Temnospondyli

Reconstruction of Dendrerpeton, from here.


Belongs within: Neostegalia.
Contains: Stereospondylomorpha, Dissorophoidea.

The Temnospondyli were one of the most significant early clades of amphibian-grade tetrapods, being recorded from the early Carboniferous (Visean) to the mid-Cretaceous (Albian). Whether they survived after this date depends on whether the modern amphibians (particularly frogs and salamanders) are accepted as derived temnospondyls. The clade as a whole is characterised by the presence of interpterygoid vacuities at least half as wide as the skull and bordered by triradiate pterygoids (Ruta et al. 2003). The majority of temnospondyls were semiaquatic, though the clade as a whole includes members varying from almost entirely terrestrial to fully aquatic.

Early Carboniferous temnospondyls include the Scottish Balanerpeton and the Nova Scotian Dendrerpeton, both medium-sized semi-terrestrial forms with the latter reaching about a metre in length. The Late Carboniferous and Early Permian Cochleosauridae were crocodile-like semi-aquatic ambush predators, reaching lengths of over 1.5 m. The Early Permian saw the appearance of even larger forms: Edops craigi reached about two metres in length, whereas the three metre-long Eryops megacephalus was one of the largest land animals of its time. Eryops had a more strongly build appendicular skeleton than other temnospondyls, indicating a greater degree of adaptation for terrestrial locomotion.

Temnospondyli [Euskelia]
    |--+--+--Edops [Edopidae, Edopoidea] PS17
    |  |  |    `--E. craigi Romer 1935 RCQ03
    |  |  `--Cochleosauridae RCQ03
    |  |       |--Chenoprosopus lewisi Hook 1993 RCQ03
    |  |       `--Cochleosaurus Fritsch 1885 D07
    |  |            |--C. bohemicus D07
    |  |            `--C. florensis Rieppel 1980 RCQ03
    |  `--Eryopoidea YW00
    |       |--Zatrachydidae YW00
    |       `--Eryopidae RCQ03
    |            |--Onchiodon YW00
    |            `--Eryops Miner 1925 PS17, D07
    |                 `--E. megacephalus Cope 1877 RCQ03
    `--+--+--Balanerpeton woodi Milner & Sequeira 1994 PS17, RCQ03
       |  `--Dendrerpeton [Dendrerpetontidae] PS17
       |       |--D. acadianum Owen 1853 RCQ03
       |       `--D. helogenes YW00
       `--Dissorophoidea PS17

Temnospondyli incertae sedis:
  Adamanterpeton ohioensis RCQ03
  Cacops Williston 1910 D07
    `--C. aspideporus D07
  Kamacops S02
  Cyclotosaurus PS88 [incl. Procyclotosaurus W70]
  Laccosaurus PS88
  Platyhystrix Berman, Reisz & Fracasso 1981 D07
    `--P. rugosus D07
  Gerrothorax Nilsson 1937 D07
    |--G. pulcherrimus D07
    `--G. rhaeticus D07
  Latiscopus [Latiscopidae] CZ79
    `--L. disjunctus Wilson 1948 CZ79
  Limnarchia [Dvinosauroidea] RCQ03
    |  i. s.: Erpetosaurus PS88
    |--Stereospondylomorpha YW00
    `--Dvinosauria YW00
         |  i. s.: Eugyrinus YW00
         |--Trimerorhachidae [Trimerorhachoidea] RCQ03
         |    |--Neldasaurus wrightae Chase 1965 YW00, RCQ03
         |    `--Trimerorhachis insignis Case 1935 YW00, RCQ03
         `--Dvinosauroidea YW00
              |--Eobrachyopidae YW00
              |    |--Isodectes obtusus (Cope 1868) YW00, RCQ03 (see below for synonymy)
              |    `--Acroplous vorax YW00
              `--+--Tupilakosaurus [Tupilakosauridae] YW00
                 `--Dvinosaurus [Dvinosauridae] YW00
                      |--*D. primus Amalitskii 1921 YW00, TT05
                      `--D. egregius YW00

Isodectes obtusus (Cope 1868) YW00, RCQ03 [=Saurerpeton obtusum YW00; incl. I. megalops YW00]

*Type species of generic name indicated

REFERENCES

[CZ79] Cosgriff, J. W., & J. M. Zawiskie. 1979. A new species of the Rhytidosteidae from the Lystrosaurus Zone and a review of the Rhytidosteoidea. Palaeontologia Africana 22: 1–27.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[PS88] Panchen, A. L., & T. R. Smithson. 1988. The relationships of the earliest tetrapods. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 1–32. Clarendon Press: Oxford.

[PS17] Pardo, J. D., M. Szostakiwskyj, P. E. Ahlberg & J. S. Anderson. 2017. Hidden morphological diversity among early tetrapods. Nature 546: 642–645.

[RCQ03] Ruta, M., M. I. Coates & D. L. J. Quicke. 2003. Early tetrapod relationships revisited. Biological Reviews 78: 251–345.

[S02] Schoch, R. R. 2002. The neurocranium of the stereospondyl Mastodonsaurus giganteus. Palaeontology 45 (3): 627–645.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27–77.

[W70] Wills, L. J. 1970. The Triassic succession in the central Midlands in its regional setting. Quarterly Journal of the Geological Society of London 126: 225–283.

