marginella liparozona is a species of colorful small sea snail , a marine gastropod mollusk in the marginellidae family .

the species is endemic to são tomé and príncipe .

marginella liparozona

marginella liparozona tomlin & shackleford , 1913 . retrieved through : world register of marine species on 31 may 2010 .
marginella jousseaumei locard , 1897 : synonym of marginella subturrita p . fischer , 1884
marginella jousseaumi locard , 1897 : synonym of marginella subturrita p . fischer , 1884
marginella gorii t . cossignani , 2012 : synonym of marginella gemma a . adams , 1850
marginella unifasciata turton , 1932 : synonym of marginella munda e . a . smith , 1904
marginella tomlini shackleford , 1916 : synonym of marginella bicatenata g . b . sowerby iii , 1914
marginella nimbosus s . g . veldsman , 2013 : synonym of marginella nimbosa s . g . veldsman , 2013
marginella intermedia g . b . sowerby ii , 1846 : synonym of marginella floccata g . b . sowerby iii , 1889
marginella savignyi . retrieved through : world register of marine species on 31 may 2010 .
marginella suezensis . retrieved through : world register of marine species on 31 may 2010 .
marginella arcana s . g . veldsman , aiken & j . h . veldsman , 2014
marginella mzimayiensis s . g . veldsman , aiken & j . h . veldsman , 2015
marginella obliqua s . g . veldsman , aiken & j . h . veldsman , 2014
marginella umzumbeensis s . g . veldsman , aiken & j . h . veldsman , 2016
marginella amydrozona melvill . retrieved through : world register of marine species on 31 may 2010 .
marginella persicula linnaeus . retrieved through : world register of marine species on 31 may 2010 .
marginella pygmaea issel , 1869 : synonym of marginella isseli g . nevill & h . nevill , 1875 : synonym of granulina isseli ( g . nevill & h . nevill , 1875 )
marginella lamarck , 1799 . 6 december 2010 . retrieved through : world register of marine species .
marginella epipolia tomlin 1921 . retrieved through : world register of marine species on 31 may 2010 .
marginella pachista tomlin 1913 . retrieved through : world register of marine species on 31 may 2010 .
marginella barnardi tomlin , 1919 : synonym of gibberula differens ( e . a . smith , 1904 )
marginella borealis verrill , 1884 : synonym of prunum boreale ( a . e . verrill , 1884 )
marginella deburghi a . adams , 1864 : synonym of persicula deburghi ( a . adams , 1864 )
marginella evelynae bayer , 1943 : synonym of prunum evelynae ( f . m . bayer , 1943 )
marginella ithychila tomlin , 1918 : synonym of gibberula dulcis ( e . a . smith , 1904 )
marginella lactea kiener , 1841 : synonym of volvarina abbreviata ( c . b . adams , 1850 )
marginella mirabilis h . adams , 1869 : synonym of glabella mirabilis ( h . adams , 1869 )
marginella nobiliana bayer , 1943 : synonym of prunum nobilianum ( f . m . bayer , 1943 )
marginella taylori shackleford , 1916 : synonym of gibberula differens ( e . a . smith , 1904 )
marginella virginiana verrill , 1885 : synonym of dentimargo smithii ( a . e . verrill , 1885 )
marginella adamkusi bozzetti , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella amazona bavay , 1912 . retrieved through : world register of marine species on 31 may 2010 .
marginella anapaulae massier , 2004 . retrieved through : world register of marine species on 31 may 2010 .
marginella anna jousseaume , 1881 . retrieved through : world register of marine species on 31 may 2010 .
marginella aurantia lamarck , 1822 . retrieved through : world register of marine species on 31 may 2010 .
marginella bavayi dautzenberg , 1910 . retrieved through : world register of marine species on 31 may 2010 .
marginella belcheri hinds , 1844 . retrieved through : world register of marine species on 31 may 2010 .
marginella broderickae hayes , 2001 . retrieved through : world register of marine species on 31 may 2010 .
marginella caterinae bozzetti , 1991 . retrieved through : world register of marine species on 31 may 2010 .
marginella cosmia bartsch , 1915 . retrieved through : world register of marine species on 31 may 2010 .
marginella croukampi hayes , 1996 . retrieved through : world register of marine species on 31 may 2010 .
marginella emmae bozzetti , 1998 . retrieved through : world register of marine species on 31 may 2010 .
