epiglaea decliva , the sloping sallow moth , is a moth of the noctuidae family .

it is found in north america , where it has been recorded from quebec and maine to south carolina , west to kansas and north to alberta .

the habitat consists of barrens , thickets , woodlots and forests .

the wingspan is 40 – 50 mm .

the forewings range from orangish to reddish-brown or purplish-brown .

the reniform and orbicular spots are filled with a slightly darker colour , and have a pale outline .

the hindwings are dirty brownish-grey with wavy red terminal line .

adults are on wing from october to december in one generation per year .

the larvae feed on the leaves of malus , prunus and quercus species .

they have a brown to reddish-brown body and a mottled black head .

they reach a length of 50 mm when full-grown .

the larvae can be found from may to july .

the species overwinters as an egg .

epiglaea decliva

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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
adults - forewing light brownish - gray with several black longitudinal lines running length of wing ; pm line curves immediately below costa toward outer margin , then breaks into dark dots before continuing to inner margin ; diffuse dark patch at anal angle ; female has thick black line running through center of wing from base to midpoint of outer margin ; fringe checkered ; hindwing white with some brownish - gray shading along upper half of outer margin ; fringe white .
catabena lineolata has thinner / finer black lines , and a more northern distribution .
becker , v . o . & s . e . miller 2002 . the large moths of guana island , british virgin islands : a survery of efficient colonizers ( sphingidae , notodontidae , noctuidae , arctiidae , geometridae , hyblaeidae , cossidae ) . jl . lep . soc . 56 ( 1 ) : 44 ( pdf )
hampson , g . f . 1909 . catalogue of the lepidoptera phalænæ in the british museum . 8 : 236 ; pl . 78 , fig . 15 . ( treats terminellus as a syn . of vitrina )
herrich - schäffer , 1868 . correspondenz - blatt des zoologisch - mineralogischen vereins in regensburg . 22 : 147
walker , f . 1857 . noctuidæ . list of the specimens of lepidopterous insects in the collection of the british museum . 11 : 718
presence in california ; list of 3 specimen records with dates and locations ( u . of california at berkeley )
additions and corrections to the check list of the noctuoidea ( insecta , lepidoptera ) of north america north of mexico iii lafontaine j . d . & b . c . schmidt . 2015 . zookeys 527 : 127 - 147 .
contributed by robin mcleod on 16 february , 2008 - 2 : 10pm additional contributions by steve nanz , maury heiman , ron m . , randy hardy last updated 29 january , 2018 - 6 : 33pm
disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .
contributors own the copyright to and are solely responsible for contributed content . click the contributor ' s name for licensing and usage information . everything else copyright © 2003 - 2018 iowa state university , unless otherwise noted .
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 .
the author ' s name is made up of a string of letters , with no blanks , and multiple authors ' names are separated by a comma . spelling of author ' s name is based on the original publication . if there are more than three authors , only the names of the first two authors are shown , followed by
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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 .
all diacritic marks , hyphens , and apostrophes are eliminated , thus only the following characters are used : a to z , a to z , 0 to 9 , blank , comma , and opening and closing square brackets . although upper and lower cases are used for the convenience of human recognition , it is not case sensitive .
created by dicky sick ki yu 1997 - 2012 please send me information about errors and omissions ( contact information ) with supporting references , possibly with pdf or hard copy .
south of canada , compare with footpath sallow ( metaxaglaea semitaria ) and roadside sallow ( m . viatica ) - see photos of these and related species by jim vargo at mpg
presence in south carolina county distribution map ( john snyder , furman u . , south carolina )
field guide to moths of eastern north america charles v . covell , jr . 2005 .
on tree trunk with beer / brown sugar bait . submitted for pa bg record .
came to bait . november ( and december ) was unusually warm in 2015 . also saw one on 27 nov 2015 .
a sloping sallow moth in anne arundel co . , maryland ( 10 / 24 / 2016 ) . verified by roger downer / bamona . photo by tyler bell . ( mbp list )
a sloping sallow moth in harford co . , maryland ( 10 / 23 / 2015 ) . verified by roger downer / bamona . photo by dave webb . ( mbp list )
a sloping sallow moth in howard co . , maryland ( 10 / 30 / 2004 ) . photo by larry line . ( mbp list )
use of images featured on maryland biodiversity project is only permitted with express permission of the photographer .
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hodges , r . w . et al . , eds . 1983 . check list of the lepidoptera of america north of mexico . e . w . classey limited and the wedge entomological research foundation , london . 284 pp .
due to latency between updates made in state , provincial or other natureserve network databases and when they appear on natureserve explorer , for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data . please refer to
distribution data for u . s . states and canadian provinces is known to be incomplete or has not been reviewed for this taxon .
an occurrence where the species occurs or has occurred with potential for persistence or regular recurrence . minimally a collection ( generally must be an adult ) associated with suitable habitat . photodocumentation except from spread specimens is strongly discouraged but sometimes will suffice high quality occurrences will occupy at least hundreds of hectares and in some areas eos often cover tens of thousands of hectares . where forests are fragmented the metapopulation concept probably applies , similarly if the foodplant is very unevenly distributed within a large forest .
general habitat boundaries will sometime suffice for mapping , especially for conifer feeders . when plausible combine populations in proximate fragments as one metapopulation . see habitat and food comments fields and / or ask local experts for species - specific habitat information . be particularly careful with pine feeding zale species which may be species - specific . note that many of conifer feeders such as feralia , panthea and pine feeding zale do not wander far from their hostplants . in general then is the densitiy of pines ( or other appropriate conifers ) drops below about five per hectare in mixed forests such parts are probably best not mapped as habitat for these moths except over short distancs between more suitable patches .
if the habitat is occurring patchily within an extensive overall wooded landscape consider all patches within half the suitable habitat separation distance of at least one other as one metapopulation occurrence . however apply the unsuitable habitat distance across cleared lands .
inferred extent , for example based on one specimens in a light trap , is all suitable habitat within two kilometers of the collection point . if the habitat is more extensive than that there is almost no chance the resulting 1000 hectare circle will close to contain the entire occurrence . however when data are minimal conservative assumptions are warranted .
care has been taken not to include species that would likely violate any of the distances given , especially inferred extent . herminiine , smaller
deltoids
and some other small weak fliers are not included in this group . this group is mostly for noctuids that are either polyphagous or feed on a dominant , codominant , or at least frequent foodplant . it may not be appropriate for species feeding on grasses , forbs , shrubs or even tree that tend to occur as localized , well separated patches within a forest , although if such patches are fairly frequent they should work ( see alternate separation procudure ) .
use the generic guidelines for the application of occurrence ranks ( 2008 ) . the key for ranking species occurrences using the generic approach provides a step - wise process for implementing this method .
zoological data developed by natureserve and its network of natural heritage programs ( see local programs ) and other contributors and cooperators ( see sources ) .
lafontaine , j . d . and b . c . schmidt . 2010 . annotated check list of the noctuoidea ( insecta , lepidoptera ) of north america north of mexico . zookeys 40 : 1 - 239 .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
data provided by natureserve in collaboration with robert ridgely , james zook , the nature conservancy - migratory bird program , conservation international - cabs , world wildlife fund - us , and environment canada - wildspace .
patterson , b . d . , g . ceballos , w . sechrest , m . f . tognelli , t . brooks , l . luna , p . ortega , i . salazar , and b . e . young . 2003 . digital distribution maps of the mammals of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
data provided by natureserve in collaboration with bruce patterson , wes sechrest , marcelo tognelli , gerardo ceballos , the nature conservancy - migratory bird program , conservation international - cabs , world wildlife fund - us , and environment canada - wildspace .
iucn , conservation international , and natureserve . 2004 . global amphibian assessment . iucn , conservation international , and natureserve , washington , dc and arlington , virginia , usa .
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myrmecia flammicollis is an australian ant which belongs to the myrmecia genus .

