deltavjatia was a procolophonid reptile from the tatarian stage of the permian time period .

it had a large body of about 1.5 m ( 5 ft ) in length .

deltavjatia was an herbivore .

deltavjatia lived in what is now russia .

the first specimen of deltavjatia was a specimen of a skull and lower mandobile ( pin 2212/1 ) , found in the urpalov formation in kotelnich , vyatka river .

deltavjatia was named a subtaxon of pareiasauridae by m.s.y. lee in 1997 .

it was named deltavjatia vjatkensis by hartmann & weinberg in 1937 .

pareiasuchus vjatkensis is the alternative combination of deltavjatia vjatkensis .

deltavjatia vjatkensis was also named anthodon rossicus by hartmann & weinberg in 1937 .

deltavjatia

first # fossil kashtanov discovered at # kotelnich locality was # pareiasaur # deltavjatia . # peoplemw # museumweek urltoken
deltavjatia was originally named as a species of pareiasuchus ( itself largely treated as a synonym to pareiasaurus )  ,  and then later as species of anthodon .  then in 1987 deltavjatia was established as a distinct genus by lebedev .  deltavjatia was a fairly standard pareiasaur ,  though the genus is noted as having several primitive features .
graphics photograph of deltavjatia vjatkensis © albert khlyupin and vyatka palaeontological museum - original page ( parareptilia ) , used with permission . skeletal reconstruction © david peters
now , more than 80 years after , we still find # deltavjatia # fossils every year in quite a big number . # paleontology # museumweek # russia
deltavjatia vjatkensis : pin 2212 / 1 , a skull ( including lower jaw ) . its type locality is kotelnich , vjatka river (
sarmin
series ) , pin 2212 , which is in a severodvinian terrestrial claystone / sandstone in the urpalov formation of the russian federation .
comments : originally placed in pareiasuchus ( hartmann - weinberg 1933 ) this smallish pareiasaur is distinct enough to warrant its own genus . the only pareiasaur known from the lower tatarian of russia , deltavjatia is an important taxon that bridges the large morphological and stratigraphic gap separating primitive south african tapinocephalus zone pareiasaurs from the more derived forms characteristic of the overlying cistecephalus zone and later deposits . it retains the long tail and feeble dermal armour of more primitive pareiasaurs , but possesses many skeletal traits characteristic of more advanced species lee 1997 . from above , the skull resembles the letter v , hence the name deltavjatia ( from the greek letter delta ) .
name : deltavjatia . phonetic : del - tah - v - yah - te - ah . named by : lebedev  -  1987 . synonyms : anthodon chlynoviensis ,  anthodon rossicus ,  pareiasuchus vjatkensis ,  scutosaurus rossicus . classification : chordata ,  reptilia ,  procolophonomorpha ,  pareiasauridae . species : d .  vjatkensis ( type )  . diet : herbivore . size : up to 1 . 5 meters long . known locations : russia  -  urpalov formation . time period : tatraian of the permian . fossil representation : several individuals .
( 3 ) in view of the paraphyletic status of bradysaurus ( tsuji & müller 2009 ) there doesn ' t seem to be a clear point where node velosauria should be placed . mikko haraama offers two alternatives , velosauria lee , 1994 sensu lee , 1997 , which is an apomorphy - based definition , everything more derived than bradysaurus , and velosauria lee , 1994 sensu jalil & janvier , 2005 , which is a node - based definition ( therischia + pumiliopareiasauria ) for everything more derived than deltavjatia . given that there is no unanimity regarding the therischia - tsuji & müller 2009 ' s unpublished analysis ( see cladogram in that paper ) did not recover it - we have gone for the former option . mak120323
these animals were probably highly gregarious , occuring in large numbers . as with many of the similarily abundant south african bradysaurs , most of the deltavjatia found at the kotelnich fossil locality were recovered as complete skeletons found fossilised in standing posture . the usual explanation is that they became bogged in mud died as a result of heavy rain fall , and were buried by silt and sediments carried by the flood water . another explanation is that these water - loving animals were crowded in small ponds when there was not enough water to swim through but the ground became series of mud traps . the deltavjatias preserved this way show not only the smallest bones of the feet and tail , but also the dermal ossicles . ( vyatka palaeontological museum )
no doubt because of its primitive nature relative to the later and more advanced scutosaurus , golubev 2000 places this genus in a stratigraphic table of russian permian tetrapod families under the
bradysauridae
. deltavjatia is also more lightly built than scutosaurus , and may have been a more active animal . despite its primitive status , it is still characterised by numerous turtle - like skull features ( e . g . , a very high palate ) , limbs , and girdles , and lateral projections flaring out some of the vertebrate in a very shell - like way . ( transitional vertebrate fossils faq by kathleen hunt ) ; this relatively small species lived immediately after the mid capitanian extinction event and represented the start of a new paraeiasaur adaptive radiation mak120331
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$ 12 or more cusps on teeth of lower jaw ; $ parallel , closely spaced rows of palatal teeth ; $ parasphenoid greatly shortened ; large , u - shaped interpterygoid vacuity ; $ < 20 caudal vertebrae ; cleithrum absent ; $ greater trochanter present and deflected ( ? ) from femoral head ; $ bony dermal armor , with large osteoderms ; $ armor covers entire dorsal surface , united over shoulder and pelvis ; $ conical studs over appendages .
references : debraga & rieppel ( 1997 ) ; lee ( 1997 ) . atw020924
images : scutosaurus karpinskii © urltoken , used with permission . atw020924 . revised mak120331
comments : ( 1 ) includes the subfamily pareiasaurinae nopcsa 1923 . oskar kuhn ( cotylosauria , p . 75 ) includes the following genera here : pareiasaurus , pareiasuchus ( dr carroll includes this genus under pareiasaurus ) , embrithosaurus , anthodon , scutosaurus , shihtienfenia , and nanoparia . von huene however places scutosaurus , shihtienfenia , and nanoparia each in their own subfamilies . cladistics substituted taxonomy with phylogeny , but this didn ' t help because hypotheses keep changing . lee ( 1997 ) argues that this group ( his clade ' d ' ) is paraphyletic and should include testudines ( or chelonia ) . the current consensus is that the pareiasaur origin is no longer valid , the two most popular turtle origin hypotheses now being thequite morphologically implausible crown archosauria of molecular phylogeny , and sauropterygian lepidosauromorpha , although we have gone for a more conservative prtocolophonoid origin
( 2 ) includes the
typical
pareiasaurs . the heads decorated by knobs and ridges . the teeth each have nine cusps . the upper margin of the ilium is flat , never vaulted . the surface of the back is armoured to varying degrees with bony scutes . some forms like the ancestral genus embrithosaurus had only weak armour ; while later forms like pareiasaurus , scutosaurus , and anthodon were well protected . the limb posture is advanced and upright to support the weight of the body ( graviportal ) .
stratigraphic range : lower to upper tapinocephalus zone , lower beaufort beds , of the karoo basin , south africa ; most common in the lower tapinocephalus zone ( lower to middle tapinocephalus zone in boonstra , 1969 pp26 , 32 , ; lower tapinocephalus to pristerognathus ( = old upper tapinocephalus ) zone in ward et al 2005 , supplem . info , range data ; both = capitanian age )
comments : [ 1 ] embrithosaurus is another huge early pareiasaur , a contemporary of bradysaurus . the skull is relatively deep and narrow . the body is lightly armoured with thin , smooth dermal scutes . this genus would seem to be ancestral to pareiasaurus . mak000124 .
[ 2 ] lee 1995 and jalil & janvier 2005 have three species , and tsuji & müller 2009 four as a scalloped paraphyletic series . four pareiasaur taxa ( bradysaurus baini , bradysaurus seeleyi , embrithosaurus ( nochelesaurus ) alexanderi and embrithosaurus schwarzi co - existing in the same environment seem excessive , so either this was intraspecies variation , gender difference , a stratigraphic and anagenetic ( chronospecies ) series , or niche partitioning mak120326
[ 3 ] cladistic analysis makes embrithosaurus as originally described paraphyletic , or in other words , an evolutionarily intermediate form between bradysaurus and higher pareiasaurs , and to avoid paraphyletic taxa the synonym nochelosaurus is restored . the present writer ( mak ) is opposed to the prohibition against representing transitional and ancestral species in accounts of earth history , as he feels that this is a too literal application of cladistic formalism , which is concerned with hypothesis testing , not with describing the actual evolutionary history of species in deep time . also , although phylogenetic nomenclature tends towards monospecific genera ( hence preference here of nochelesaurus over embrithosaurus , this creates an incongruency with taxonomy of plants , invertebrates , and quaternary and recent vertebrates , all of which ares based multi - species genera . for this reason , embrithosaurus is retained for all species in these evolutionary series ( ditto for bradysaurus and pareiasaurus ) ; in any case there is no restriction on paraphyletic genera in linnaean ( binomial ) taxonomy . mak120329
comments : the skull is large and deep , pointed at the front , and elevated in the jugal region . this species was originally made the type species of nochelosaurus by haughton and boonstra . boonstra later ( 1969 ) moved it into the genus bradysaurus , on the basis of the primitive tooth structure . kuhn however considers it belongs under embrithosaurus
comments : this species was made the type for
dolichopareia
. as the name indicates , the skull is long and narrow . this would seem to indicate a different lifestyle or diet to other parieasaurs . more recently , it has been used as the monotypal species for the genus nochelesaurus ( it is not clear what the status of embrithosaurus strubeni is , this may be a further transitional species ) the bottom skull is about 48 cm long , and was originally described as brachypareia watsoni , in cladistic analyses , this species is phylogenetically intermediate between bradysaurus seeleyi and embrithosaurus schwarzi mak000124 120329