[YW00] Yates, A. M., & A. A. Warren. 2000. The phylogeny of the ‘higher’ temnospondyls (Vertebrata: Choanata) and its implications for the monophyly and origins of the Stereospondyli. Zoological Journal of the Linnean Society 128: 77–121.

Last updated: 3 August 2017.

Chelidae

Murray turtles Emydura macquarii, photographed by David Kleinert.


Belongs within: Testudinata.

The Chelidae are a family of side-necked freshwater turtles found in South America and Australia.

Synapomorphies (from Gaffney & Meylan 1988): Lateral cheek emargination unusually well developed, reaching the parietal; quadratojugal absent; mesoplastra absent.

<==Chelidae
    |--Pseudemydura umbrina GM88, USDI77
    `--Chelinae GM88
         |--+--Emydura macquarrii GM88
         |  |--Elseya GM88
         |  `--Rheodytes GM88
         `--Chelodd GM88
              |--Platemys platycephala GM88, DS86
              |    |--P. p. platycephala DS86
              |    `--P. p. melanonota DS86
              `--Chelini GM88
                   |--+--Batrachemys GM88
                   |  |--Mesoclemmys GM88
                   |  `--Phrynops GM88
                   |       |--P. gibbus DS86
                   |       `--P. nasutus DS86
                   `--Chelina GM88
                        |--Chelus fimbriatus GM88, DS86
                        `--Hydromedusad GM88
                             |--Hydromedusa GM88
                             `--Chelodina GM88
                                  |--C. oblonga GM88
                                  `--C. steindachneri HD04

*Type species of generic name indicated

REFERENCES

[DS86] Dixon, J. R. & P. Soini. 1986. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Milwaukee Public Museum: Milwaukee.

[GM88] Gaffney, E. S., & P. A. Meylan. 1988. A phylogeny of turtles. In The Phylogeny and Classification of the Tetrapods, vol. 1. Amphibians, Reptiles, Birds (M. J. Benton, ed.) pp. 157-219. Clarendon Press: Oxford.

[HD04] How, R. A., & J. Dell. 2004. Reptile assemblage of the Abydos Plain, north-eastern Pilbara, Western Australia. Journal of the Royal Society of Western Australia 87 (3): 85-95.

[OC03] Olson, P. D., T. H. Cribb, V. V. Tkach, R. A. Bray & D. T. J. Littlewood. 2003. Phylogeny and classification of the Digenea (Platyhelminthes: Trematoda). International Journal for Parasitology 33: 733-755.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants - republication of list of species. Federal Register 42: 36420-36431.

Caudata

Two-toed amphiuma Amphiuma means, photographed by J. D. Wilson.


Belongs within: Euskelia.
Contains: Cryptobranchoidei, Plethodontidae, Salamandridae.

The Caudata are the salamanders, one of the three major groups of living amphibian-grade tetrapods. The earliest known caudates appear in the Jurassic, with both stem- and crown-group taxa known from this period. The neotenic Sirenidae (sirens) have sometimes been regarded as the sister taxon to the remaining caudates, or even classified outside the caudates, but most recent phylogenies have placed them in a more nested position.

Synapomorphies (from Milner 1988): Vertebrae with bicipital rib-bearers and basapophyses; atlas with tuberculum interglenoideum with distinct articulation facets; stapes with large footplate and short slender stylus; basibranchial I with radial horns in metamorphosed individuals; co-ossified scapulocoracoid.