it is native to australia .

they are usually distributed in the north of queensland .

they were described by william brown in 1953 .

myrmecia flammicollis resembles the m. petiolata .

the body back of the myrmecia flammicollis is black while prothorax is largely or completely an orange-red colour .

myrmecia flammicollis

the above specimen data are provided by antweb . please see myrmecia flammicollis for further details
the following myrmecia species groups are based on ogata & taylor ( 1991 [ 1 ] ) .
die gattung myrmecia ist unterteilt in 9 artengruppen damit sich die vielen arten dieser gattung leichter klassifizieren lassen .
the genus myrmecia is sub - divided in 9 species groups for an easier classification of the many species .
brown , w . l . , jr . 1953j . revisionary notes on the ant genus myrmecia of australia . bull . mus . comp . zool . 111 : 1 - 35 pdf
ogata , k . , taylor , r . w . ( 1991 ) ants of the genus myrmecia fabricius : a preliminary review and key to the named species ( hymenoptera : formicidae : myrmeciinae ) . journal of natural history , 25 , 1623–1673 .
the following information is derived from barry bolton ' s new general catalogue , a catalogue of the world ' s ants .
holotype , worker ,
the rocky scrub
around the headwaters of the rocky river , in the mcilwraith range , ne of coen , cape york rocky peninsula , queensland , australia , museum of comparative zoology .
paratype , 1 worker , lankelly creek , mcilwraith range , cape york peninisula , queensland , australia , museum of comparative zoology .
paratype , 2 workers , rocky scrub , mcillwraith range , cape york peninsula , queensland , australia , museum of comparative zoology .
paratype , 1 worker , rocky scrub , mcillwraith range , cape york peninsula , queensland , australia , queensland museum .
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from urltoken
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antweb is funded from private donations and from grants from the national science foundation , deb - 0344731 , ef - 0431330 and deb - 0842395 . c : 0
confused by a class within a class or an order within an order ? please see our brief essay .
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 .
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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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urn : lsid : biodiversity . org . au : afd . taxon : 63b90975 - 2edd - 4a90 - 8f13 - 61f8be7feafc
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urn : lsid : biodiversity . org . au : afd . taxon : f7b9376a - 762b - 4b90 - 8804 - 4de674639b68
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diadectomorpha are a clade of large reptile-like amphibians that lived in euramerica during the carboniferous and early permian periods and in asia during late permian ( wuchiapingian ) , and are very close to the ancestry of the amniota .

they include both large ( up to 2 meters long ) carnivorous and even larger ( to 3 meters ) herbivorous forms , some semi-aquatic and others fully terrestrial .

the diadectomorpha seem to have evolved during late mississippian times , although they only became common after the carboniferous rainforest collapse and flourished during the late pennsylvanian and early permian periods .