graphics sketches of skulls of embrithosaurus angustus ( top ) and bradysaurus watsoni ( bottom ) . these two species are considered synonyms of embrithosaurus alexanderi , which indicates either artistic license or a great deal of individual variation , postmortem distortion , and so on . from kuhn 1969 mak120401
characters : the bony dermal plates are thin and smooth . the ulna is without an olecranon process . ( zittel & eastman 1932 p . 243 ) mak000124 .
anterior expansion of the iliac blade is flat rather than everted ; the two iliac blades are not parallel but diverge anteriorly , making an angle of approximately 40° with the sagittal plane ; pelvic symphysis is extremely thick , being almost half as deep as long .
- lee 1997
comments : the type species . this is the most advanced species of this genus , as indicated by the teeth , which have nine cusps ( in three groups of three ) . in cladistic analyses it is used as the monotypal species for the genus .
in spite of the large eye sockets , the eyeballs were small , as is shown by the size of the sclerotic ring that reinforced the cornea . skulls of pareiasaurs with the sclerotic ring preserved are very rare . as with all pareiasaurs , the surface of skull is uneven . being covered with horn - like bumps and ossicles ( bony plates ) of different sizes . the largest of these are usually on the cheek , parietal and nosal bones . sometimes there are large bosses on the frontal bones , and almost always on the lower jaw where it articulates with the skull . attached to the largest dermal ossicles are nodules or bosses of various sizes . the spaces between the dermal ossicles probably contained skin glands , which exuded specific muscus for keeping the skin moist , or even , as with some toads , poisonous warts to deter predators . ( vyatka palaeontological museum )
as with all pareiasaurs , the skull does not contain a separation of mouth and nose cavities . the animals had to bolt their food without chewing , because when the food was in mouth cavity , there was impossible to breathe . ( vyatka palaeontological museum )
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full reference : a . hartmann - weinberg . 1937 . pareiasauriden als leitfossilien [ pareiasaurids as guide fossils ] . problemy paleontologii 213 : 649 - 712
anthodon rossicus : pin 2212 / 2 , a skull ( without lower jaw ) . its type locality is kotelnich , vjatka river (
sarmin
series ) , pin 2212 , which is in a severodvinian terrestrial claystone / sandstone in the urpalov formation of the russian federation .
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further reading -  evolution der pareiasauriden .   -  trudy palaeontological institute academe nauk ,  sssr .   -  a .  p .  hartmann - weinberg  -  1933 .
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( hartmann - weinberg 1937 , lebedev 1987 ) ~ 1m in length was another basal pareiasaur , but without a fused tarsus and without a distinct acromion process , among other distinctions . a variety of specimens have been assigned to this ' wastebasket ' genus that will someday be assigned to new species or genera .
hartmann - weinberg ap 1933 . evolution der pareiasauriden : trudy palaeontological institute academe nauk , sssr , 1933 , n . 3 , p . 1 - 66 .
lee msy 1997 . pareiasaur phylogeny and the origin of turtles . zoological journal of the linnean society 120 : 197 - 280 .
owen r 1876 . descriptive and illustrated catalogue of the fossil reptilia of south africa in the collection of the british museum . london , british museum ( natural history ) .
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etymology : in reference to the triangular shape of its skull , resembling the greek letter “delta” , and the vyatka river ( tributary of the kama and volga ) , near where the specimen was found .
locality : kotel’nich locality , approximately 20 km on the western bank of the jjatka river south of the town of kotel ' nich , kotel’nichskii district , kirov region , eastern european russia .
age : upper capitanian to lowermost wuchiapingian , lowermost part of the upper tatarian age , lopingian subepoch , upper zechstein epoch , late permian .
smns 58317 : complete skull and lower jaw , incomplete postcranial including articulated presacral vertebrae , partial shoulder girdle including clavicle , humerus , and partial pelvic girdle .
rtmp 95 . 44 . 10 : complete skull and lower jaw , partially articulated and mostly complete postcranial , missing posterior caudals , right and left manus and some ribs .
umzc t1321 : complete skull and lower jaw , vertebral column missing posterior sacrals and caudals , ribs , complete shoulder girdle and both forelimbs , partial eft hind lime and osteoderms .
kmr ( unnumbered ) : a skull , lower jaw , vertebral column missing tail , and portions of both girdls , forelimbs and hind limbs .
there are at least 4 skulls of this animal , one in the palaeontological institute , moscow , the labels have been mixed up , but the catalogue number is probably pin 2212 / 3 , another is in the private collection of mr . terry manning , and 2 more are currently undergoing prepartion in the deparment of zoology , erindale collage , univeristy of toronto ( modesto , 1994 ) .
kpm / n11 / 99 : a fragment of skull belonging to a neonate , vertebral column missing posterior dorsal and caudal vertebrae , patial shoulder girdle and ribs , osteoderms .
kpm 232 : complete skull and lower jaw , complete postcranial missing posterior caudals and right hindlimb , as well as some phalanges from the right manus .
kpm 11 / 94 : some disarticulated skull elements including maxilla and quadratojugal , partially articulated dorsal vertebrae with osteoderms , disarticulated limb elements and interclavicle .
age : lower tatarian age , capitanian stage , upper guadelupian stage , middle zechstein epocn , late middle permian .
number : not given : a small partial skull of eather a neonate or embronyic animal , a larger adult specimen was also found .
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this article has been cited by the following publications . this list is generated based on data provided by crossref .
liu , jun bever , gabriel s . and angielczyk , kenneth 2018 . the tetrapod fauna of the upper permian naobaogou formation of china : a new species of elginia ( parareptilia , pareiasauria ) . papers in palaeontology , vol . 4 , issue . 2 , p . 197 .
müller , johannes bickelmann , constanze and sobral , gabriela 2018 . the evolution and fossil history of sensory perception in amniote vertebrates . annual review of earth and planetary sciences , vol . 46 , issue . 1 , p . 495 .
araújo , ricardo fernandez , vincent polcyn , michael j . fröbisch , jörg and martins , rui m . s . 2017 . aspects of gorgonopsian paleobiology and evolution : insights from the basicranium , occiput , osseous labyrinth , vasculature , and neuroanatomy . peerj , vol . 5 , issue . , p . e3119 .
benton , michael j . 2016 . the chinese pareiasaurs . zoological journal of the linnean society , vol . 177 , issue . 4 , p . 813 .
verrière , antoine brocklehurst , neil and fröbisch , jörg 2016 . assessing the completeness of the fossil record : comparison of different methods applied to parareptilian tetrapods ( vertebrata : sauropsida ) . paleobiology , vol . 42 , issue . 04 , p . 680 .
sobral , gabriela reisz , robert neenan , james m . müller , johannes and scheyer , torsten m . 2016 . evolution of the vertebrate ear . vol . 59 , issue . , p . 207 .
turner , morgan l . tsuji , linda a . ide , oumarou and sidor , christian a . 2015 . the vertebrate fauna of the upper permian of niger—ix . the appendicular skeleton ofbunostegos akokanensis ( parareptilia : pareiasauria ) . journal of vertebrate paleontology , vol . 35 , issue . 6 , p . e994746 .
from the kotel ' nich locality , kirov province , russia , is described in detail . the taxon is characterised by a distinctive pattern of dermal sculpture and the exaggerated embayment of the posterior skull roof , resulting in the dorsal exposure of the braincase . postcranially ,
shares some aspects of its morphology with basal pareiasaurs , including the osteoderm pattern . features such as the forward - slanting and pointed iliac blade are shared with stratigraphically younger , more derived forms . well - preserved material of the taxon spans a wide size - range , allowing an assessment of ontogenetic trends . a geometric morphometric analysis of the skull roof of
reveals an allometric increase in snout length and postorbital area , a result that can serve as a basis for examining morphological trends within pareiasaurs . a reassessment of pareiasauromorph relationships , using both parsimony and bayesian methods of phylogenetic inference , recovers similar topologies in both cases . four bayesian analyses were completed , with and without a gamma - shaped parameter and with and without the inclusion of autapomorphies . despite differing taxon and outgroup selection , the recovered topologies are similar to previous phylogenies of pareiasaurian relationships , with
pareiasaurian studies . part viii . the osteology and mycology of the locomotor apparatus b . fore limb
owenetta kitchingorum , sp . nov . , a small parareptile ( procolophonia : owenettidae ) from the lower triassic of south africa
researches on the structure , organisation , and classification of the fossil reptilia . ii . on pareiasaurus bombidens ( owen ) , and the significance of affinities to amphibians , reptiles and mammals
the vertebrate fauna of the upper permian of niger - vii . cranial anatomy and relationships of bunostegos akokanensis ( pareiasauria )
ein schädelrest von parasaurus geinitzi h . v . meyer ( reptilia , cotylosauria ) aus dem kupferschiefer ( perm ) von richelsdorf ( hessen )
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somatohelix is a genus of cuvaceous , spiralling tubular fossil , 2 – 7 mm wide and 3 – 14 cm long , from the ediacaran deposites of the south australia that was originally interpreted as a trace fossil ; a larger amount of better-preserved material since facilitated its reconstruction as the remains of an animal .