marginella eucosmia bartsch , 1915 . retrieved through : world register of marine species on 31 may 2010 .
marginella felixi massier , 2004 . retrieved through : world register of marine species on 31 may 2010 .
marginella festiva kiener , 1841 . retrieved through : world register of marine species on 31 may 2010 .
marginella fishhoenkensis massier , 2004 . retrieved through : world register of marine species on 31 may 2010 .
marginella gabrielae bozzetti , 1998 . retrieved through : world register of marine species on 31 may 2010 .
marginella geraldi lussi , 2006 . retrieved through : world register of marine species on 31 may 2010 .
marginella gloriosa jousseaume , 1884 . retrieved through : world register of marine species on 31 may 2010 .
marginella hayesi bozzetti , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella helmatina rang , 1832 . retrieved through : world register of marine species on 31 may 2010 .
marginella huberti clover , 1972 . retrieved through : world register of marine species on 31 may 2010 .
marginella hybrida cossignani , 2006 . retrieved through : world register of marine species on 31 may 2010 .
marginella irrorata menke , 1828 . retrieved through : world register of marine species on 31 may 2010 .
marginella joanae bozzetti , 2001 . retrieved through : world register of marine species on 31 may 2010 .
marginella joanmassierae bozzetti , 1992 . retrieved through : world register of marine species on 31 may 2010 .
marginella lamarcki boyer , 2004 . retrieved through : world register of marine species on 31 may 2010 .
marginella limbata lamarck , 1822 . retrieved through : world register of marine species on 31 may 2010 .
marginella lineatolabrum gaskoin , 1840 . retrieved through : world register of marine species on 31 may 2010 .
marginella lineofasciata bozzetti , 1992 . retrieved through : world register of marine species on 31 may 2010 .
marginella marocana locard , 1897 . retrieved through : world register of marine species on 31 may 2010 .
marginella millardi lussi , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella monicae bozzetti , 1997 . retrieved through : world register of marine species on 31 may 2010 .
marginella musica hinds , 1844 . retrieved through : world register of marine species on 31 may 2010 .
marginella natalcinerea massier , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella nevillana kilburn , 1977 . retrieved through : world register of marine species on 31 may 2010 .
marginella ornata redfield , 1870 . retrieved through : world register of marine species on 31 may 2010 .
marginella orstomi coomans , 1975 . retrieved through : world register of marine species on 31 may 2010 .
marginella peelae bozzetti , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella petitii duval , 1841 . retrieved through : world register of marine species on 31 may 2010 .
marginella piperata hinds , 1844 . retrieved through : world register of marine species on 31 may 2010 .
marginella pseudornata bozzetti , 1992 . retrieved through : world register of marine species on 31 may 2010 .
marginella pseudosebastiani mattavelli , 2001 . retrieved through : world register of marine species on 31 may 2010 .
marginella purpurea cossignani , 2006 . retrieved through : world register of marine species on 31 may 2010 .
marginella rosea lamarck , 1822 . retrieved through : world register of marine species on 31 may 2010 .
marginella roseafasciata massier , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella roseolineata turton , 1932 . retrieved through : world register of marine species on 31 may 2010 .
marginella senegalensis clover , 1990 . retrieved through : world register of marine species on 31 may 2010 .
marginella sergioi bozzetti , 1997 . retrieved through : world register of marine species on 31 may 2010 .
marginella stuarti kilburn , 1977 . retrieved through : world register of marine species on 31 may 2010 .
marginella tuguriana lussi , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella verrilli morrison , 1967 . retrieved through : world register of marine species on 31 may 2010 .
marginella vexillum redfield , 1852 . retrieved through : world register of marine species on 31 may 2010 .
marginella werneri bozzetti , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella westhuizeni massier , 1993 . retrieved through : world register of marine species on 31 may 2010 .
marginella decaryi bavay , 1920 . retrieved through : world register of marine species on 31 may 2010 .
marginella delphinica bavay , 1920 . retrieved through : world register of marine species on 31 may 2010 .
marginella virgula jousseaume , 1922 . retrieved through : world register of marine species on 31 may 2010 .
marginella granum philippi , 1849 . retrieved through : world register of marine species on 31 may 2010 .
marginella angustata sowerby , 1846 . retrieved through : world register of marine species on 31 may 2010 .
marginella burnupi sowerby , 1897 . retrieved through : world register of marine species on 31 may 2010 .