diadectomorpha

diadectomorpha have been nested in and out of the amniota . they’re in here .
links : diadectomorpha ; phylogeny of stegocephalians ( note figures of limnoscelis ) ; biology 356 .
diadectomorpha : ( limnoscelis + diadectidae ) + * . or limnoscelis + ( diadectidae + * ) .
kissel , r . a . , 2010 : morphology , phylogeny , and evolution of diadectidae ( cotylosauria : diadectomorpha ) . –phd dissertation , university of toronto , canada .
kissel , r . 2010 . morphology , phylogeny , and evolution of diadectidae ( cotylosauria : diadectomorpha ) . thesis ( graduate department of ecology & evolutionary biology university of toronto ) .
kissel , r . a . , 2010 : morphology , phylogeny , and evolution of diadectidae ( cotylosauria : diadectomorpha ) . –phd dissertation , university of toronto , canada , [ url : urltoken ]
“similarly , michel laurin ( 1996 ) uses the term in a cladistic sense to refer to only the most advanced reptile - like amphibians . thus his definition include the ( diadectomorpha and solenodonsauridae ) and the amniotes . ”
is the taxon basal to all other lepidosauromorpha including diadectomorpha , chelonia and lepidosauria . ( sorry , cut off from bottom of cladogram , fig . 1 ) . it was considered the most complete captorhinid from the late permian .
limnoscelis paludis ( diadectomorpha - limnosceliidae ) , early pemian , texas and new mexico an animal very close to the ancestry of amniotes . length about 1 . 5 meters life reconstruction , by dmitry bogdanov ( larger image ( wikimedia ) )
berman , d . s , s . s . sumida , and t . martens . 1998a . diadectes ( diadectomorpha : diadectidae ) from the early permian of central germany , with description of a new species . annals of carnegie museum 67 : 53 - 93 .
“however , michael benton ( 2000 , 2004 ) makes the anthracosaurs a paraphyletic order within the superorder reptiliomorpha , along with the orders seymouriamorpha and diadectomorpha , thus making the anthracosaurians the “lower” reptile - like amphibians . in his definition , the group encompass the embolomeri , chroniosuchia and possibly the family gephyrostegidae . ”
ty - jour ti - new cranial material of the rare diadectid desmatodon hesperis ( diadectomorpha ) from the late pennsylvanian of central colorado t2 - annals of the carnegie museum . vl - 64 is - 4 ur - urltoken pb - published by authority of the board of trustees of the carnegie institute , cy - [ pittsburgh ] : py - 1995 sp - 315 ep - 336 sn - 0097 - 4463 au - berman , david s au - sumida , stuart s er -
figure 1 . most parsimonious tree . the nodes are numbered in alphabetical order from the in - group node up . when a name is given , the letter designating the node is omitted but it is still reserved for this node . several clades are named in the text but could not be labeled on the figure . these include the following clades : k ( embolomeri ) , n ( seymouriamorpha ) , y ( apoda ) , z ( gymnophiona ) , aa ( salientia ) , ab ( anura ) and ad ( diadectomorpha ) . 4
synapsida ^ diadectomorpha ( ? ) sauropsida amphibian ( anamniote ) - grade taxon has ever managed to invade the niche of terrestrial herbivore , and a causal reason has been suggested for this . the endosymbionts required for digestion of high fiber vegetation must be obtained immediately after hatching fromterrestrial microbial decomposers ( modesto , 1992 ; zug , 1993 ; hotton and beerbower , 1994 ; hotton et al . , this volume ) . this constraint therefore prevents anamniote ( aquatically reproducing ) taxa from feeding on high - fiber vegetation . the fact that advanced diadectomorphs invaded this niche suggests that diadectomorphs had evolved the amniote egg .
@ article { bhlpart226640 , title = { new cranial material of the rare diadectid desmatodon hesperis ( diadectomorpha ) from the late pennsylvanian of central colorado } , journal = { annals of the carnegie museum . } , volume = { 64 } , copyright = { in copyright . digitized with the permission of the rights holder } , url = urltoken publisher = { [ pittsburgh ] : published by authority of the board of trustees of the carnegie institute , 1901 - } , author = { berman , david s and sumida , stuart s } , year = { 1995 } , pages = { 315 - - 336 } , }
relatively few large - scale phylogenetic studies on early tetrapods have been performed , and the origin of amniotes has generally been studied separately from the origin of lissamphibians . studies on the origin of amniotes usually only dealt with anthracosaurs ( carroll , 1970a ; gauthier et al . , 1988 ) because these have been presumed to be the closest relatives of amniotes . a consensus emerged that diadectomorpha is the sister - group of amniota whereas seymouriamorphs are among the next closest relatives of amniotes ( panchen and smithson , 1988 ; laurin and reisz , 1995 ) . research on the origins of lissamphibians concluded that
dissorophoids
were the closest relatives of lissamphibians ( trueb and cloutier , 1991 ) .
name : diadectes ( penetrating bite ) . phonetic : die - ah - deck - tees . named by : edward drinker cope - 1878 . synonyms : nothodon , empecocles , helodectes , empedias , chilonyx , bolbodon , diadectoides , animasaurus . classification : chordata , amphibia , reptiliomorpha , diadectomorpha , diadectidae . species : d . sideropelicus , d . lentus , d . tenuitectus , d . carinatus , d . sanmiguelensis , d . absitus . type : herbivore . size : between 1 . 5 and 3 meters long . known locations : usa , concentration in the texas red beds . time period : asselian through to the kungurian of the permian . fossil representation : many specimens are known allowing for accurate reconstruction .
comments : traditionally considered the most primitive diadectomorphs , although a recent analysis ( kissel 2010 ) makes them the sister group to the diadectidae . limnoscelid - like in appaerance , but tending to an omnivorous or herbivorous diet . only one species is known , a single late surviving form , essentially a
living fossil
co - existing with its more advanced and successful contemporaries . from wikipedia : tseajaia was described from a single , fairly complete specimen and was given its own family by robert l . carroll . it was originally thought to be an seymouriamorph ( moss 1972 ) additional finds allowing for a better taxonomic analysis indicate they belong in the diadectomorpha , as the sister group to the large and more derived diadectidae . tseajaia itself being a fairly generalized form , gives a reasonable indication of the build and looks of the closest relatives of the amniotes ( berman et al 1992 , kissel 2010 )