<==Caudata [Batrachoidei, Caudati, Mecodonta, Saurobatrachii, Urodelia]
    |  i. s.: Liaoxitriton zhongjiani ZBH03
    |         Ramonellus Nevo & Estes 1969 RCQ03
    |         Comonecturoides M88
    |         Prosiren M88
    |--Marmorerpeton GS03
    `--+--+--Karaurus sharovi Ivakhnenko 1978 GS03, RCQ03
       |  `--Kokartus GS03
       `--Urodela [Ambystomatoidea, Neocaudata, Salamandroidea] GS03
            |  i. s.: Noterpeton Rage et al. 1993 [Noterpetontidae] GM01
            |           `--*N. bolivianum Rage et al. 1993 GM01
            |         Galverpeton M88
            |         Diemictylus viridescens D56
            |         Valdotriton gracilis Evans & Milner 1996 GS01, RCQ03
            |--+--Cryptobranchoidei GS03
            |  `--+--Jeholotriton paradoxus GS03, ZBH03
            |     `--+--Laccotriton subsolanus GS03, GS01
            |        `--Sinerpeton Gao & Shubin 2001 GS01
            |             `--*S. fengshanensis Gao & Shubin 2001 GS01
            `--Diadectosalamandroidei FG06
                 |--Plethosalamandroidei FG06
                 |    |--Rhyacotriton Dunn 1920 [Rhyacotritonidae] FG06
                 |    |    |--R. cascadae FG06
                 |    |    |--R. kezeri FG06
                 |    |    |--R. olympicus FG06
                 |    |    `--R. variegatus FG06
                 |    `--Xenosalamandroidei FG06
                 |         |--Plethodontidae FG06
                 |         `--Amphiuma Garden 1821 [Amphiumidae] FG06
                 |              |--A. means BF88
                 |              |--A. pholeter FG06
                 |              `--A. tridactylum HNM93
                 `--Hydatinosalamandroidei FG06
                      |--Perennibranchia (see below for synonymy) FG06
                      |    |--Sirenidae [Sirenes, Sirenina, Trachystomata] FG06
                      |    |    |--Pseudobranchus Gray 1825 FG06
                      |    |    |    `--P. striatus FG06
                      |    |    `--Siren Österdam 1766 [=Sirena] FG06
                      |    |         |--S. intermedia FG06
                      |    |         `--S. lacertina FG06
                      |    `--Proteidae [Proteoidea] FG06
                      |         |--Phanerobranchus Leuckart 1821 [Phanerobranchoidea] FG06
                      |         |--Hylaeobatrachus Dollo 1884 [Hylaeobatrachidae] FG06
                      |         |--Menobrachus Harlan 1825 [Menobranchida] FG06
                      |         |--Necturus Rafinesque 1819 [incl. Exobranchia; Necturi, Necturina] FG06
                      |         |    |--N. alabamensis AA87
                      |         |    |--N. beyeri FG06
                      |         |    `--N. maculosus (Rafinesque 1818) W08
                      |         `--Proteus Laurenti 1768 Sw86 (see below for synonymy)
                      |              `--P. anguinus Laurenti 1768 Sw86 (see below for synonymy)
                      |                   |--P. a. anguinus N10
                      |                   `--P. a. parkelj N10
                      `--Treptobranchia FG06
                           |--Salamandridae FG06
                           `--Ambystomatidae [Ambystomina] FG06
                                |--Dicamptodon Strauch 1870 [Dicamptodontidae, Dicamptodontinae] FG06
                                |    |--D. aterrimus FG06
                                |    |--D. ensatus FG06
                                |    `--D. tenebrosus FG06
                                `--Ambystoma Tschudi 1838 FG06 (see below for synonymy)
                                     |  i. s.: A. annulatum Cope 1886 W08
                                     |         A. californiense FG06
                                     |         A. dumerilii FG06
                                     |         A. gracile (Baird 1859) W08
                                     |         A. jeffersonianum W-CL06
                                     |         A. laterale Hallowell 1875 M68
                                     |         A. macrodactylum Baird 1849 W08
                                     |           |--A. m. macrodactylum W08
                                     |           |--A. m. columbianum Baird 1849 W08
                                     |           `--A. m. croceum Baird 1849 W08
                                     |         A. maculatum (Shaw 1802) W08
                                     |         A. opacum (Gravenhorst 1807) W08
                                     |         A. ordinarium FG06
                                     |         A. platineum Cope 1867 M68
                                     |         A. talpoideum (Holbrook 1838) W08
                                     |         A. texanum (Matthes 1855) W08
                                     |         A. tremblayi Comeau 1943 M68
                                     |--A. cingulatum FG06
                                     `--+--A. mexicanum FG06
                                        `--A. tigrinum (Green 1825) FG06, W08
                                             |--A. t. tigrinum W08
                                             `--A. t. mavortium (Baird 1850) W08

Ambystoma Tschudi 1838 FG06 [incl. Bathysiredon Sh86, Rhyacosiredon Sh86, Siredon Wagler 1830 Sh86, FG06; Soredontina]

Perennibranchia [Amphipneusta, Branchiata, Branchiuromalgaei, Diplopneumena, Dysmolgae, Externibranchia, Manentibranchia, Meantia, Perennibranchiata, Ramibranchia] FG06

Proteus Laurenti 1768 Sw86 [incl. Hypochthon Merrem 1820 FG06, Larvarius FG06; Hypochthonina]

Proteus anguinus Laurenti 1768 Sw86 [incl. Hypochthon carrarae Fitzinger 1850 N10, H. freyeri Fitzinger 1850 N10, H. haidingeri Fitzinger 1850 N10, H. laurentii Fitzinger 1850 N10, H. schreibersii Fitzinger 1850 N10, H. xanthostictus Fitzinger 1850 N10, H. zoisii Fitzinger 1850 N10]

*Type species of generic name indicated

REFERENCES

[AA87] Avise, J. C., J. Arnold, R. M. Ball, E. Bermingham, T. Lamb, J. E. Neigel, C. A. Reeb & N. C. Saunders. 1987. Intraspecific phylogeography: The mitochondrial DNA bridge between population genetics and systematics. Annual Review of Ecology and Systematics 18: 489-522.

[BF88] Bishop, M. J., & A. E. Friday. 1988. Estimating the interrelationships of tetrapod groups on the basis of molecular sequence data. In The Phylogeny and Classification of the Tetrapods, vol. 1. Amphibians, Reptiles, Birds (M. J. Benton, ed.) pp. 33-58. Clarendon Press: Oxford.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1-370.

[GS01] Gao, K.-Q., & N. H. Shubin. 2001. Late Jurassic salamanders from northern China. Nature 410: 574-577.

[GS03] Gao, K.-Q., & N. H. Shubin. 2003. Earliest known crown-group salamanders. Nature 422: 424-428.

[GM01] Gayet, M., L. G. Marshall, T. Sempere, F. J. Meunier, H. Cappetta & J-C. Rage. 2001. Middle Maastrichtian vertebrates (fishes, amphibians, dinosaurs and other reptiles, mammals) from Pajcha Pata (Bolivia). Biostratigraphic, palaeoecologic and palaeobiostratigraphic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 169: 39-68.