marginella cherubini bavay , 1922 . retrieved through : world register of marine species on 31 may 2010 .
marginella henrikasi bozzetti , 1995 . retrieved through : world register of marine species on 31 may 2010 .
marginella lantzi jousseaume , 1875 . retrieved through : world register of marine species on 31 may 2010 .
marginella marianae bozzetti , 1999 . retrieved through : world register of marine species on 31 may 2010 .
marginella pulchella kiener , 1834 . retrieved through : world register of marine species on 31 may 2010 .
marginella pumila redfield , 1870 . retrieved through : world register of marine species on 31 may 2010 .
marginella sandwicensis pease , 1860 . retrieved through : world register of marine species on 31 may 2010 .
marginella alabaster reeve , 1865 : synonym of volvarina fauna ( g . b . sowerby i , 1846 )
marginella albina gaskoin , 1853 : synonym of mesoginella turbinata ( g . b . sowerby ii , 1846 )
marginella attenuata weinkauff , 1879 : synonym of austroginella translucida ( g . b . sowerby ii , 1846 )
marginella candida bivona ant . , 1832 : synonym of ringicula auriculata ( ménard de la groye , 1811 )
marginella ignota jousseaume , 1875 : synonym of dentimargo neglecta ( g . b . sowerby ii , 1846 )
marginella imperatrix sykes , 1905 : synonym of glabella pseudofaba ( g . b . sowerby ii , 1846 )
marginella inflexa sowerby g . b . i , 1846 : synonym of volvarina mitrella ( risso , 1826 )
marginella leia cotton , 1944 : synonym of mesoginella turbinata ( g . b . sowerby ii , 1846 )
marginella lepta bartsch , 1915 : synonym of gibberula burnupi ( g . b . sowerby iii , 1897 )
marginella marianae bozzetti , 1999 : [ 123 ] synonym of prunum pyrumoides lussi & g . smith , 1999
marginella multizonata krauss , 1848 : synonym of hyalina cylindrica ( g . b . sowerby ii , 1846 )
marginella pattisoni cotton , 1944 : synonym of mesoginella turbinata ( g . b . sowerby ii , 1846 )
marginella pellucida weinkauff , 1879 : synonym of volvarina fauna ( g . b . sowerby i , 1846 )
marginella petterdi beddome , 1883 : synonym of ovaginella ovulum ( g . b . sowerby ii , 1846 )
marginella reevei krauss , 1852 : synonym of dentimargo neglecta ( g . b . sowerby ii , 1846 )
marginella rufula gaskoin , 1853 : synonym of dentimargo neglecta ( g . b . sowerby ii , 1846 )
marginella volutiformis reeve , 1865 : synonym of austroginella translucida ( g . b . sowerby ii , 1846 )
marginella claudoni bavay , date unknown . retrieved through : world register of marine species on 31 may 2010 .
marginella scalariformis duclos , date unknown . retrieved through : world register of marine species on 31 may 2010 .
marginella bellii sowerby ii , 1846 : synonym of glabella bellii ( g . b . sowerby ii , 1846 )
marginella harpaeformis sowerby ii , 1846 : synonym of glabella harpaeformis ( g . b . sowerby ii , 1846 )
marginella pseudofaba sowerby ii , 1846 : synonym of glabella pseudofaba ( g . b . sowerby ii , 1846 )
marginella aequinoctialis boyer & simbille , 2004 . retrieved through : world register of marine species on 31 may 2010 .
marginella aronnax bouchet & warén , 1985 . retrieved through : world register of marine species on 31 may 2010 .
marginella carquejai gofas & fernandes , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella celestae massier & rosado , 2008 . retrieved through : world register of marine species on 31 may 2010 .
marginella chalmersi tomlin & shackleford , 1912 . retrieved through : world register of marine species on 31 may 2010 .
marginella cloveri rios & matthews , 1972 . retrieved through : world register of marine species on 31 may 2010 .
marginella colomborum ( bozzetti , 1995 ) . retrieved through : world register of marine species on 31 may 2010 .
marginella desjardini marche - marchad , 1957 . retrieved through : world register of marine species on 31 may 2010 .
marginella gemma a . adams , 1850 . retrieved through : world register of marine species on 31 may 2010 .
marginella gemmula bavay in dautzenberg , 1913 . retrieved through : world register of marine species on 31 may 2010 .
marginella genessi h . fisher , 1901 . retrieved through : world register of marine species on 31 may 2010 .