to study the impact of the topology ( that only represents a consensus of current phylogenies ) and internal branch lengths ( that are not known and difficult to estimate within nested clades that appear simultaneously in the fossil record ) , two strategies have been used . a second topology , incorporating relationships between the main clades from panchen and smithson ( 1988 ) was examined . this was done because the topology suggested from these authors represents the “traditional” point of view and because it presents a great number of differences with the preferred phylogeny ( fig . 2 ) . in this phylogeny , only the relationships between the large clades are different ; the topology within temnospondyls , “microsaurs , ” nectridea , baphetidae , seymouriamorpha , anthracosauria ( embolomeres and gephyrostegus ) , and cotylosauria ( diadectomorpha and amniota ) is identical to that of the main phylogeny ( fig . 1 ) . this reflects the fact that few phylogenies ( sometimes only one ) are present within these clades ( temnospondyls are an exception ) .
further reading - a new diadectes . - the american naturalist 12 : 565 . - e . d . cope - 1878 . - new or little known reptiles and amphibians from the permian ( ? ) of texas . - bulletin of the american museum of natural history 28 : 163 - 181 . - e . c . case - 1910 . - a description of the skulls of diadectes lentus and animasaurus carinatus . - american journal of science 33 ( 30 ) : 339 - 348 . - e . c . case & s . w . williston - 1912 . - early permian vertebrates from the cutler formation of the placerville area , colorado . - united states geological survey professional papers 503 - c : c1 - c46 - g . e . lewis & p . p . vaughn - 1965 . - diadectes ( diadectomorpha : diadectidae ) from the early permian of central germany , with description of a new species . - annals of carnegie museum 67 ( 1 ) : 53 - 93 . d . s . berman , s . s . sumida & t . martens - 1998 .
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( eds sumida , s . s . & martin , k . l . m . ) 353–398 ( academic , san diego , ( 1997 ) .
deposition , diagenesis and structures of the cheese bay shrimp bed , lower carboniferous , east lothian .
shrimp - bearing sedimentary successions in the lower carboniferous ( dinantian ) cementstone and oil shale groups of northern britain .
report on the fossil fishes collected by the geological survey of scotland from the shales exposed on the shore near gullane , east lothian .
from the viséan of east kirkton , west lothian , scotland and the amniote stem .
we thank s . finney for preparation of the specimen and assistance with photographs . this work was supported by an nerc grant ( to j . a . c . ) .
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$ more derived axis - atlas complex ; $ sacrum with at least 2 vertebrae , $ robust claws on pes ; $ loss of lateral line system .
links : phylogeny of stegocephalians ; amphibia et reptilia ngiana ( chinese ) ; v the four other mass extinctions . atw020623 .
cotylosauria : stem amniota + * : tseajaia + ( limnoscelis + diadectidae ) .
1 . 5 - 3 m ; some fully terrestrial ;
swollen
neural arches ; some ( diadectes ) herbivorous .
comments : primitive , small to large , carnivorous diadectomorphs . these animals must have lived a very similar life to ophaicodontid , varanopsid , and sphenacodontid pelycosaurs , which they resembled in overall size and shape ; presumably relied on niche partitioning . teeth retained labyrinthodont infolding of the enamel , and were pointed and slightly recurved at the tip ( carroll 1967 , via wikipedia ) traditionally divided into two genera , limnoscelis and limnostygis , although these may be synonyms
comments : highly successful clade of large herbivores , include the largest animals of their time and teh first exploitation of vegetable matter by tetrapods . previously tentrapods ( amphibians and reptiles ( ) had been carnivores only , feeding on insects , crustacea and other invertebrates , fish , and smaller tetrapods . this means that eco \ ystems were inefficent ( becaise of long food chains , and tied to water or water margins . while edphosaurs were also herbivores , they never challenged the supremacy of the diadectids . diadectid extinction at the end of the early permian allowed caseid pelycosaurs to replace them as the large herbivores ; they were in turn replaced by herbivousrous therapsids ( dinocephalia and anomodonts ) later in the permian .
left : life reconstruction of diasparactus zenos , by dmitry bogdanov . length 1 . 35 metres . wikipedia , gnu free documentation / creative commons attribution below left : the skulls of orobates , oradectes , silvadectes and diadectes from kissel 2010 , via david peters , reptile evolution . scale bar = 2 cm . below right : cladogram from kissel 2010 via wikipedia :
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parent taxon : cotylosauria according to m . laurin and r . r . reisz 1995
see also colbert 1946 , gauthier et al . 1989 , gauthier et al . 1988 , haughton and brink 1954 , huene 1954 , kissel and lehman 2002 , lewis and vaughn 1965 , martens et al . 2005 , olson 1947 , price 1947 and romer 1956
scientists aim to describe a single ' tree of life ' that reflects the evolutionary relationships of living things . however , evolutionary relationships are a matter of ongoing discovery , and there are different opinions about how living things should be grouped and named . eol reflects these differences by supporting several different scientific ' classifications ' . some species have been named more than once . such duplicates are listed under synonyms . eol also provides support for common names which may vary across regions as well as languages .
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levis & vaughn , 1965 ; l . perm . swnam . | | - - †
genera et species indet . ( berman & henrici , 2003 ) ] l . perm . ceu . ` - - + ? - †
liu , j . & bever , g . s . , 2015 : the last diadectomorph sheds light on late palaeozoic tetrapod biogeography . –biology letters : vol . 11 , # 5 , in press [ doi : 10 . 1098 / rsbl . 2015 . 0100 ]
berman , d . s . & henrici , a . c . , 2003 : homology of the astragalus and structure and function of the tarsus of diadectidae . –journal of paleontology : vol . 77 , # 1 , pp . 172 - 188
carroll , r . l . , 1988 : vertebrate paleontology and evolution . –w . h . freeman and company , new york , 1988 , 698 .
carroll , r . l . , 1988 : appendix . 594 - 648 . in carroll , r . l . 1988 : vertebrate paleontology and evolution . –w . h . freeman and company , new york , 1988 , 698 .
carroll , r . l . , 1967 : a limnoscelid reptile from the middle pennsylvanian . –journal of paleontology : vol . 41 , # 5 , pp . 1256 - 1261