somatohelix

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here you will find one or more explanations in english for the word somatohelix . also in the bottom left of the page several parts of wikipedia pages related to the word somatohelix and , of course , somatohelix synonyms and on the right images related to the word somatohelix .
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this article has been cited by the following publications . this list is generated based on data provided by crossref .
evans , s . d . dzaugis , p . w . droser , m . l . and gehling , j . g . 2018 . you can get anything you want from alice ' s restaurant bed : exceptional preservation and an unusual fossil assemblage from a newly excavated bed ( ediacara member , nilpena , south australia ) . australian journal of earth sciences , p . 1 .
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droser , mary l . tarhan , lidya g . and gehling , james g . 2017 . the rise of animals in a changing environment : global ecological innovation in the late ediacaran . annual review of earth and planetary sciences , vol . 45 , issue . 1 , p . 593 .
darroch , simon a . f . boag , thomas h . racicot , rachel a . tweedt , sarah mason , sara j . erwin , douglas h . and laflamme , marc 2016 . a mixed ediacaran - metazoan assemblage from the zaris sub - basin , namibia . palaeogeography , palaeoclimatology , palaeoecology , vol . 459 , issue . , p . 198 .
buatois , luis a . and mángano , m . gabriela 2016 . the trace - fossil record of major evolutionary events . vol . 39 , issue . , p . 27 .
kirkland , christopher l . macgabhann , breandán a . kirkland , brian l . daly , j . stephen and macchiarelli , roberto 2016 . cryptic disc structures resembling ediacaran discoidal fossils from the lower silurian hellefjord schist , arctic norway . plos one , vol . 11 , issue . 10 , p . e0164071 .
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joel , lucas v . droser , mary l . and gehling , james g . 2014 . a new enigmatic , tubular organism from the ediacara member , rawnsley quartzite , south australia . journal of paleontology , vol . 88 , issue . 02 , p . 253 .
moczydłowska , małgorzata westall , frances and foucher , frédéric 2014 . microstructure and biogeochemistry of the organically preserved ediacaran metazoan sabellidites . journal of paleontology , vol . 88 , issue . 02 , p . 224 .
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macdonald , francis a . pruss , sara b . and strauss , justin v . 2014 . trace fossils with spreiten from the late ediacaran nama group , namibia : complex feeding patterns five million years before the precambrian–cambrian boundary . journal of paleontology , vol . 88 , issue . 02 , p . 299 .
chen , zhe zhou , chuanming meyer , mike xiang , ke schiffbauer , james d . yuan , xunlai and xiao , shuhai 2013 . trace fossil evidence for ediacaran bilaterian animals with complex behaviors . precambrian research , vol . 224 , issue . , p . 690 .
cai , yaoping hua , hong and zhang , xingliang 2013 . tube construction and life mode of the late ediacaran tubular fossil gaojiashania cyclus from the gaojiashan lagerstätte . precambrian research , vol . 224 , issue . , p . 255 .
clites , e . c . droser , m . l . and gehling , j . g . 2012 . the advent of hard - part structural support among the ediacara biota : ediacaran harbinger of a cambrian mode of body construction . geology , vol . 40 , issue . 4 , p . 307 .
cai , yaoping schiffbauer , james d . hua , hong and xiao , shuhai 2011 . morphology and paleoecology of the late ediacaran tubular fossil conotubus hemiannulatus from the gaojiashan lagerstätte of southern shaanxi province , south china . precambrian research , vol . 191 , issue . 1 - 2 , p . 46 .
university of california , riverside , department of earth sciences , 900 university ave . , riverside , 92521 , usa
ediacaran trace fossils are becoming an increasingly less common component of the total precambrian fossil record as structures previously interpreted as trace fossils are reinterpreted as body fossils by utilizing qualitative criteria . two morphotypes , form e and form f of glaessner ( 1969 ) , interpreted as trace fossils from the ediacara member of the rawnsley quartzite in south australia are shown here to be body fossils of a single , previously unidentified tubular constructional morphology formally described herein as
on a single surface and the few distinct characteristics preserved within this relatively indistinct fossil also provides the necessary data required to define a taphonomic gradient for this fossil . armed with this information , structures which have been problematic in the past can now be confidently identified as
based on morphological criteria . this suggests that the original organism that produced this fossil was a widespread and abundant component of the ediacaran ecosystem .
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yang , ben ; steiner , michael ; zhu , maoyan ; . . .
south australia is one of the world ' s richest ediacaran fossil localities . fossils are found in a rock unit called the ediacara member throughout the flinders ranges , most notably at ediacara conservation park , where the first discoveries were made , and at the remarkable site at nilpena , where serial excavations
the ediacaran fossil sites lie within the cryogenian to cambrian sedimentary sequence of the adelaide geosyncline ( cf . sprigg , 1952 ; see
for a concise recent summary of the geology of this region ) . late cryogenian tillites of the elatina formation ( a marinoan glacial equivalent ) are capped by a mixed carbonate and siliciclastic succession of the wilpena group . rare ediacaran fossils can be found in the wonoka formation ( such as
) , but the most spectacular occurrences occur within the ediacara member of the rawnsley quartzite .
due to a lack of suitable lithologies , there has been little in the way of radiometric dating of the australian succession . the ediacara member is considered to be a shallow - marine to deltaic succession , with fossils found in sandstone facies reflecting fair - weather to storm - wave - base depths (
the formal boundary for the base of the ediacaran period ( the basal ediacaran global stratotype section and point ; gssp ) can be found within the flinders ranges national park , at enorama creek . the gssp is located at the base of the cap carbonate immediately above the elatina glacial horizon , and is lithostratigraphically correlated to rocks elsewhere that have been dated to ~ 635 million years in age .
the basal ediacaran gssp , enorama creek , south australia . the gssp sits above the purple diamictite of the elatina formation , at the base of the cap carbonate nuccaleena formation .
the view from the razorback lookout , brachina trail , looking sw towards the abc ranges and wilpena pound , taking in the entire cryogenian to upper ediacaran sedimentary sequence from the trezona fm . at left , to the rawnsley quartzite ( tips of the hills in the far distance ) .
, earth sciences , university of california , riverside , 900 university ave , riverside , ca 92521 , aaron . sappenfield @ email . ucr . edu , droser , mary l . , department of earth sciences , university of california , riverside , 900 university ave , riverside , ca 92521 , and gehling , james , south australian museum , adelaide , 5000 , australia
session no . 245 - - booth # 103 paleontology : taphonomy , diversity & extinction ( posters ) oregon convention center : hall a 9 : 00 am - 6 : 00 pm , wednesday , 21 october 2009 geological society of america abstracts with programs , vol . 41 , no . 7 , p . 629
© copyright 2009 the geological society of america ( gsa ) , all rights reserved . permission is hereby granted to the author ( s ) of this abstract to reproduce and distribute it freely , for noncommercial purposes . permission is hereby granted to any individual scientist to download a single copy of this electronic file and reproduce up to 20 paper copies for noncommercial purposes advancing science and education , including classroom use , providing all reproductions include the complete content shown here , including the author information . all other forms of reproduction and / or transmittal are prohibited without written permission from gsa copyright permissions .
journal of paleontology ( 2011 ) 85 ( 2 ) : 256 - 265 .
aaron sappenfield , mary l . droser , james g . gehling ; problematica , trace fossils , and tubes within the ediacara member ( south australia ) : redefining the ediacaran trace fossil record one tube at a time . journal of paleontology ; 85 ( 2 ) : 256–265 . doi : urltoken
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the leptotyphlopidae ( commonly called slender blind snakes or thread snakes ) are a family of snakes found in north and south america , africa , and asia .