[HNM93] Hedges, S. B., R. A. Nussbaum & L. R. Maxson. 1993. Caecilian phylogeny and biogeography inferred from mitochondrial DNA sequences of the 12S rRNA and 16S rRNA genes (Amphibia, Gymnophiona). Herpetological Monographs 7: 64-76.

[M68] Maslin, T. P. 1968. Taxonomic problems in parthenogenetic vertebrates. Systematic Zoology 17 (3): 219-231.

[M88] Milner, A. R. 1988. The relationships and origin of living amphibians. In The Phylogeny and Classification of the Tetrapods, vol. 1. Amphibians, Reptiles, Birds (M. J. Benton, ed.) pp. 59-102. Clarendon Press: Oxford.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[RCQ03] Ruta, M., M. I. Coates & D. L. J. Quicke. 2003. Early tetrapod relationships revisited. Biological Reviews 78: 251-345.

[Sh86] Shaffer, H. B. 1986. Utility of quantitative genetic parameters in character weighting. Systematic Zoology 35 (1): 124-134.

[Sw86] Sweet, S. S. 1986. Caudata. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 734-740. E. J. Brill / Dr. W. Backhuys: Leiden.

[W08] Warburg, M. R. 2008. Temporal variations in body dimensions followed for 25 years in a breeding population of adult salamanders, with a partial review on other Urodela (Amphibia, Urodela, Salamandridae). Senckenbergiana Biologica 88 (1): 81-101.

[W-CL06] Watkins-Colwell, G. J., T. A. Leenders, B. T. Roach, D. J. Drew, G. Dancho & J. Yuckienuz. 2006. New distribution records for amphibians and reptiles in Connecticut, with notes on the status of an introduced species. Bulletin of the Peabody Museum of Natural History 47 (1-2): 47-62.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807-814.

Lioconcha

Lioconcha (Lioconcha) castrensis, photographed by Guido & Philippe Poppe.


Belongs within: Veneridae.

Lioconcha is a genus of venerid bivalves found in the tropical Pacific and East Indies, with a fossil record dating to the Pliocene. It is divided between two subgenera, Lioconcha (Lioconcha) with a smooth shell surface with zigzag streaks or spots, and Sulcilioconcha with concentric ribs (Cox et al. 1969).

Characters (from Cox et al. 1969): Hinge strong, resembling Pitar; lunule present; pallial sinus shallow.

<==Lioconcha
    |  i. s.: L. dautzenbergi LS96
    |         L. fastigiata (Sowerby 1851) [=Cytherea fastigiata] H09
    |         L. tigrina (Lamarck 1816) BW09
    |--L. (Lioconcha) LS96
    |    |--L. (L.) annettae Lamprell & Whitehead 1990 LS96
    |    |--L. (L.) castrensis (Linnaeus 1758) (see below for synonymy) LS96
    |    |--L. (L.) ornata (Dillwyn 1817) [=Venus ornata; incl. Cytherea picta Lamarck 1818] LS96
    |    `--L. (L.) polita (Röding 1798) [=Venus polita; incl. Cytherea sulcatina Lamarck 1818] LS96
    `--L. (Sulcilioconcha) LS96
         |--L. (S.) melharteae Lamprell & Stanisic 1996 LS96
         |--L. (S.) philippinarum (Hanley 1844) [=Cytherea philippinarum] LS96
         `--L. (S.) richerdeforgesi Lamprell & Stanisic 1996 LS96

Lioconcha (Lioconcha) castrensis (Linnaeus 1758) [=Venus castrensis; incl. Cytherea ornata Lamarck 1818 non Dillwyn 1817] LS96

*Type species of generic name indicated

REFERENCES

[BW09] Bryce, C. & C. Whisson. 2009. The macromolluscs of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 177-208.

Cox, L. R., N. D. Newell et al. 1969. Treatise on Invertebrate Paleontology pt N. Bivalvia vol. 2. The Geological Society of America, Inc. and The University of Kansas.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[LS96] Lamprell, K. L., & J. Stanisic. 1996. Callista, Lioconcha and Pitar in New Caledonia and adjacent waters (Mollusca, Veneridae). Molluscan Research 17: 27-48.

Chione

Chione cancellata, photographed by Edward T. Babinski.


Belongs within: Veneridae.

Chione is a genus of venerid bivalves found in the Americas and the South Pacific, with the earliest records from the Oligocene (Cox et al. 1969).

Characters (from Cox et al. 1969): Cordate; lunule and escutcheon mostly well-defined; concentric sculpture strong, frilled in some forms.