marginella gilva goud & neefs , 1996 . retrieved through : world register of marine species on 31 may 2010 .
marginella glabella ( linnaeus , 1758 ) . retrieved through : world register of marine species on 31 may 2010 .
marginella himburgae massier & zettler , 2009 . retrieved through : world register of marine species on 31 may 2010 .
marginella immelmani hart m . , 2001 . retrieved through : world register of marine species on 31 may 2010 .
marginella impudica p . fischer , 1883 . retrieved through : world register of marine species on 31 may 2010 .
marginella joostei liltved & millard , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella lemaitrei liltved & millard , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella luculenta gofas & fernandes , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella lussii hayes & millard , 1995 . retrieved through : world register of marine species on 31 may 2010 .
marginella marimba gofas & fernandes , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella mauretanica boyer & neefs , 1999 . retrieved through : world register of marine species on 31 may 2010 .
marginella melvilli tomlin & shackleford , 1913 . retrieved through : world register of marine species on 31 may 2010 .
marginella minuscula ( turton , 1932 ) . retrieved through : world register of marine species on 31 may 2010 .
marginella poppei boyer & neefs , 1999 . retrieved through : world register of marine species on 31 may 2010 .
marginella simulata gofas & fernandes , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella spinacia gofas & fernandes , 1988 . retrieved through : world register of marine species on 31 may 2010 .
marginella spiralineata b . hayes , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella subturrita p . fischer , 1883 . retrieved through : world register of marine species on 31 may 2010 .
marginella susanae veldsman & jooste , 2009 . retrieved through : world register of marine species on 31 may 2010 .
marginella undulans gofas & fernandes , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella verdascai hayes & rosado , 2007 . retrieved through : world register of marine species on 31 may 2010 .
marginella xhosa liltved & millard , 1994 . retrieved through : world register of marine species on 31 may 2010 .
marginella eveleighi tom & schackelford , 1913 . retrieved through : world register of marine species on 31 may 2010 .
marginella extra ( jousseaume , 1894 ) . retrieved through : world register of marine species on 31 may 2010 .
marginella lateritia melvill & sykes , 1903 . retrieved through : world register of marine species on 31 may 2010 .
marginella reeveana ( petit , 1851 ) . retrieved through : world register of marine species on 31 may 2010 .
marginella signali dautzenberg & fischer , 1896 . retrieved through : world register of marine species on 31 may 2010 .
marginella angustata sowerby , 1846 : [ 115 ] synonym of volvarina angustata ( g . b . sowerby , 1846 )
marginella cypraeoides tenison - woods , 1878 : synonym of ovaginella ovulum ( g . b . sowerby ii , 1846 )
marginella cleryi petit de la saussaye , 1836 . retrieved through : world register of marine species on 31 may 2010 .
marginella matthewsi van mol & tursch , 1967 . retrieved through : world register of marine species on 31 may 2010 .
marginella abbreviata c . b . adams , 1850 : synonym of volvarina abbreviata ( c . b . adams , 1850 )
marginella algoensis e . a . smith , 1901 : synonym of crithe algoensis ( e . a . smith , 1901 )
marginella burnupi sowerby , 1897 : [ 116 ] synonym of gibberula burnupi ( g . b . sowerby iii , 1897 )
marginella carinata e . a . smith , 1891 : synonym of alaginella carinata ( e . a . smith , 1891 )
marginella differens e . a . smith , 1904 : synonym of gibberula differens ( e . a . smith , 1904 )
marginella fallax e . a . smith , 1903 : synonym of canalispira fallax ( e . a . smith , 1903 )
marginella fluctuata c . b . adams , 1850 : synonym of gibberula fluctuata ( c . b . adams , 1850 )
marginella guillaini petit de la saussaye , 1851 : synonym of glabella obtusa ( g . b . sowerby ii , 1846 )
marginella ingloria e . a . smith , 1910 : synonym of volvarina ingloria ( e . a . smith , 1910 )
marginella onychina a . adams & reeve , 1850 : synonym of cryptospira onychina ( a . adams & reeve , 1850 )
marginella pygmaea g . b . sowerby , 1846 : synonym of mesoginella pygmaea ( g . b . sowerby , 1846 )
marginella rubella c . b . adams , 1845 : synonym of volvarina rubella ( c . b . adams , 1845 )
marginella shepstonensis e . a . smith , 1906 : synonym of persicula shepstonensis ( e . a . smith , 1906 )
marginella walkeri e . a . smith , 1899 : synonym of dentimargo walkeri ( e . a . smith , 1899 )
marginella diadochus a . adams & reeve , 1848 . retrieved through : world register of marine species on 31 may 2010 .