all are fossorial and adapted to burrowing , feeding on ants and termites .

two genera are recognized comprising 87 species .

leptotyphlopidae

hahn de . liste der rezenten amphibien und reptilien . anomalepididae , leptotyphlopidae , typhlopidae .
roze ja . contribución al conocimiento de los ophidios de las familias typhlopidae y leptotyphlopidae en venezuela .
morphology of the lower jaw and suspensorium in the texas blindsnake , leptotyphlops dulcis ( scolecophidia : leptotyphlopidae ) .
description of a second specimen of leptotyphlops parkeri ( squamata : leptotyphlopidae ) , with comments on its generic placement .
stephen r goldberg , reproduction in the western threadsnake , leptotyphlops humilis ( serpentes : leptotyphlopidae ) , from arizona : .
goldberg , s . 2003 . reproduction in the western thread snake , leptotyphlops humilis ( serpentes : leptotyphlopidae ) , from arizona .
morphology of the lower jaw and suspensorium in the texas blindsnake , leptotyphlops dulcis ( scolecophidia : leptotyphlopidae ) . - pubmed - ncbi
description of a second specimen of leptotyphlops parkeri ( squamata : leptotyphlopidae ) , with comments on its generic placement . - pubmed - ncbi
family leptotyphlopidae ( slender blind snakes and thread snakes ) .
animal diversity web . urltoken ( accessed on september 29 , 2004 ) .
from surinam with notes on the other surinam species of the genus ( leptotyphlopidae , serpentes ) . – notes on the herpetofauna of surinam v .
to cite this page : ast , j . 2003 .
leptotyphlopidae
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
slender blind snakes , thread snakes , or worm snakes : leptotyphlopidae .
grzimek ' s student animal life resource . . retrieved july 09 , 2018 from urltoken urltoken
adalsteinsson sa , branch wr , trape s , vitt lj , hedges sb . molecular phylogeny , classification , and biogeography of snakes of the family leptotyphlopidae ( reptilia , squamata )
pinto rr , passos p , portilla jrc , arredondo jc , fernandes r . taxonomy of the threadsnakes of the tribe epictini ( squamata : serpentes : leptotyphlopidae ) in colombia .
hedges , s . b . , 2011 : the type species of the threadsnake genus tricheilostoma jan revisited ( squamata , leptotyphlopidae ) . –zootaxa : 3027 , pp . 63 - 64
broadley , donald g . & broadley , sheila 1999 . a review of the arican wormsnakes from south of latitude 12°s ( serpentes : leptotyphlopidae ) . syntarsus 5 : 1 - 36
slender blind snakes , thread snakes , or worm snakes : leptotyphlopidae .
grzimek ' s student animal life resource . . encyclopedia . com . ( july 9 , 2018 ) . urltoken
broadley , donald g . & wallach , v . 2007 . a revision of the genus leptotyphlops in northeastern africa and southwestern arabia ( serpentes : leptotyphlopidae ) . zootaxa 1408 : 1–78 - get paper here
hahn d . e . & v . wallach , 1998 . comments on the systematics of old world leptotyphlops ( serpentes : leptotyphlopidae ) , with description of a new species . hamadryad 23 : 50 - 62
broadley d g . wallach v . 1997 . a review of the worm snakes of mozambique ( serpentes : leptotyphlopidae ) with the description of a new species . arnoldia zimbabwe 10 ( 11 ) : 111 - 119
despite blind snakes’ abundance ( more than 300 species ) and diversity , they are real strangers to herpetoculture . depending on your references , there are three families of blind snakes : the typhlopidae , leptotyphlopidae and anomalepidae .
adalsteinsson sa , branch wr , trape s , vitt lj , hedges sb . molecular phylogeny , classification , and biogeography of snakes of the family leptotyphlopidae ( reptilia , squamata ) . zootaxa . 2009 ; 2244 : 1–50 .
broadley d g . wallach v . 1997 . a review of the genus leptotyphlops ( serpentes : leptotyphlopidae ) in kwazulu - natal , south africa , with the description of a new forest - dwelling species . durban museum novitates 22 : 37 - 42 .
lehr , e . , wallach , v . , köhler , g . & aguilar , c . , 2002 : new species of tricolor leptotyphlops ( reptilia : squamata : leptotyphlopidae ) from central peru . –copeia : vol . 102 , # 1 , pp . 131 - 136
broadley , d . g . & watson , g . 1976 . a revision of the worm snakes of south - eastern africa ( serpentes : leptotyphlopidae ) . occ . pap . nation . mus . rhodesia bulawayo , ( bs ) 1976 : ( 8 ) : 465 - 510
leptotyphlopidae comprises about 90 species in two genera ( leptotyphlops and rhinoleptis ) . they have a tropical distribution , and are found in tropical north and south america , africa , and the middle east and northwest india . they live in a variety of habitats , from rainforests to semideserts , and from termite burrows to trees tops .
adalsteinsson , s . a . ; branch , w . r . ; trapé , s . ; vitt , l . j . & hedges , s . b . 2009 . molecular phylogeny , classification , and biogeography of snakes of the family leptotyphlopidae ( reptilia , squamata ) . zootaxa 2244 : 1 - 50 - get paper here
leptotyphlopidae is generally considered to be sister to the clade of typhlopidae and anomalepididae , although a recent analysis of molecular data strongly supports them as sister to typhlopidae . some morphological evidence , such as characteristics of the tongue , also support their sister relationship to typhlopidae . other authors have also questioned the monophyly of scolecophidia , based on other morphological data .
adalsteinsson , s . a . , branch , w . r . , trape , s . , vitt , l . j . & hedges , s . b . , 2009 : molecular phylogeny , classification , and biogeography of snakes of the family leptotyphlopidae ( reptilia , squamata ) . –zootaxa : vol . 2244 , pp . 1 - 50
families are listed in order according to fig 1 . for the taxonomy of families and subfamilies , we use adalsteinsson et al , [ 26 ] for anomalepididae and leptotyphlopidae , pyron and wallach [ 29 ] for gerrhopilidae , typhlopidae , and xenotyphlopidae , pyron et al [ 30 ] for booidea , and pyron et al [ 15 ] for alethinophidia . the number of species per clade was taken from the reptile database ( urltoken ) on 10 / 01 / 2015 . percentages of the number of species sampled do not include taxa not assigned to species status . paraphyletic taxa are included under their traditional family and / or subfamily . in the total cell for total number of species , the number not in parentheses equals the sum of the values in the table and the number in the parentheses equals the number returned when a search for serpentes is conducted in the reptile database . percentage for total number of species sampled is based on 3566 species .
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banks , r . c . , r . w . mcdiarmid , a . l . gardner , and w . c . starnes
checklist of vertebrates of the united states , the u . s . territories , and canada , draft ( 2004 )
banks , r . c . , r . w . mcdiarmid , and a . l . gardner
checklist of vertebrates of the united states , the u . s . territories , and canada
flores - villela , oscar / mccoy , c . j . , ed .
mcdiarmid , roy w . , jonathan a . campbell , and t ' shaka a . touré
snake species of the world : a taxonomic and geographic reference , vol . 1
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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a new species of fish , pseudoliparis swirei , published in zootaxa ( 4358 : 161 - 177 ) by gerringer , m . e et al . was voted among top 10 new species described in 2017 .
a new species of madagascan crickets described by mustafa ünal and george beccaloni in zootaxa was featured in a national geographic story . well done mustafa and george !