clack , j . a . , 1998 : a new early carboniferous tetrapod with a mélange of crown - group characters . –nature : vol . 394 , 2 july , pp . 66 - 69
laurin , m . & reisz , r . r . , 1992 : a reassessment of the pennsylvanian tetrapod romeriscus . –journal of vertebrate paleontology : vol . 12 , # 4 , pp . 524 - 527
laurin , m . & reisz , r . r . , 1997 : a new perspective on tetrapod phylogeny . 9 - 59 in sumida , s . s . & martin , k . l . m . , 1997 : amniote origins : completing the transition to land . –academic press , san diego , 1997 , pp . x - 510
lee , m . y . s . & spencer , p . s . , 1997 : crown - clades , key characters and taxonomic stability : when is an amniote not an amniote ? 61 - 84 in sumida , s . s . & martin , k . l . m . , 1997 : amniote origins : completing the transition to land . –academic press , san diego , 1997 , pp . x - 510
paton , r . l . , smithson , t . r . , & clack , j . a . , 1999 : an amniote - like skeleton from the early carboniferous of scotland . –nature : vol . 398 , 8 april , pp . 508 - 513
ruta , m . & milner a . r . & coates , m . i . , 2000 : the scottish carboniferous tetrapod caerorhachis bairdi –journal of vertebrate paleontology : vol . 20 , # 3 , suppl . to # 3 ,
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limnoscelis ( new mexico and colorado ) and limnostegis ( nova scotia ) ; williston ( 1911b ) , carroll ( 1967 ) , fracasso ( 1980 ) , berman and sumida ( 1990 ) , and berman ( 1993 ) .
desmatodon ( colorado , new mexico , and pennsylvania ) and diasparactus ( new mexico ) ; case ( 1908 ) , case and williston ( 1913 ) , romer ( 1952 ) , vaughn ( 1969 , 1972 ) , fracasso ( 1980 ) , berman ( 1993 ) , eberth and berman ( 1993 ) , and berman and sumida ( 1995 ) .
limnosceloides ( new mexico and west virginia ) limnoscelops ( colorado ) , and limnoscelid ( utah ) ; romer ( 1952 ) , vaughn ( 1962 ) , lewis and vaughn ( 1965 ) , langston ( 1966 ) , and berman ( 1993 ) .
tseajaia ( new mexico and utah ) ; vaughn ( 1964 ) , moss ( 1972 ) , and berman ( 1993 ) .
figure 3 . late carboniferous distributions of the diadectomorph families limnoscelidae ( l ) and diadectidae ( d ) . localities , latitidue and longitude may be found in appendix 1 .
( 1860 ) , geinitz and deichmuller ( 1882 ) , olson ( 1947 , 1967 , 1975 ) , langston ( 1963 ) , lewis and vaughn ( 1965 ) , berman ( 1971 , 1993 ) , and berman and martens ( 1993 ) .
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diadectes was a large reptile - like tetrapod that lived in the late carboniferous and early permian periods . fossils of diadectes have been discovered in new mexico , colorado , oklahoma and texas ( usa ) and one species described from germany .
diadectes latibuccatus cope 1878a ; empecocles latibuccatus ( cope 1878a ) ; empedias latibuccatus ( cope 1878a ) ; bolosaurus rapidens cope 1878a ; chilonyx rapidens ( cope 1878a ) ; empedocles alatus cope 1878a ; empedias alatus ( cope 1878a ) ; diadectes molaris cope 1878b ; empedocles molaris ( cope 1878b ) ; empedias molaris ( cope 1878b ) ; diadectes phaseolinus cope 1880b ; empedias phaseolinus ( cope 1880b ) ; helodectes paridens cope 1880b ; helodectes isaaci cope 1880b ; empedias fissus cope 1883 ; diadectes biculminatus cope 1896 .
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the phylogeny of early groups of amniotes has been in a state of flux in the last few years , but the first phylogeny based on a data matrix was published in the eighties ( gauthier et al . , 1988 ) . the origins of mammals and saurians from early synapsids and early diapsids , respectively , has been relatively non - controversial , but the origin of turtles has been problematic . the most widespread view , at least until the last few years , was that the closest extinct relatives of turtles were the captorhinids ( gaffney & mc kenna , 1979 ; gaffney & meylan , 1988 ; gaffney , meylan , & wyss , 1991 ; gauthier et al . , 1988 ) . the study of gauthier et al . ( 1988 ) is probably the best documented study supporting close affinities between captorhinids and turtles :
this study is also significant in giving the first phylogenetic definitions for the names of many amniote taxa . in this tree , parareptiles are not closely related to turtles , but gauthier et al . ( 1988 ) were not confident in this part of their tree ( which is why they did not erect a formal parareptilia ) .
laurin & reisz ( 1995 ) suggested that many of the taxa that gauthier et al . ( 1988 ) considered parareptiles represented early relatives of turtles . furthermore , laurin & reisz ( 1995 ) suggested that mesosaurs were the closest known relatives of reptiles :
this study used a terminology derived from olson ( 1947 ) . unfortunately , this terminology includes some junior synonyms ( according to the rules of priority suggested by de queiroz & gauthier in 1990 , 1992 , and 1994 ) of the taxa defined by gauthier et al . ( 1988 ) and the terminology used in the main phylogeny ( in the page on amniota ) corrects this .
lee ( 1995 , 1996 ) studied in detail the phylogeny of the presumed relatives of turtles , and argued that pareiasaurs are the closest known relatives of turtles , rather than procolophonids ( as argued by laurin & reisz , 1995 ) . furthermore , he believed that pareiasaurs are the stem - group of turtles ( some pareiasaurs are more closely related to turtles than to other pareiasaurs ) . his phylogeny is as follows :
all the studies mentioned above agree that turtles are the sister - clade of saurians among extant amniotes . however , the old idea that turtles are closely related to placodonts ( broom , 1924 ) has been revived recently ( rieppel , 1994 , 1995 ; rieppel & debraga , 1996 ; debraga & rieppel , 1997 ) . if this theory is accurate , turtles are saurians and are more closely related to lepidosaurs than to archosaurs .
an exhaustive list of autapomorphies of various amniote taxa cannot be provided here , but a sample of these characters can be given . the list given below assumes that the phylogeny presented at the beginning of the page on amniota is essentially correct . phylogenetic definitions , when available , are also given for the main taxa . amniote classification can be summarized as such :
parts of the phylogeny are poorly documented ( most of the unnamed clades ) and will not be discussed below . furthermore , if the suggestion of modesto ( 1999 ) that mesosaurs are anapsids is accepted , this classification and the list of apomorphies will have to be modified .
amniota is defined as
the most recent common ancestor of extant mammals and reptiles , and all its descendants
