower , 1899 ; proc . linn . soc . n . s . w . 24 ( 1 ) : 112 ; tl : broken hill , new south wales
macrobathra syncoma lower , 1899 ; proc . linn . soc . n . s . w . 24 ( 1 ) : 112 ; tl : broken hill , new south wales
macrobathra trimorpha meyrick , 1889 ; proc . linn . soc . n . s . w . ( 2 ) 3 : 1675 ; tl : carnarvon and geraldton , west australia
macrobathra zonodesma lower , 1900 ; proc . linn . soc . n . s . w . 25 ( 3 ) : 414 ; tl : broken hill , new south wales
macrobathra lychnophora turner , 1932 ; proc . linn . soc . n . s . w . 57 : 269 ; tl : queensland , bunya mts ; new south wales , allyn river
macrobathra callipetala turner , 1932 ; proc . linn . soc . n . s . w . 57 : 271 ; tl : n . australia , darwin ; n . queensland , cape york
macrobathra nephelomorpha meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 820 ; tl : toowoomba , queensland ; sydney , new south wales ; hobart , tasmania
macrobathra trithyra meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 808 ; tl : sydney and cooma , new south wales ; mt lofty , south australia
macrobathra ceraunobola meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 818 ; tl : sydney and blackheath , new south wales ; melbourne , victoria ; hobart , tasmania
macrobathra melanomitra meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 807 ; tl : bowenfels , new south wales ; mt lofty , wirrabara and quorn , south australia
macrobathra desmotoma meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 806 ; tl : brisbane and toowoomba , queensland ; newcastle and sydney , new south wales ; melbourne , victoria
macrobathra chrysotoxa meyrick , 1886 ; proc . linn . soc . n . s . w . 10 ( 4 ) : 804 ; tl : brisbane and toowoomba , queensland ; newcastle and sydney , new south wales ; melbourne and warragul , victoria ; launceston and hobart , tasmania
the adult moths have off - white forewings with bold dark brown bands and patches . the hindwings are dark brown . the wingspan is about 1 . 5 cms .
series 1 , volume 10 , part 4 ( 1886 ) , pp . 804 - 805 ,
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urn : lsid : biodiversity . org . au : afd . taxon : cee00aef - 1c98 - 42a7 - 87a6 - 1275e1d296d3
urn : lsid : biodiversity . org . au : afd . name : 382205
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aspasiodes allophyla turner , 1944 ; proc . linn . soc . n . s . w . 69 ( 1 - 2 ) : 51 ; tl : queensland , injune
s . queensland , new south wales , s . australia . see [ maps ]
macronemata basisticha turner , 1936 ; proc . linn . soc . n . s . w . 61 : 309 ; tl : queensland , brisbane
gelechia constrictella walker , 1864 ; list spec . lepid . insects colln br . mus . 29 : 647 ; tl : australia
deltozona meyrick , 1932 ² ; exotic microlep . 4 ( 8 - 9 ) : 276
stagmatophora distincta walsingham , 1891 ; trans . ent . soc . lond . 1891 ( 1 ) : 119 , pl . 6 , f . 57 ; tl : bathurst , gambia
gelechia epimela lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 106 ; tl : brisbane
borkhausenia ( ? ) erythrocephala lower , 1904 ; trans . r . soc . s . austr . 28 : 169 ; tl : broken hill , n . s . w :
stagmatophora fasciata walsingham , 1891 ; trans . ent . soc . lond . 1891 ( 1 ) : 118 , pl . 6 , f . 56 ; tl : bathurst , gambia
hamata meyrick , 1931 ² ; exotic microlep . 4 ( 5 ) : 116
leurozancla humilis turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 173 ; tl : queensland , national park
larva on quercus serrata , q . glauca moriuti , 1973 , tyô to ga 23 ( 2 ) : 38
lymnaecia subharpalea legrand , 1966 ; mém . mus . hist . nat . paris ( a ) 37 : 65 , pl . 4 , f . 10
lymnaecia superharpalea legrand , 1966 ; mém . mus . hist . nat . paris ( a ) 37 : 65 , pl . 4 , f . 11
laverna vividella felder & rogenhofer , 1875 ; reise fregatte novara , bd 2 ( abth . 2 ) ( 5 ) : pl . 140 , f . 6
gelechia xylopterella walker , 1864 ; list spec . lepid . insects colln br . mus . 29 : 650 ; tl : moreton bay
[ afromoths ] de prins , j . & de prins , w . , 2013
sangmi lee , richard brown & sibyl bucheli . gelechioidea - a global framework ;
[ ² ] this may require parentheses or not . i don ' t have the necessary information for this taxon .
reise der österreichischen fregatte novara um die erde in den jahren 1857 , 1858 , 1859 unter den behilfen des commodore b . von wüllerstorf - urbair . zoologischer theil . band 2 . abtheilung 2 . lepidoptera . rhopalocera
- 120 , ( inhalts - verz . ) 1 - 9 ( pl . 1 - 74 ) , ( felder & rogenhofer , 1874 ) , ( 5 ) : pl .
in la faune entomologique de i ' ile de lar réunion . lépidoptères ( excepté les tordeuses et les géométrides )
walsingham , 1891 african micro - lepidoptera trans . ent . soc . lond . 1891 ( 1 ) : 63 - 132 , pl . 3 - 7
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
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and the number of omitted authors .
attribute is enclosed in square brackets . this is rarely needed , but to differentiate homo - identifiers , this will contain the page , line or plate number of original publication .
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