a new species of wolf spider , lycosa aragogi , is named after aragog—the famous fictional spider from “harry potter” book series by j . k . rowling . the new species is similar to the animatronic puppet version of the character used in the film “harry potter and the chamber of secrets” , which is actually based on a wolf spider . the naming of the new species is dedicated to the 20th anniversary of harry potter book series .
mariah o . pfleger , r . dean grubbs , charles f . cotton , toby s . daly - engel
identification of nipaecoccus ( hemiptera : coccomorpha : pseudococcidae ) species in the united states , with descriptions of nipaecoccus bromelicola sp . n . and the male of n . floridensis beardsley
the number of species increased from 10 , 711 to 10 , 793 , i . e . an increase of 82 species . 66 new species have been described , 9 species have been revalidated from synonymy and 16 subspecies were elevated to full species . . .
over the past 4 months , the number of species increased from 10 , 639 to 10 , 711 .
the number of species has grown from 10 , 544 in the may release to now 10 , 639 ( + 95 species ) .
overall , 212 new taxa have been added or changed their status or name .
the reptile database is a taxonomic database that provides basic information about all living reptile species , such as turtles , snakes , lizards , and crocodiles , as well as tuataras and amphisbaenians , but does not include dinosaurs .
currently there are more than 10 , 000 species and an additional 2 , 700 subspecies . this is making reptiles the largest vertebrate group after fish ( ~ 25 , 000 species ) and birds ( ~ 10 , 000 species ) , and significantly larger than mammals ( ~ 5 , 000 species ) or amphibians ( ~ 6 , 000 species ) .
the reptile database provides taxonomic information for the catalogue of life and the encyclopedia of life . our taxonomic information has also been used by genbank and many other resources and is the only comprehensive reptile database on the web .
the reptile database can be used to find all species within a certain geographic area ( e . g . all snakes of egypt ) . its collection of more than 2 , 500 images allow users to identify a species or at least get an idea how the species or genus may look like . more than 30 , 000 references provide a guide to further information .
this database is maintained by peter uetz ( database content ) and jakob hallermann , zoological museum hamburg ( new species and updates ) .
leptotyphlopids are small and very slender snakes , with a maximum recorded total length of 40cm but more typically attaining lengths of 15 - 20cm . like other blindsnakes , the eyes of anomalepedids are vestigial and covered by scales . the highly flexible dentaries posses four or five teeth , although teeth are absent form the immobile maxillae . the hyobranchium is y - shaped . the left lung is absent ( as it is in most snakes ) , as is a tracheal lung . some species retain rather well developed pelvic elements , including paired ilia , ischia , pubes , and femurs . females lack a left oviduct .
leptotyphlopids eat soft - bodied invertebrates , and secretions that fool termites allow them to take up residence in termite nests and eat eggs and larvae . leptotyphlopids have also been found in screech owl nests consuming arthropods - - the texas blindsnake leptotyphlops dulci is often brought to the nest as a prey item by these owls , although the snakes sometimes escape . nests with these resident blindsnakes enjoy a lower chick mortality rate , presumably because the snakes are eating parasitic arthropods . the feeding mechanism of leptotyphlops ( mandibular raking ) has been well characterized .
leptotyphlopids are oviparous , laying up to 13 eggs per clutch . female snakes have been observed coiled around their eggs , possibly demonstrating parental care .
from one point of view all living snakes may be regarded as degenerate quadrupeds , but some have gone further in having the appearance if not actuality of degenerated degenerates ! no reptile approaches so nearly the stage of being ' sans teeth , sans eyes , sans taste , sans everything ' as the members of the typhlops and leptotyphlops genera .
greene , h . 1997 . snakes : the evolution of mystery in nature . university of california press , berkeley .
heise , p . j . , l . r . maxson , h . g . dowling , and s . b . hedges . 1995 . higher - level snake phylogenies inferred from mitochondrial dna sequences and 12s rrna and 16s rrna genes . mol . biol . evol . 12 : 259 - 265 .
kley , n . j . and e . l . brainerd . 1999 . feeding by mandibular raking in a snake . nature 402 : 369 - 370 .
list , j . c . 1955 . external limb vestiges in leptotyphlops . herpetologica 11 : 15 - 16 .
mcdowell , s . b . 1972 . the evolution of the tongue of snakes , and its bearing on snake origins . in evolutionary biology ( t . dobzhansky , m , hecht , and w . steere , eds ) 6 : 191 - 273 . appleton - century - crofts , new york .
rieppel , o . 1988 . a review of the origin of snakes . evol . biol . 22 : 37 - 130 .
zug , g . 1993 . herpetology : an introductory biology of amphibians and reptiles . academic press , san diego
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 .
slender blind snakes , which are also known as thread snakes or worm snakes , are thin snakes with smooth , shiny scales . members of this family look much like blind snakes of the family typhlopidae and the early blind snakes of the family anomalepididae , since all have tube - shaped bodies that are about the same diameter from head to tail , and all have short heads with mouths that open downward instead of right on the front end of the head . species within the three blind snake families have small eyes and bodies that are covered with small scales that are the same size on the belly as they are on the sides and back . in most snakes , the belly scales , or ventrals , are noticeably larger . in the three blind snake families , only the scales on the snout are larger .
the slender blind snakes are different from the other two families in several ways . while all are slim , the slender blind snakes are the thinnest . the bodies of most species within this family are no wider than 0 . 2 inches ( 0 . 5 centimeters ) , and some are as little as 1 / 25th of an inch ( 1 millimeter ) wide . this gives them the appearance of moving string or thread . the largest species in the family , such as the western slender blind snake ( also known as the southwestern thread snake ) and the western thread snake , may reach more than 15 inches ( 38 centimeters ) in length , but most of the 93 species in the family are much smaller . the typical adult ranges from 4 to 10 inches ( 10 to 25 centimeters ) in length and no more than 0 . 05 ounces ( 1 . 4 grams ) in weight . another characteristic that sets the slender blind snakes apart from the other blind snakes—and indeed from all other snakes—is their teeth . slender blind snakes are the only snakes that have teeth on the lower jaw , but none on the upper jaw .
most slender blind snakes have backs that are all one color . they may be pink , light or dark brown , black , or gray . a few south american species are colorfully striped . the tails vary in length from about 2 percent of the overall body length to 19 percent , but most have tails that take up about 5 to 10 percent of the total length . the tail on a snake begins at the vent , which is a slitlike opening on the underside and toward the rear of the animal . the tail in almost all species of slender blind snakes has a sharp spine on the end . most have rounded snouts , but some have hooked and / or pointed snouts . in one unusual species , known as the western slender blind snake , the largest of its snout scales glows under ultraviolet light . inside the body , the skeleton of most slender blind snakes includes pieces of hip and upper leg bones that are leftover reminders of its ancient ancestors , which had working hips and legs . in boas and other snakes that have similar structures , the bony bits sometimes stick out of the body near the vent and look like small claws . these