( gauthier et al . , 1988 ) . it is divided into two stem - based taxa : synapsida ( mammals and their extinct relatives ) and sauropsida ( reptiles and their fossil relatives ) . the autapomorphies of amniota are listed in the
characteristics
section of the tree of life page .
sauropsida is defined as
reptiles plus all other amniotes more closely related to them than they are to mammals
( gauthier , 1994 ) . the characters supporting sauropsida include the following :
presence of a single coronoid . synapsids and the diadectomorph limnoscelis have two coronoid elements . the coronoids are bones on the dorsomedial surface of the lower jaw .
supinator process parallel to humeral shaft and separated from it by a groove . in synapsids and diadectomorphs , the supinator process is strongly angled relative to the shaft .
presence of a single pedal centrale . two centralia were present in the tarsus ( ankle ) of synapsids and diadectomorphs .
reptilia is defined as
the most recent common ancestor of extant turtles and saurians , and all of its descendants
( gauthier et al . , 1988 ) . characters supporting reptilia include :
tabular small . the tabular ( a bone on the posterolateral corner of the skull table ) of early synapsids and diadectomorphs is large , but reptiles have only a small tabular , when it is present .
suborbital foramen present ( fig . 1b - d ) . the suborbital foramen is a small hole near the lateral edge of the palate , between the pterygoid , palatine , and ectopterygoid ( or jugal , when the ectopterygoid is absent ) . this structure was not found in early synapsids and diadectomorphs ( fig . 1a ) .
supraoccipital anterior crista present . the supraoccipital of mesosaurs , synapsids , and diadectomoprhs lacks anterior parasagittal flanges . the supraoccipital of reptiles has a paired anterior parasagittal flange called an anterior crista .
supraoccipital plate narrow . the supraoccipital is a bone in the back of the braincase . the supraoccipital plate of mesosaurs , synapsids , and diadectomorphs is broad and extends farther laterally than the postparietal .
figure 1 . amniote skulls in palatal view . a , cotylorhynchus , a lower permian synapsid ; b , captorhinus , a lower permian romeriid ; c , procolophon , a triassic anapsid ; d , proganochelys , the oldest known turtle ( upper triassic ) . redrawn from a , laurin & reisz ( 1995 ) ; b , heaton ( 1979 ) ; c , carroll & lindsay ( 1985 ) ; and d , gaffney ( 1990 ) . abbreviations : bc , braincase ; ec , ectopterygoid ; m , maxilla ; n , nasal ; pa , palatine ; pm , premaxilla ; ps , parasphenoid ; pt , pterygoid ; q , quadrate ; qj , quadratojugal ; s , stapes ; sq , squamosal ; v , vomer .
anapsida is defined as
extant turtles , and all other extinct taxa that are more closely related to them than they are to other [ extant ] reptiles
( gauthier et al . , 1988 ) . characters supporting anapsida include :
foramen orbito - nasale enclosed between prefrontal , lacrimal , and palatine . there is no foramen between these bones in diapsids , synapsids , and diadectomorphs .
anterior lateral maxillary foramen distinctly larger than other foramina . in romeriids , mesosaurs , synapsids , and diadectomorphs , several small foramina ( holes ) may be present on the lateral surface of the maxilla , but none of these foramina is much larger than the others .
quadratojugal expanded dorsally ( fig . 2c , d ) . the quadratojugal of romeriids , mesosaurs , synapsids , and diadectomorphs is a low bone narrowly exposed on the lateral surface of the cheek ( fig . 2a , b ) .
figure 2 . amniote skulls in lateral view . a , cotylorhynchus , a lower permian synapsid ; b , captorhinus , a lower permian romeriid ; c , procolophon , a triassic anapsid ; d , proganochelys , the oldest known turtle ( upper triassic ) . redrawn from a , laurin & reisz ( 1995 ) ; b , heaton ( 1979 ) ; c , carroll & lindsay ( 1985 ) ; and d , gaffney ( 1990 ) . abbreviations : bc , braincase ; e , epipterygoid ; f , frontal ; l , lacrimal ; m , maxilla ; n , nasal ; p , parietal ; pm , premaxilla ; po , postorbital ; pof , postfrontal ; prf , prefrontal ; pt , pterygoid ; q , quadrate ; qj , quadratojugal ; sq , squamosal ; st , supratemporal ; t , tabular .
temporal emargination bordered by quadratojugal and squamosal ( fig . 2c , d ) . the temporal emargination supports a tympanum in turtles , and the presence of a slender stapes in several anapsids suggests that a tympanum appeared fairly early in this group . there is no temporal emargination in romeriids , mesosaurs , and synapsids ( fig . 2a , b ) .
jaw articulation anterior to occiput ( fig . 1c , d ) . the jaw articulation of romeriids , mesosaurs , synapsids , and diadectomorphs is at the level of the occiput ( fig . 1a , b ) .
edentulous ectopterygoid . the ectopterygoid bears a shagreen of small denticles in romeriids , early synapsids , and diadectomorphs .
stapedial dorsal process unossified . the stapes of most early romeriids , mesosaurs , synapsids , and diadectomorphs has an ossified dorsal process .
sacral ribs with narrow distal contact . the sacral ribs of romeriids , mesosaurs , and synapsids are broad distally and a broad contact between successive ribs leaves only a small gap . the sacral ribs of anapsids are more slender and contact each other distally .
iliac blade dorsally expanded . the iliac blade of romeriids , mesosaurs , and synapsids has a long , low posterodorsal process and lacks an anterior expansion . the iliac blade of anapsids has a dorsal expansion directly above the acetabulum , a short anterior process , and it is short posteriorly .
figure 3 . skulls of early amniotes in dorsal view . a , cotylorhynchus , a lower permian synapsid ; b , captorhinus , a lower permian romeriid ; c , procolophon , a triassic anapsid ; d , proganochelys , the oldest known turtle ( upper triassic ) . redrawn from a , laurin & reisz ( 1995 ) ; b , heaton ( 1979 ) ; c , carroll & lindsay ( 1985 ) ; and d , gaffney ( 1990 ) . abbreviations : bc , braincase ; f , frontal ; j , jugal ; l , lacrimal ; m , maxilla ; n , nasal ; p , parietal ; pa , palatal ; pm , premaxilla ; po , postorbital ; pof , postfrontal ; pp , postparietal ; prf , prefrontal ; qj , quadratojugal ; sq , squamosal ; st , supratemporal ; t , tabular .
procolophonia is defined as
the most recent common ancestor of pareiasaurs , procolophonids , and testudines ( chelonia ) , and all its descendants
( laurin & reisz , 1995 ) . it is diagnosed by several autapomorphies , including the following :
pineal foramen close to fronto - parietal suture ( fig . 3c ) . the pineal foramen of most other amniotes is located close to the mid - length of the interparietal suture , or slightly anterior to this ( fig . 3a , b ) .