<==Chione Megerle 1811 M27
    |--*C. cancellata M27
    |--C. californiensis M27
    |--C. (Timoclea) concinna Viader 1951 L96
    |--C. costellifera (Adams 1850) [=Venus costellifera] H09
    |--C. dysera F66
    |--C. fluctifraga M27
    |--C. gayi M27
    |--C. halli Pritchard 1895 F27
    |--C. infans Smith 1885 H09
    |--C. latilirata GS79
    |--C. marica (Linnaeus 1758) [=Venus marica] H09
    |--C. martini Finlay 1927 [=Venus pulcherrima Martin 1883 non Deshayes 1860] H09
    |--C. nuttalli M27
    |--C. phoenicopterus (Romer 1869) [=Caryatis phoenicopterus] H09
    |--C. recognita (Smith 1885) [=Venus recognita] H09
    |--C. regularis (Smith 1885) [=Cytherea regularis] H09
    |--C. roberti Pritchard 1906 [=C. halli Tate 1900 non Pritchard 1895] F27
    |--C. scabra (Hanley 1845) [=Venus scabra] H09
    |--C. semperi Dunker 1871 H09
    |--C. similicentrifuga Viader 1951 L96
    |--C. subnodulosa (Hanley 1845) [=Venus subnodulosa] H09
    |--C. torresica Reeve 1863 H09
    `--C. undulosa (Lamarck 1819) [=Venus undulosa] H09

Nomen nudum: Chione multitaeniata Tate 1893 F27

*Type species of generic name indicated

REFERENCES

Cox, L. R., N. D. Newell et al. 1969. Treatise on Invertebrate Paleontology pt N. Bivalvia vol. 2. The Geological Society of America, Inc. and The University of Kansas.

[F27] Finlay, H. J. 1927. New specific names for austral Mollusca. Transactions and Proceedings of the New Zealand Institute 57: 488-533.

[F66] Fraser, L. 1866. Communication of a list of mollusks collected by R. Swinhoe, Esq., in Formosa. Proceedings of the Zoological Society of London 1866: 146.

[GS79] Gore, R. H., & L. E. Scotto. 1979. Crabs of the family Parthenopidae (Crustacea Brachyura: Oxyrhyncha) with notes on specimens from the Indian River region of Florida. Memoirs of the Hourglass Cruises 3 (6): 1-98.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[L96] Loch, I. 1996. Notice: R. Viader collection. Molluscan Research 17: 110.

[M27] Marwick, J. 1927. The Veneridae of New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 567-636.

Pitar

Pitar (Pitarina) citrina. Photo from here.


Belongs within: Veneridae.

Pitar is a genus of venerid bivalves found worldwide, first recorded from the Eocene (Cox et al. 1969).

Characters (from Cox et al. 1969): Oval or subtrigonal, smooth or finely concentrically lamellate; lunule superficial, escutcheon not defined; hinge with 2b triangular, joined to a thin 2a, 3a and 1 separate.

<==Pitar
    |  i. s.: P. nipponica Kuroda & Habe 1971 LS96
    |         P. subpellucidus LS96
    |--*P. (Pitar) tumens [=Venus tumens] M27
    |--P. (Hyphantosoma) LS96
    |    |--P. (H.) intricata (Dautzenberg 1907) [=Meretrix (Pitar) intricata] LS96
    |    `--P. (H.) sculpturatus (Marshall 1918) [=Macrocallista sculpturata] M27
    `--P. (Pitarina) LS96
         |--P. (P.) affinis (Gmelin 1791) LS96 (see below for synonymy)
         |--P. (P.) citrina (Lamarck 1818) (see below for synonymy) LS96
         |--P. (P.) healyi Lamprell & Stanisic 1996 LS96
         |--P. (P.) japonicus Kuroda & Kawamoto 1956 [=P. (Agriopoma) japonicum] LS96
         |--P. (P.) nancyae Lamprell & Whitehead 1990 LS96
         |--P. (P.) noguchii Habe 1958 [=P. (Agriopoma) noguchii] LS96
         |--P. (P.) pellucidus (Lamarck 1818) LS96 (see below for synonymy)
         |--P. (P.) potteri Healy & Lamprell 1992 LS96
         |--P. (P.) prora (Conrad 1837) [=Venus prora] LS96
         |--P. (P.) sophiae (Angas 1877) [=Cytherea sophiae] LS96
         `--P. (P.) spoori Lamprell & Whitehead 1990 LS96

Pitar (Pitarina) affinis (Gmelin 1791) LS96 [=Venus affinis LS96; incl. Cytherea inflata Sowerby 1853 LS96, Gafrarium inflatum H09]

Pitar (Pitarina) citrina (Lamarck 1818) [=Cytherea citrina, Dione citrina, Pitar (Pitarina) citrinus; incl. D. ustulata Reeve 1863] LS96

Pitar (Pitarina) pellucidus (Lamarck 1818) LS96 [=Cytherea pellucidus LS96; incl. Dione australica Reeve 1863 LS96, Gafrarium australica H09]

*Type species of generic name indicated

REFERENCES

Cox, L. R., N. D. Newell et al. 1969. Treatise on Invertebrate Paleontology pt N. Bivalvia vol. 2. The Geological Society of America, Inc. and The University of Kansas.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[LS96] Lamprell, K. L., & J. Stanisic. 1996. Callista, Lioconcha and Pitar in New Caledonia and adjacent waters (Mollusca, Veneridae). Molluscan Research 17: 27-48.

[M27] Marwick, J. 1927. The Veneridae of New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 567-636.

Cobanocythere

Features of Cobanocythere labiata, from Hartmann (1959). 117: Right valve; 118: hinge; 119: end of body; 120: female antenna; 121: male copulatory organ.


Belongs within: Cytheroidea.