claws
are called spurs . in slender blind snakes , the leftover bones usually do not poke through the skin . a few species have another odd feature in their skeletons : the top of the skull is missing .
slender blind snakes live in south , central , and north america , as well as africa and southwest asia . north american species live in mexico and the southwestern united states . certain species also live on the island of socotra in the northwestern indian ocean , in the west indies , and on islands off the coast of africa , mexico , and central america .
slender blind snakes eat small invertebrates ( in - ver - teh - brehts ) , which are insects and other animals without backbones . many of the species will eat almost anything , including insects such as caterpillars , fly maggots , beetles , cockroaches , and crickets , as well as spiders , harvestmen , which include daddy longlegs , and the many - legged centipedes and millipedes . most species , however , tend to prefer ants and termites . the snakes are able to find ant and termite hills by following the chemical trails that these insects leave on the ground as they travel to and from the nest . once the snake tracks down the ant or termite hill , it slithers inside and eats as much as it can . in ant hills , they especially like the eggs , larvae ( lar - vee ) , and pupae ( pyu - pee ) . ant eggs hatch into larvae , which are the maggotlike life stage of ants . eventually , the larvae transform into the motionless pupae stage before becoming adult ants . the slender blind snakes are able to jut out and pull in the lower jaw very quickly , which allows them to eat hundreds of eggs , larvae , and pupae in a very short time .
many animals avoid ant hills because these insects , which are very protective of their nests , can bite and sting . the slender blind snake , however , is able to defend itself . when attacked , the snake rolls into a ball and smears itself with its body ' s own ant repellant : a mixture of slime and feces . the ants shy away from the smelly mess , leaving the snake to return to its meal .
some animals need to live in a moist environment . the slender blind snakes , for example , burrow underground , hide beneath rocks , or slither into rotting logs or piles of dead leaves . one of the reasons a slender blind snake needs moisture is its very high surface - to - volume ratio . this is a mathematical formula that shows how much outer surface , or surface area , an animal has compared to the space , or volume , the entire animal takes up . because the slender blind snake is so long and thin , it has a great deal of surface area compared to its overall tiny body . if the snake were round like a ball rather than long and thin , its surface area would be much , much smaller . the outside weather has a greater effect on animals with higher surface - to - volume ratios , because a higher percentage of their total body volume is exposed . this means that they can dry out especially fast and may even die . for this reason , these animals frequently live in moist habitats or underground where their surroundings are damp .
slender blind snakes spend most of their time out of sight and below ground , sometimes as much as 49 feet ( 15 meters ) down , but they will crawl out of their burrows at night or after a heavy rain . they are able to dig through loose , sandy soil but cannot dig into harder ground , so they probably use other animals ' burrows or perhaps crawl along the paths of plant and tree roots when they are in tougher soil conditions . if they feel threatened , the snakes will squirm wildly , and those with tail spines will jab their attacker . some species may also stiffen up their bodies and play dead in an attempt to survive an attack .
although they are not sure , scientists suspect that all species lay eggs , rather than give birth to live baby snakes . the few snakes that have been studied mate in the spring and lay one to twelve eggs at a time in the summer .
physical characteristics : with their brownish pink to dark brown coloration , texas blind snakes look much like earthworms , except that the snakes have noticeable scales and lack the worm ' s segments . the snakes have a lighter colored , sometimes almost white , underside . also known as texas thread snakes , they have a long , thin body and a small head with eyes that appear as little more than tiny dark spots . adults range from 2 . 6 to 10 . 7 inches ( 6 . 6 to 27 centimeters ) long . the tail is short , just 5 to 6 percent of total body length , and has a spine at the end .
geographic range : texas blind snakes are found in the southwestern united states and northeastern mexico .
habitat : also known as a texas worm snake , the texas blind snake spends much of its time in the dirt , under rocks , or in some other hiding place . it can live in dry areas , including deserts and rocky mountainsides , but often chooses a spot near a water source .
diet : texas blind snakes most often eat ant larvae and pupae and termites , but they sometimes eat other insects and spiders . they always eat ant larvae and pupae whole , but they often refuse to eat the heads of termites and sometimes only chew the juices out of the back portion of the termite . once in a while , a small owl known as a screech owl will swoop down to snatch a texas blind snake and , keeping it alive , bring it back to its nest . there , the snake cleans out the nest by eating small invertebrates that might otherwise nibble on the owl .
behavior and reproduction : texas blind snakes live mainly underground but sometimes crawl out of their burrows at night or after a rain downpour . they are not especially good at slithering above ground and sometimes jab the tail spine into the ground to push off . males and females group together in the spring for the mating season . the female lays two to seven eggs in june or july and then coils around them . often , several females lay their eggs near one another . the eggs hatch in late summer into baby snakes about 2 . 6 to 3 inches ( 6 . 6 to 7 . 6 centimeters ) long .
texas blind snakes and people : texas blind snakes and people rarely encounter one another .
brazaitis , p . , and m . watanabe . snakes of the world . new york : crescent books , 1992 .
burnie , david , and don wilson , eds . the definitive visual guide to the world ' s wildlife . new york : dk publishing , 2001 .
grace , eric , ed . snakes . san francisco : sierra club books for children , 1994 .
greene , h . snakes : the evolution of mystery in nature . berkeley : university of california press , 1997 .
mcdiarmid , r . w . , j . a . campbell , and t . a . touré . snake species of the world . a taxonomic and geographic reference . vol . 1 . washington , dc : herpetologists ' league , 1999 .
mehrtens , john m . living snakes of the world in color . new york : sterling publishing , 1987 .
shaw , c . e . , and s . campbell . snakes of the american west . new york : alfred a . knopf , 1974 .
werler , j . e . , and j . r . dixon . texas snakes : identification , distribution , and natural history . austin : university of texas press , 2000 .
other interesting aspects of ant biology .
rice university . urltoken ( accessed on october 7 , 2004 ) .
photographs of blindsnakes .
comparative physiology and biomechanics lab at the university of massachusetts amherst . urltoken ( accessed on october 7 , 2004 ) .
texas blind snake .
national wildlife federation . urltoken ( accessed on october 7 , 2004 ) .