tabular absent ( fig . 3c , d ) . most other amniotes have a tabular ( fig . 3a ) , but this bone has been lost convergently in captorhinids ( fig . 3b ) .
cranio - quadrate space large . in most other amniotes , the cranio - quadrate space ( the space between the braincase and the quadrate ramus of the pterygoid ) is narrow and the paroccipital process and the quadrate ramus of the pterygoid converge posterolaterally ( fig . 1a , b ) . in procolophonoids , pareiasaurs , and turtles , the cranio - quadrate space is wide and the paroccipital process is parallel to the quadrate ramus of the pterygoid ( fig . 1c , d ) .
pterygoid palatal ramus not reaching level of internal naris ( fig . 1c , d ) . the palatal ramus of millerettids , romeriids , mesosaurs , and early synapsids extends anteriorly medial to the internal naris ( fig . 1a , b ) .
transverse flange of pterygoid directed anterolaterally ( fig . 1c , d ) . the transverse flange of other amniotes extends transversely or posterolaterally from the area of the basicranial articulation ( fig . 1a , b ) .
cultriform process short ( fig . 1c , d ) . the cultriform process of other amniotes is long ( fig . 1a , b ) .
paroccipital process antero - posteriorly expanded . the paroccipital process of most other amniotes is a broad , thin , vertical flange .
quadrate condyle articular surfaces nearly flat and antero - posteriorly short . in other amniotes , the articular surfaces of the quadrate are strongly convex and short .
astragalus and calcaneum ( two ankle bones ) sutured or fused to each other .
romeriida is defined as
extant saurians , and all other taxa that are more closely related to them than they are to anapsids
( gauthier et al . , 1988 ) . romeriida possesses the following autapomorphies :
postorbital separated from supratemporal ( fig . 3b ) . the postorbital contacts the supratemporal in most anapsids ( fig . 3c ) , in mesosaurs , in early synapsids ( fig . 3a ) , and in diadectomorphs .
supratemporal small ( fig . 3b ) . the supratemporal is large in most other groups of amniotes ( fig . 3a , c ) .
caniniform maxillary tooth present . romeriids have a large anterior maxillary tooth ( fig . 2b ) . most other early amniotes have a relatively homodont dentition and no anterior maxillary tooth is much larger than the other teeth ( fig . 2a , c ) .
quadrate anterior process short ( fig . 1b ) . the anterior process of the quadrate extends anteriorly along more than 55 % of the quadrate ramus of the pterygoid in most other amniotes ( fig . 1a , d ) .
anterior pleurocentra keeled ventrally . the anterior pleurocentra of other amniotes is rounded ventrally .
metapodials overlapping . the proximal heads of the metapodials of other amniotes barely contact each other . the metapodials of
protorothyridids
and diapsids are expanded proximally and overlap the metapodial lateral to them .
we thank miss diane scott for scanning the figures of amniote skulls , and ms . patricia lai for proof - reading this page .
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scientists have found evidence that about 2 . 4 million years ago a gene regulating jaw muscles mutated and may have led to the more graceful human jaw we see today . which statement most closely explains the link between jaw size and hominid evolution
some characteristics of recently discovered organism are listed in the table . based on the given characteristics , this organism would be classified in which kingdom
the classification of four birds are listed . based on the classification which two birds are most closely related
scientists are studying the evolutionary history of a group of plants in the us . what information about the organisms best helps the scientists to determine the evolutionary relationships among them
many scientists think that the mitochondria in eukaryotic cells originated in independent prokaryotes . which characterisitc of mitochondira most strongly supports this conclusion
modern classification of the related organisms in a diagram . which two groups are most closely related
- like taxon first crawled out on dry land . then , according to the widely accepted paradigm , certain lineages returned to the water while others ventured forth onto higher and drier environs .
( lrt , 1033 taxa ) documents a bushier conquest of land , occurring in at least seven devonian waves until the beachhead was secured by our reptile ancestors .
dr . jennifer clack and her team have shown us that fish / amphibians can have limbs ( acanthostega and ichthyostega ) and not be interested in leaving the water . that comes later and later and , well , seven times all together .
figure 1 . colosteus relatives according to the lrt scaled to a common skull length . their sizes actually vary quite a bit , as noted by the different scale bars . only pholidogaster and colosteus are taxa in common with traditional colosteid lists .
had small limbs with toes . the smaller , but equally scaly and eel - like
, reduced those limbs to vestiges , showing they were not that important for getting around underwater in that wriggly clade . neither shows signs of ever leaving the water and phylogenetically neither led to the crawling land tetrapods . however , like the living peppered moray eel (
figure 2 . greererpeton reduced to a blueprint of body parts . here there may be one extra phalanx on pedal digit 5 and one missing on pedal digit 2 compared to sister taxa . so an alternate is shown with that repair . the skulls at left are juveniles .
were likewise flat - and long - bodied aquatic forms that seem unlikely to have been able to support themselves without the natural buoyancy of water . their descendants in the lrt likewise look like they were more comfortable lounging underwater like living
the hellbender has working lungs , but gill slits are often retained , although only immature specimens have true gills ; the hellbender absorbs oxygen from the water through capillaries of its side frills . ”
only rarely do hellbenders leave the water , perhaps to climb on low pond rocks . if the
figure 3 . pederpes is a basal taxon in the whatcheeria + crassigyrinus clade .
( with its flattened skull ) appear to have opted for a return to a watery environment . and who could blame them ? in any case , their big lumbering bodies were not well adapted to clambering over dry obstacles , like rocks and plants , that made terrestrial locomotion more difficult . and the biggest best food was still in the water . no doubt limbs helped many of them find new ponds and swamps when they felt the urge to do so , like living crocs . and they probably left the water after some of the smaller and more able taxa listed below .