Cobanocythere is a genus of marine interstitial ostracods found around the world. Members of this genus are united by the presence of three adductor muscle scars, a rudimentary or absent bracnhial plate on the mandible, and a copulatory organ with a ventro-median extension and short penis. The genera Microcytheridea and Platymicrocythere, each previously distinguished by features of the external shell, share these characteristics and hence were synonymised by Gottwald (1983).

Characters (from Hartmann 1959*): Shell similar to Microcythere, with large radial pore canals; hinge with cardinal teeth or bar. Eyes reduced. Distinct adductor muscle scars. First antenna 6-segmented; distalmost segment reduced. Second antenna 4-segmented, distally with one large and one small seta. Mandibular palp slender, 3-segmented; basis of mandible with long tooth and several shorter teeth. Maxilla with small exopod, with only a few rays. Maxilliped well-developed, exopod one-segmented, distal segment not developed. Bristles of maxilliped and exopod slim and long. Thoracopods 4-segmented.

*Translated from original German with the help of Google Translate; any errors of translation are my own.

<==Cobanocythere Hartmann 1959 DH86 [incl. Microcytheridea Hartmann 1965 G83, Platymicrocythere Schormikov 1975 G83]
    |  i. s.: C. angularis Gottwald & Scherner 1980 DH86
    |         C. axi Gottwald 1983 DH86
    |         C. chilensis Gottwald 1983 [=Microcytheridea labiata Hartmann 1965 non C. labiata Hartmann 1959] DH86
    |         C. cuneiformis Hartmann 1974 DH86
    |         C. hartmanni Reys 1961 DH86
    |         C. longiantennata (Marinov 1962) DH86 [=Microcythere longiantennata G83]
    |         ‘Microcythere’ subterranea Hartmann 1954 [=Cobanocythere parva Gottwald 1983] DH86
    |         C. subterranoides Hartmann 1974 DH86
    |         C. syltensis Gottwald 1983 DH86
    |--C. elongata Gottwald 1983 G83
    |--C. guttaeformis Gottwald 1983 G83
    |--C. santacruzensis Gottwald 1983 G83
    |--+--*C. ‘subterranea’ Hartmann 1959 non Microcythere subterranea Hartmann 1954 G83
    |  |--C. arenicola Gottwald 1983 G83
    |  `--C. japonica Schornikov 1975 G83
    |--+--C. tokiokai (Schornikov 1975) [=Platymicrocythere tokiokai] G83
    |  `--+--C. labiata Hartmann 1959 G83
    |     `--C. mielkei Gottwald 1983 G83
    `--+--C. fernandinensis Gottwald 1983 G83
       |--C. hoodensis Gottwald 1983 G83
       |--C. lanceolata Gottwald 1983 G83
       |--C. pacifica Gottwald 1983 G83
       |--C. psammophila Gottwald 1983 G83
       `--C. sublitoralis Gottwald 1983 G83

*Type species of generic name indicated

REFERENCES

[DH86] Danielopol, D. L., & G. Hartmann. 1986. Ostracoda. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 265–294. E. J. Brill/Dr. W. Backhuys: Leiden.

[G83] Gottwald, J. 1983. Interstitielle Fauna von Galapagos. XXX. Podocopida 1 (Ostracoda). Mikrofauna des Meeresbodens 90: 1–187.

Hartmann, G. 1959. Zur Kenntnis der lotischen Lebensbereiche der pazifischen Küste von El Salvador unter besonderer Berücksichtigung seiner Ostracodenfauna. Kieler Meeresforschungen 76 (2): 187–241.

Marama

Marama murdochi, from Marwick (1927).


Belongs within: Veneridae.

Marama is a genus of venerid bivalve recorded from the Miocene to the Pliocene of New Zealand. It contains two subgenera: Marama (Marama) with the left median cardinal tooth joined to the anterior, the right posterior cardinal broad and deeply divided, and the lunule very large and pouting, and Hina with the median and anterior cardinals not joined, th escutcheon well marked, the posterior end abruptly truncated, and the right posterior cardinal much stronger than the median (Marwick 1927).

Characters (from Marwick 1927): Moderate to tubercular anterior lateral tooth in left valve, hinge narrow, marginal crenulations very fine.

<==Marama Marwick 1927 M27
    |--M. (Marama) M27
    |    |--*M. (M.) murdochi Marwick 1927 M27
    |    |--M. (M.) hurupiensis Marwick 1927 M27
    |    |--M. (M.) ovata Marwick 1927 M27
    |    `--M. (M.) pristina Marwick 1927 M27
    `--M. (Hina Marwick 1927) M27
         |--M. (*H.) pinguis Marwick 1927 M27
         |--M. (H.) hendersoni Marwick 1927 M27
         |--M. (H.) mackenziei Marwick 1927 M27
         |--M. (H.) tumida (Marshall 1918) [=Dosinia tumida non Artemis tumida Gray 1838 (not preoc. if dif. gen.)] M27
         |--M. (H.) vaga Marwick 1927 M27
         `--M. (H.) williamsi Marwick 1927 M27

*Type species of generic name indicated

REFERENCES

[M27] Marwick, J. 1927. The Veneridae of New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 567-636.

Eumarcia

Eumarcia fumigata, photographed by Denis Riek.


Belongs within: Veneridae.