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, phyllodactylidae , scincidae y viperidae presentaron solamente una especie , donde cada una representa el 3 , 5 % de la riqueza total .
, typhlopidae , boidae , colubridae , elapidae , hydrophiidae , viperidae and crotalidae ( khan , 2004 , 2006 ) .
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discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
with over 3 , 500 species encompassing a diverse range of morphologies and ecologies , snakes make up 36 % of squamate diversity . despite several attempts at estimating higher - level snake relationships and numerous assessments of generic - or species - level phylogenies , a large - scale species - level phylogeny solely focusing on snakes has not been completed . here , we provide the largest - yet estimate of the snake tree of life using maximum likelihood on a supermatrix of 1745 taxa ( 1652 snake species + 7 outgroup taxa ) and 9 , 523 base pairs from 10 loci ( 5 nuclear , 5 mitochondrial ) , including previously unsequenced genera ( 2 ) and species ( 61 ) .
increased taxon sampling resulted in a phylogeny with a new higher - level topology and corroborate many lower - level relationships , strengthened by high nodal support values ( > 85 % ) down to the species level ( 73 . 69 % of nodes ) . although the majority of families and subfamilies were strongly supported as monophyletic with > 88 % support values , some families and numerous genera were paraphyletic , primarily due to limited taxon and loci sampling leading to a sparse supermatrix and minimal sequence overlap between some closely - related taxa . with all rogue taxa and incertae sedis species eliminated , higher - level relationships and support values remained relatively unchanged , except in five problematic clades .
our analyses resulted in new topologies at higher - and lower - levels ; resolved several previous topological issues ; established novel paraphyletic affiliations ; designated a new subfamily , ahaetuliinae , for the genera ahaetulla , chrysopelea , dendrelaphis , and dryophiops ; and appointed hemerophis ( coluber ) zebrinus to a new genus , mopanveldophis . although we provide insight into some distinguished problematic nodes , at the deeper phylogenetic scale , resolution of these nodes may require sampling of more slowly - evolving nuclear genes .
citation : figueroa a , mckelvy ad , grismer ll , bell cd , lailvaux sp ( 2016 ) a species - level phylogeny of extant snakes with description of a new colubrid subfamily and genus . plos one 11 ( 9 ) : e0161070 . urltoken
copyright : © 2016 figueroa 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 .
data availability : genbank accession numbers are contained within the paper . all other relevant data are within the paper and its supporting information files .
funding : this work was supported by national science foundation east asia & pacific summer institute ( oise - 1107819 ) ; university of new orleans college of sciences graduate student research grant ; university of new orleans department of biological science dissertation enhancement award ; university of new orleans dissertation improvement grant ; and university of new orleans latin american studies abroad program award . the funders had no role in the study design , data collection and analysis , decision to publish , or preparation of the manuscript .
we extracted genomic dna from tissue samples following the standard protocol provided for qiagen® dneasy kits . we sequenced six genes : 16s , c - mos , cyt - b , nd4 , nt3 , and rag - 1 . a list of the primers used , their source , and annealing temperatures are provided in s3 table . we aliquoted a 2 μl portion of each purified dna extract and combined it with gotaq green mastermix ( promega corp . ) , primers from respective gene , and deionized water to create a 10 μl reaction to be used in the polymerase chain reaction ( pcr ) . we placed all pcr reactions on a thermal cycler under the following protocol : 95°c for 2 min ; 95°c for 30 s ; 50°c for 30 s for 40 cycles ; 72°c for 1 : 15 min ; 72°c for 3–5 min ; and chilled at 4°c until taken off cycler . next , we cleaned the pcr products using 1 μl of exosap - it ( usb corp . ) per 10 μl of pcr product . we performed cycle sequencing on purified pcr products using 1 μl primer ( 10 μm ) , 2 μl template , and 5 μl deionized water along with a big dye terminator 3 . 1 ( amersham pharmacia biotech ) reaction premix for 50 cycles of 96°c for 10 s ; 45°c for 5 s ; and 60°c for 4 min and purified using a sephdex column , then used an abi 3130xl genetic analyzer to determine nucleotide sequences of each sample .
we aligned all sequences using the default parameters of the geneious alignment , and refined alignments using the default parameters of the muscle alignment [ 72 ] in the program geneious v4 . 8 . 4 ( urltoken ; [ 73 ] ) . we then edited alignments by eye and trimmed ambiguous end regions . for some genes , a few species had identical sequences with other taxa so we retained the first taxon in alphabetical order ( [ 15 ] ; s1 table ) . finally , we used geneious to concatenate all genes to create a supermatrix . this matrix contained 71 . 41 % of missing data ; however , previous studies have shown that missing data does not negatively influence topology , branch length estimates , and node support [ 15 , 23 , 40 , 41 ] . we deposited all sequences generated from this study in genbank ( s2 table ) . the final alignment is available in phylip format in s1 file .
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 : 3966804e - d532 - 4c52 - 92ac - becae776e434 . 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 .
abridged phylogeny on final dataset of 1652 snake species and seven outgroup taxa displaying higher - level relationships .
maximum - likelihood phylogenetic estimate based on 10 concatenated genes . tips represent families and sub - families . commonly recognized higher - level clades are labeled in all caps and bold . species classified as lamprophiidae incertae sedis are also shown since they did not place within a subfamily . node values represent shl support values . skeleton of the species tree is displayed on the left , colored and labeled as they appear in figs 2 – 10 .
maximum - likelihood phylogenetic estimate based on 10 concatenated genes . node values represent shl support values . seven outgroup taxa are not shown . colors of clades indicate their position in the overall tree , shown at left . newly sequenced taxa are highlighted in bold . skeleton of the species tree is displayed on the left with displayed subfamilies / families highlighted . letters denoted by i and ii represent parts of the tree where external branches do not connect to the part of the tree immediately preceding it . a ) anomalepididae , epictinae , leptotyphlopinae , gerrhopilidae , xenotyphlopidae , and typhlopinae . b ) asiatyphlopinae i , afrotyphlopinae ; madatyphlopinae , and asiatyphlopinae ii .
a ) aniliidae , tropidophiidae , calabariidae , candoiidae , sanziniidae , charininae , ungaliophiinae , erycidae , and boidae . b i ) cylindrophiidae + anomochilidae , uropeltidae , xenopeltidae , loxocemidae , and pythonidae . b ii ) bolyeridae , xenophidiidae , acrochordidae , xenodermatidae , and pareatidae .
a i ) viperinae . a ii ) azemiopinae and crotalinae . b ) crotalinae continued .
a ) homalopsidae , psammophiinae , buhoma procterae , prosymninae , pseudaspidinae , atractaspidinae , and aparallactinae . b i ) oxyrhabdium leporinum and lamprophiinae . b ii ) ditypophis sp . + micrelaps bicoloratus and pseudoxyrhophiinae .