took the first steps toward more of a land - living life . the nostrils shifted forward , but were still tiny , at first . bur the ribs were slender without any overlap . perhaps this signaled improvements in lung power . larger nostrils appeared in
. all these taxa were still rather large and lumbering and so were probably more at home in the water .
figure 5 . eucritta in situ and reconstructed . note the large pes in green .
like ancestors . one descendant clade begins with a several long - bodied , short - legged salamander - like taxa . discosauriscus is one of these . it begins life in water , but grows up to prefer dry land . seymouria is the culmination of this clade .
, a short - legged , flat - bodied aquatic taxon . that plesiomorphic taxon gives rise to legless
their marriage to or divorce from water varies across a wide spectrum in living taxa .
figure 7 . various stem amniotes ( reptiles ) that precede tulerpeton in the lrt . so these taxa likely radiated in the late devonian . and taxa like acanthostega and ichthyostega are late - survivors of earlier radiations documented by the earlier trackways .
no one should have ever said you have to look like a typical reptile to lay an amnion - covered egg . and if they did , they were not guided by a large gamut phylogenetic analysis .
this clade become fully divorced from needing water for reproduction , but basal members still liked the high humidity and wet substrate of the swamp .
. these were small agile taxa with relatively long legs that would have had their genesis in the late devonian . their first appearance in the fossil record was much later . the development of the amnion - enclosed embryo may have taken millions of years . the first phylogenetic reptiles appear in the form of amphibian - like
, according to the lrt , from distinct clades that were more or less ready to do so and in different ways . and , of course , odd extant fish , like the peppered moray eel (
) continue this tradition . what will they eventually evolve into , given enough time ?
the fin to limb transition : new data , interpretations , and hypotheses from paleontology and developmental biology . annual review of earth and planetary sciences . 37 : 163–179 .
jarvik : postcranial anatomy , basal tetrapod interrelationships and patterns of skeletal evolution . earth and environmental science transactions of the royal society of edinburgh .
polydactly in the earliest known tetrapod limbs . nature 347 : 66 - 69 .
diurnal , land - based predation on shore crabs by moray eels in the chagos archipelago . coral reefs 28 ( 2 ) : 387–397 .
on the fish - like tail in the ichtyhyostegid stegocephalians . meddelelser om grønland 114 : 1–90 .
figure 1 . the complete skull of anthracosaurus greatly resembles its relative , neopteroplax .
anthracosaurus russelli ( huxley 1863 , panchen 1977 , clack 1987 ; westphalian , late carboniferous , 310 mya , skull length 40cm ; figs . 1 , 2 ) was originally considered a labyrinthodont . the wide , yet pointed , triangular skull and tall orbits recall traits found in labyrinthodonts , like sclerocephalus , and in the basal tetrapod , tiktaalik . here , in the large reptile tree ( lrt , 967 taxa ) , anthracosaurus nests with neopteroplax ( fig . 3 ) as a derived embolomere , the clade that likely gave rise to seymouriamorpha , lepospondyli and reptilia
has an inverted teardrop shape . the marginal and palatal fangs are quite large . although flattened in dorsal view , comparisons suggest the jaw margin was convex , as in
is basal in the embolomeri . those giant marginal and palatal fangs indicate a predatory niche .
figure 2 . left : anthracosaurus chimaera from clack 1987 . right : older tracing in dorsal view of the complete skull and palatal view attributed to anthracosaurus from an online photo . the narrower skull is made of several different specimens ( chimaera ) and produces a loss of resolution in the lrt .
( fig . 2 ) based on a chimaera of specimens . unfortunately , plugging that data into the lrt produced loss of resolution over several nodes . using the older single skull in dorsal view had no such problems .
figure 3 . neopteroplax has a skull quite similar to the older single skull of anthracosaurus and they nest together in the lrt .
“gauthier , kluge and rowe ( 1988 ) defined anthracosauria as ‘amniota plus all other tetrapods that are more closely related to amniotes than they are to amphibians” ( amphibia in turn was defined by these authors as a clade including lissamphibia and those tetrapods that are more closely related to lissamphibians than they are to amniotes ) . ”
in this definition non - amniote anthracosauria does not include anthracosaurus , but only silvanerpeton and gephyrostegus in the lrt because more basal taxa are also basal to amphibians .
“as ruta , coates and quicke ( 2003 ) pointed out , this definition is problematic , because , depending on the exact phylogenetic position of lissamphibia within tetrapoda , using it might lead to the situation where some taxa traditionally classified as anthracosaurs , including even the genus anthracosaurus itself , wouldn’t belong to anthracosauria .
laurin ( 2001 ) created a different phylogenetic definition of anthracosauria , defining it as “the largest clade that includes anthracosaurus russelli but not ascaphus truei” .
in the lrt ascaphus , the tailed frog , is derived from the large clade , the embolomeri , that includes anthracosaurus . however the small clade that includes just anthracosaurus and neopteroplax does not include the tailed frog .
in the lrt the embolomeri are basal to eucritta and the seymouriamorpha , which are basal to the reptilia ( = amniota ) and lepospondyli ( including amphibia ) . the chroniosuchia and gephyrostegus are both amphibian - like reptiles in the lrt .
s the exact phylogenetic position of lissamphibia within tetrapoda remains uncertain , it also remains controversial which fossil tetrapods are more closely related to amniotes than to lissamphibians , and thus , which ones of them were reptiliomorphs in any meaning of the word . ”
( amphibia : anthracosauria ) from the northumberland coal measures . palaeontology 30 ( 1 ) : 15 - 26 .
description of anthracosaurus russelli , a new labyrinthodont from the lanarkshire coalfield . quartery journal of the geological society 19 : 56 - 58 .
huxley . philosophical transactions of the royal society of london , b . 269 : 581 - 640 .
huxley ( amphibia : labyrinthodontia ) and the family anthracosauridae . philosophical transactions of the royal society b . 279 ( 968 ) : 447–512 .