Eumarcia is a genus of venerid bivalves found from the Oligocene to the present of Australia and New Zealand. The genus is divided into two subgenera, Eumarcia (Eumarcia) with a superficial lunule and escutcheon and no concentric grooves on the shell surface, and Atamarcia with a well-impressed lunule and escutcheon and generally strong concentric grooves on the shell surface (Marwick 1927).

Characters (from Marwick 1927): Shell moderate to large size, oval, margins smooth, pallial sinus ample.

<==Eumarcia Iredale 1925 M27
    |--E. (Eumarcia) M27
    |    |--*E. (E.) fumigata [=Venus fumigata; incl. V. nitida] M27
    |    |--E. (E.) altilunula Marwick 1927 M27
    |    |--E. (E.) kaawaensis Marwick 1927 M27
    |    |--E. (E.) pareoraensis (Suter 1917) [=Macrocallista pareoraensis] M27
    |    `--E. (E.) plana Marwick 1927 M27
    `--E. (Atamarcia Marwick 1927) M27
         |--E. (*A.) sulcifera Marwick 1927 M27
         |--E. (A.) benhami Marwick 1927 (see below for synonymy) M27
         |--E. (A.) crassa Marwick 1927 M27
         |--E. (A.) crassatelliformis Marwick 1927 M27
         |--E. (A.) curta (Hutton 1873) [=Tapes curta, Paphia (Ruditapes) curta] M27
         |--E. (A.) enysi (Hutton 1873) [=Cytherea enysi] M27
         |--E. (A.) thomsoni Marwick 1927 M27
         `--E. (A.) trigonalis Marwick 1927 M27

Eumarcia (Atamarcia) benhami Marwick 1927 [=Venus sulcata Hutton 1875 non Lamarck 1835, Cytherea (Circomphalus) sulcata] M27

*Type species of generic name indicated

REFERENCES

[M27] Marwick, J. 1927. The Veneridae of New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 567-636.

Periglypta

Periglypta puerpera, from here.


Belongs within: Veneridae.

Periglypta is a genus of venerid bivalves that first appeared in the Oligocene (Cox et al. 1969).

Characters (from Cox et al. 1969): Quadrate, heavy, pallial sinus rounded; sculpture cancellate.

<==Periglypta Jukes-Brown 1914 KS02 [=Cytherea (preoc.) M27]
    |  i. s.: ‘Venus’ anadyomene Anton 1839 [=Cytherea anadyomene] H09
    |         ‘Venus’ calophylla Philippi 1836 [=Cytherea calophylla] H09
    |         ‘Chione’ capricornea Hedley 1908 [=Cytherea capricornea] H09
    |         ‘Venus’ chemnitzii Hanley 1845 [=Cytherea chemnitzii] H09
    |         ‘Chione’ embrithes Melvill & Standen 1899 [=Cytherea embrithes] H09
    |         ‘Venus’ foliacea Philippi 1846 [=Cytherea foliacea] H09
    |         ‘Cytherea’ hebraea Lamarck 1818 LS96 [=Macrocallista hebraea H09]
    |         ‘Venus’ laqueata Sowerby 1853 [=Cytherea laqueata] H09
    |         ‘Venus’ lionota Smith 1885 [=Cytherea lionota] H09
    |         ‘Dosinia’ listeri Gray 1838 [=Cytherea listeri] H09
    |         ‘Venus’ reticulata Linnaeus 1758 [=Cytherea reticulata] H09
    |         ‘Cytherea’ tenuis Hall & Meek 1854 F27
    |         ‘Venus’ tiara Dillwyn 1817 [=Cytherea tiara] H09
    |         ‘Venus’ toreuma Gould 1850 [=Cytherea toreuma] H09
    |         ‘Venus’ torresiana Smith 1884 [=Cytherea torresiana] H09
    |--*P. (Periglypta) puerpera (Linnaeus 1758) KS02 [=Venus puerpera KS02, *Cytherea puerpera M27]
    `--P. (Proxichione Iredale 1929) KS02

*Type species of generic name indicated

REFERENCES

Cox, L. R., N. D. Newell et al. 1969. Treatise on Invertebrate Paleontology pt N. Bivalvia vol. 2. The Geological Society of America, Inc. and The University of Kansas.

[F27] Finlay, H. J. 1927. New specific names for austral Mollusca. Transactions and Proceedings of the New Zealand Institute 57: 488-533.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[KS02] Kanjilal, S., & M. S. Srinivasan. 2002. New bivalves from the Miocene of Little Andaman Island, Bay of Bengal. Journal of the Geological Society of India 60: 527-536.

[LS96] Lamprell, K. L., & J. Stanisic. 1996. Callista, Lioconcha and Pitar in New Caledonia and adjacent waters (Mollusca, Veneridae). Molluscan Research 17: 27-48.

[M27] Marwick, J. 1927. The Veneridae of New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 567-636.

Tellinoidea

Abra nitida, from the National Museum of Wales.


Belongs within: Heterodonta.
Contains: Semele, Tellina, Macoma.

The Tellinoidea are a group of infaunal bivalves that first appeared in the Upper Triassic (Cox et al. 1969). With their elongate, mobile siphons, tellinoids tend to burrow relatively deep within the sediment, using the siphons to pick up food particles from the sediment surface. The Tellinidae have more or less elongate shells with unequal valves and an external ligament (Cox et al. 1969).