marginella goodalli g . b . sowerby , 1825 . retrieved through : world register of marine species on 31 may 2010 .
marginella mosaica g . b . sowerby , 1846 . retrieved through : world register of marine species on 31 may 2010 .
marginella munda e . a . smith , 1904 . retrieved through : world register of marine species on 31 may 2010 .
marginella nebulosa ( bolten in röding , 1798 ) . retrieved through : world register of marine species on 31 may 2010 .
marginella punctilineata e . a . smith , 1899 . retrieved through : world register of marine species on 31 may 2010 .
marginella sebastiani marche - marchad & rosso , 1979 . retrieved through : world register of marine species on 31 may 2010 .
marginella inconspicua g . b . sowerby ii , 1846 : synonym of mesoginella inconspicua ( g . b . sowerby , 1846 )
marginella vignali dautzenberg & fischer , 1896 : [ 132 ] synonym of gibberula vignali ( dautzenberg & h . fischer , 1896 )
marginella bairstowi g . b . sowerby iii , 1886 . retrieved through : world register of marine species on 31 may 2010 .
marginella bicatenata g . b . sowerby iii , 1914 . retrieved through : world register of marine species on 31 may 2010 .
marginella floccata g . b . sowerby iii , 1889 . retrieved through : world register of marine species on 31 may 2010 .
marginella lineolata g . b . sowerby iii , 1886 . retrieved through : world register of marine species on 31 may 2010 .
marginella lutea g . b . sowerby iii , 1889 . retrieved through : world register of marine species on 31 may 2010 .
marginella curta g . b . sowerby i , 1832 : synonym of prunum curtum ( g . b . sowerby i , 1832 )
marginella cylindrica g . b . sowerby ii , 1846 : synonym of hyalina cylindrica ( g . b . sowerby ii , 1846 )
marginella electrina g . b . sowerby iii , 1892 : synonym of hyalina electrina ( g . b . sowerby iii , 1892 )
marginella evanida g . b . sowerby ii , 1846 : synonym of volvarina evanida ( g . b . sowerby ii , 1846 )
marginella fauna g . b . sowerby i , 1846 : synonym of volvarina fauna ( g . b . sowerby i , 1846 )
marginella neglecta g . b . sowerby ii , 1846 : synonym of dentimargo neglecta ( g . b . sowerby ii , 1846 )
marginella obtusa g . b . sowerby ii , 1846 : synonym of glabella obtusa ( g . b . sowerby ii , 1846 )
marginella ovulum g . b . sowerby ii , 1846 : synonym of ovaginella ovulum ( g . b . sowerby ii , 1846 )
marginella perminima g . b . sowerby iii , 1894 : synonym of granulina perminima ( g . b . sowerby iii , 1894 )
marginella ponsonbyi g . b . sowerby iii , 1897 : synonym of hyalina cylindrica ( g . b . sowerby ii , 1846 )
marginella princeps g . b . sowerby iii , 1901 : synonym of closia princeps ( g . b . sowerby iii , 1901 )
marginella ringicula g . b . sowerby iii , 1901 : synonym of dentimargo rincigula ( g . b . sowerby iii , 1901 )
marginella robusta g . b . sowerby iii , 1904 : synonym of persicula robusta ( g . b . sowerby iii , 1904 )
marginella sauliae g . b . sowerby ii , 1846 : synonym of volvarina sauliae ( g . b . sowerby ii , 1846 )
marginella taeniata g . b . sowerby ii , 1846 : synonym of volvarina taeniata ( g . b . sowerby ii , 1846 )
marginella translucida g . b . sowerby ii , 1846 : synonym of austroginella translucida ( g . b . sowerby ii , 1846 )
marginella turbinata g . b . sowerby ii , 1846 : synonym of mesoginella turbinata ( g . b . sowerby ii , 1846 )
marginella picturata g . nevill & h . nevill , 1874 . retrieved through : world register of marine species on 31 may 2010 .