a ) grayiinae , calamariinae , ahaetullinae subfam . nov . , and colubrinae . b i ) colubrinae continued . b ii ) colubrinae continued .
paraphyly at the lower - level of the tree emerged due to various reasons . for some clades paraphyly is well - established and confirmed here , more notably in brachyophidium , pseudotyphlops , rhinophis , and uropeltis in uropeltidae ( fig 3b i ) [ 15 , 41 , 53 , 90 ] ; ovophis and trimeresurus in respect to ovophis okinavensis + trimeresurus gracilis as basal to gloydius ( fig 4b ) [ 61 , 91 ] ; adelophis , amphiesma , atretium , nerodia , regina , thamnophis , tropidoclonion , and xenochrophis in natricinae ( fig 7a ) [ 15 , 68 , 92 , 93 ] ; and dipsas , geophis , and sibynomorphus in dipsadinae ( fig 7b ) [ 15 , 49 , 65 , 66 ] . additional taxa include : variable placement of morelia viridis ( fig 3b i ) [ 15 , 38 , 52 , 94 ] and bothrocophias campbelli ( fig 4b ) [ 95 ] ; and suta with parasuta ( fig 6b ) [ 15 , 96 ] . clearly , these clades require further inspection . on the other hand , we were able to rectify other paraphyletic taxa with strong support , specifically within colubrinae : boiga , chironius , coronella , crotaphopeltis , dasypeltis , dipsadoboa , hapsidophrys , and philothamnus , rhinechis , and scaphiophis .
subfamily ahaetuliinae subfam . nov . urn : lsid : zoobank . org : act : 22c47597 - 1def - 45a4 - abac - 11c4911557ad
content : four genera containing 56 species . ahaetulla ( 8 species ) , chrysopelea ( 5 species ) , dendrelaphis ( 41 species ) , and dryophiops ( 2 species ) .
etymology : from the sri lankan language sinhala , ahaetulla / ahata gulla / as gulla , meaning “eye plucker” or “eye picker” for belief that they pluck out the eyes of humans as accounted by the portuguese traveler joão ribeiro in 1685 ( as cited in [ 119 ] ) .
sister taxon : previously placed within colubrinae , ahaetuliinae forms a strong ( shl = 95 ) sister relationship with colubrinae , also weakly supported by pyron et al [ 15 ] .
distribution : members of this subfamily inhabit various habitats , but are mostly associated with forests distributed from pakistan , sri lanka and india , north to nepal and bangladesh , eastwards all throughout southeast asia to southern china , philippines , papua new guinea , and northeast australia .
remarks : the name ahaetulla has suffered from a tumultuous nomenclatural history [ 122 ] . in addition , members of these genera have historically been grouped with unrelated taxa based on absence or presence of hypapophyses [ 123 , 124 ] .
genus mopanveldophis gen . nov . urn : lsid : zoobank . org : act : 3b0cb6a0 - 1eec - 4512 - 9e77 - b105c22acabb
etymology : the generic nomen mopanveldophis is derived from the word “mopanveld” , the name of the type of habitat the specimens were found in , and the greek adjective ophis , meaning “snake” . this name refers to veld habitat distributed in southern africa , from the afrikaans word “field” , that is dominated by the mopane tree , colophospermum mopane , from the sechuana word “mopani” .
sister taxa : m . zebrinus is basal lineage to a clade including bamanophis + macroprotodon , placed within a larger clade of old world racers [ 15 , 40 , 107 , 108 ] .
distribution : currently recognized as endemic to northern namibia , africa [ 127 ] , but its range may extend into angola , africa [ 126 ] .
remarks : first described from a dead specimen collected in 1991 [ 125 ] , the species is currently known from only three specimens [ 126 ] . upon its description it was assigned to the genus coluber , presumably on basis of similar morphology , but then switched to hemerophis [ 24 , 106 ] with no published reasoning . schätti and trape [ 105 ] provide an account detailing the differences of bamanophis to other racer species , including m . zebrinus .
s1 fig . abridged phylogeny displaying higher - level relationships with all rogue taxa and incertae sedis species eliminated .
maximum - likelihood phylogenetic estimate based on 10 concatenated genes . tips represent families and sub - families . commonly recognized higher - level clades are labeled in all caps and bold . node values represent shl support values . skeleton of the species tree is displayed on the left , colored and labeled as they appear in fig 1 .
s1 file . data supermatrix comprising 1745 taxa representing 1652 snake species and 7 outgroup taxa , and 9523 base pairs from 10 loci .
s2 file . newick format maximum - likelihood phylogeny for 1745 taxa representing 1652 snake species and 7 outgroup taxa displayed in figs 2 – 10 .
s1 table . list of genbank accession numbers for 7 outgroup taxa and 1615 snake species .
two sequences were deleted during preliminary tree searches and 21 were identified as rogue taxa and pruned from the dataset leaving 1592 snake species from genbank in the tree . names represent species names as listed on the reptile database ( urltoken ) as of october 2015 . refer to s4 table for list of rogue taxa . taxa deleted during preliminary tree searches are highlighted in red , rogue taxa are highlighted in yellow , and sequences that were deleted because they were identical to other sequences are highlighted in green .
s2 table . list of taxa , institutional voucher numbers , and genbank accession numbers for tissue samples extracted and sequenced in this study .
tissue samples for boiga siamensis fmnh267726 , chrysopelea ornata lsuhc7158 , and psammodynastes pictus fmnh267940 were represented by clear chromatograms , but placed poorly in preliminary phylogenetic trees , so they were not included in the final data matrix . tropidolaemus subannulatus ku327425 was identified as a rogue taxon by roguenarok and was pruned from the dataset and thus , is not represented in the phylogeny .
s3 table . six loci , gene type , gene length , primer name , pcr annealing temperature and primer source .
s4 table . rogue taxa as identified by roguenarok web - server ( urltoken ) .
each taxon is associated with a raw improvement score ( r . i . s . ) , which represents the fraction of improvement in bootstrap support values throughout the tree when the selected taxon is pruned and all rogue taxa above it are also pruned . we performed one run and chose to sacrifice relatively lower node support values to maximize the number of taxa represented in the phylogeny . thus we elected to only prune taxa with r . i . s . greater than 0 . 8 , resulting in a total of 22 pruned taxa ( highlighted in bold ) .
we are indebted to the many researchers who donated tissue samples to museum collections and uploaded their sequences on genbank . for supporting this research in the way of loaning tissue samples , we would like to acknowledge curators and staff at the ambrose monell cryo collection ( amcc ) at the american museum of natural history , new york ( amnh ) , the california academy of sciences ( cas ) , the field museum of natural history ( fmnh ) , the natural history museum & biodiversity research center at the university of kansas ( ku ) , and the lsu museum of natural science collection of genetic resources ( lsumns ) and l . j . vitt and his nsf grants deb - 9200779 and deb - 9505518 , the museum of vertebrate zoology ( mvz ) , and the peabody museum of natural history yale university ( ypm ) .
we thank robin rowe at the university of new orleans w . m . keck conservation and molecular genetics laboratory for sequencing molecular samples , and caño palma biological research station in costa rica and david bickford at the national university of singapore for providing lab space during field research . we also thank several anonymous reviewers for providing useful comments that helped improve the manuscript .
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