marginella isseli g . nevill & h . nevill , 1875 : synonym of granulina isseli ( g . nevill & h . nevill , 1875 )
marginella bella auct . : synonym of volvarina lactea ( kiener , 1841 ) : synonym of volvarina abbreviata ( c . b . adams , 1850 )
marginella jousseaumi rochebrune , 1881 : synonym of prunum sauliae ( g . b . sowerby ii , 1846 ) : synonym of volvarina sauliae ( g . b . sowerby ii , 1846 )
marginella is a genus of small tropical and temperate sea snails , marine gastropod molluscs in the family marginellidae , the margin snails . it is the type genus of the family . [ 1 ]
marginella pseustes e . a . smith , 1904 : synonym of granulina pseustes ( e . a . smith , 1904 ) : synonym of cystiscus pseustes ( e . a . smith , 1904 )
malacolog ( four marginella species names are listed for the western atlantic ocean , one name is not available ) . malacolog is created by dr . gary rosenberg , the academy of natural sciences , philadelphia
the higher classification of the family marginellidae has long been in a state of confusion . many popular works still treat all members of this family under the single genus marginella , basing them primarily on superficial similarities of the shell .
tomlin j . r . le b . & shackleford l . j . 1913 . description of new species of marginella and mucronalia from são thomé . journal of conchology , 14 : 43 , pl . 1 . , available online at urltoken page ( s ) : 43 [ details ]
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cossignani t . ( 2006 ) . marginellidae & cystiscidae of the world . l ' informatore piceno . 408pp . [ details ]
gofas s . & fernandes f . 1988 . the marginellids of são tomé , west africa . journal of conchology 33 ( 1 ) : 1 - 30 , pls . 1 - 2 . page ( s ) : 8 - 9 [ details ]
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label : s . thomé . tomlin , 1913 : island of sao thomé .
t & s ( pl . i . , f . 5 , 6 ) , and
t . & s . ( pl . i . , 1 , 2 ) . ” two paratypes - nmw 1955 . 158 . 01177
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the shells of species in this genus are rounded , smooth and glossy , with a large aperture that appears to be toothed because it shows the edge of the columellar folds . in many species the shells are colorful . the glossy surface of the shell results from the fact that the mantle covers most of the shell when the animal is active . as is typical in the neogastropoda , the animal has a long siphon . when the animal is active , the foot extends much further out than the edge of the shell .
as is also typical for the neogastropoda , species in this genus are carnivorous and predatory .
the shells of the species in this genus have spires which range from moderately elevated to flattened . the surface of the shell is glossy and porcellaneous , and it is often but not always colourful . the columella has four definite , subequal plaits on its anterior half . the outer lip is thickened , and generally denticulate inside , with distinct teeth or folds . [ 2 ] the siphonal canal is not deeply incised .
in the living animal , the mantle only partly extends over the shell when the animal is moving .
the head is bifurcated , with slender tentacles and eyes in small bulges lateral to the base of tentacles . the siphon is large and protrudes over the head . the foot is large and flat , and when it is extended is slightly longer than the shell .
bartsch , p . 1915 . report on the turton collection of south african marine mollusks , with additional notes on other south african shells contained in the united states national museum . bulletin of the united states national museum , issued july 28 , 1915 .
cossignani t . ( 2006 ) . marginellidae & cystiscidae of the world . l ' informatore piceno . 408pp
this page was last edited on 27 march 2018 , at 16 : 59 .
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marginellidae , or the margin shells , are a taxonomic family of small , often colorful , sea snails , marine gastropod molluscs in the clade neogastropoda .
the confusion over the classification stems from the fact that the earlier classifications were based rather crudely on shell characters . although many good differential shell characters do exist within this group , those characters were generally misinterpreted or not recognized as significant . such information as did exist on the radulae and the external anatomy of the living animals was widely scattered in the scientific literature , and internal anatomical descriptions were not available until fairly recently .
the shell of marginellidae is usually small , but varies in different species from minute to medium - sized . the external color of the shell can be white , cream , yellow , orange , red , or brown , and can be uniformly colored , or patterned in various ways . the protoconch is paucispiral . the lip of the shell is thickened , and can be smooth or denticulate . an external varix may be present or absent , a siphonal notch may be present or absent . the columella may have 2 - 6 plications . the operculum is absent in this family .
subfamily granulininae : this subfamily was originally placed in family cystiscidae by coovert & coovert ( 1995 ) but placed back in marginellidae , following la perna ( 1999 ) and based on the morphology of living animals .


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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urltoken
$ 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
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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 .
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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 .
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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 .