Characters (from Cox et al. 1969): Shells mostly inequilateral with external ligament or, if sunken, in pit on hinge plate, not in chondrophore; cardinal hinge teeth two in either valve, tending to be bifid, lateral teeth well developed in most families; adductor muscle scars connected by pallial line with distinct sinus. Animal with two elongate siphons that are not fused.

<==Tellinoidea [Tellinacea]
    |--Donacidae G88
    |    |--Iphigenia G88
    |    |    |--I. centralis G88
    |    |    `--I. truncata G88
    |    `--Donax P98
    |         |--D. brazieri Smith 1891 H09
    |         |--D. deltoides Lamarck 1818 H09
    |         |--D. faba Gmelin 1791 H09
    |         |--D. nitida Reeve 1854 H09
    |         |--D. trunculus M62
    |         `--D. vittatus D56
    |--Solecurtidae G88
    |    |--Novaculininae G88
    |    |    |--Novaculina G88
    |    |    `--Sinonovacula constricta G88, KC11
    |    `--Solecurtus Blainville 1824 F27a
    |         |--*S. strigillatus [=Solen strigillatus] F27a
    |         |--S. bensoni Finlay 1924 F27a
    |         |--S. chattonensis Finlay 1924 F27a
    |         |--S. ellipticus Dana 1849 F27b
    |         |--S. evolutus Finlay 1924 F27a
    |         |--S. guaymasensis Lowe 1935 PP78
    |         |--S. murrayvianus Finlay 1927 [=S. ellipticus Tate 1887 non Dana 1849] F27b
    |         `--S. (Psammodia) planulatus Dana 1849 F71
    |--Semelidae GW02
    |    |--Semele BW09
    |    |--Theora HJ08
    |    |    |--T. fragilis (Adams 1856) [=Neaera fragilis] H09
    |    |    `--T. lubrica Gould 1861 HJ08
    |    |--Abra GW02
    |    |    |--A. nitida GO06
    |    |    |--A. prismatica GW02
    |    |    |--A. profundorum TH77
    |    |    `--A. truncata Hedley 1906 H09
    |    `--Leptomya Adams 1864 P61
    |         |--*L. cochlearis P61
    |         |--L. gravida Hamley 1879 MG-H11
    |         |--L. psittacus Hanley 1883 H09
    |         |--L. retiaria (Hutton 1885) (see below for synonymy) F27a
    |         |    |--L. r. retiaria P61
    |         |    `--L. r. aucklandica Powell 1955 P61
    |         |--L. simplex Marwick 1926 F27a
    |         `--L. suteri (Smith 1898) [=Macoma suteri] F27a
    `--Tellinidae MG-H11
         |--Eurytellina F27a
         |--Elliptotellina F27a
         |--Tellina BW09
         |--Palaeomoera inconspicua [=Psammobia inconspicua] F27b
         |--Quadrans gargadia (Linnaeus 1758) BW09
         |--Tellinides cockburnensis (Kendrick & Brearley 1984) MG-H11
         |--Macoma F27a
         |--Pseudarcopagia F27b
         |    |--*P. decussata (Lamarck 1818) [=Tellina decussata; incl. P. victoriae] F27a
         |    `--P. marshalli Finlay 1927 (see below for synonymy) F27b
         |--Barytellina Marwick 1924 F27a
         |    |--*B. crassidens Marwick 1924 F27a
         |    `--B. anomalodonta Finlay 1924 F27a
         |--Zearcopagia Finlay 1927 F27a
         |    |--*Z. disculus (Deshayes 1855) [=Tellina disculus, Arcopagia disculus, Pseudarcopagia disculus] F27a
         |    `--Z. piratica (Hedley 1918) [=Pseudarcopagia piratica] F27a
         |--Exotica BW09
         |    |--E. obliquaria (Deshayes 1854) BW09
         |    `--E. rhomboides (Quoy & Gaimard 1833) BW09
         |--Macomona Finlay 1927 F27a
         |    |--*M. liliana (Iredale 1915) P61 [=Tellina liliana P61, Macoma liliana F27a]
         |    |--M. basedowi [=Tellina basedowi] F27a
         |    `--M. deltoidalis [=Tellina deltoidalis] F27a
         |--Solyma Conrad 1870 S98
         |    |--*S. lineolatum Conrad 1870 S98
         |    |--S. flemingae Stilwell 1998 S98
         |    |--S. levis Stephenson 1923 S98
         |    `--S. stewarti Stephenson 1952 S98
         `--Angulus Megerle 1811 P61
              |  i. s.: A. valtonis (Hanley 1844) BD86
              |--A. (Angulus) P61
              |    |--*A. (A.) lanceolatus [=Tellina lanceolata] P61
              |    |--A. (A.) gaimardi (Iredale 1915) P61
              |    |--A. (A.) meropsis (Dall 1900) [=Tellina (A.) meropsis] PP78
              |    `--A. (A.) spenceri (Suter 1907) P61
              `--A. (Peronidia Dall 1900) P61
                   |--A. (*P.) albicans [=Tellina albicans] P61
                   `--A. (P.) edgari (Iredale 1915) P61

Leptomya retiaria (Hutton 1885) [=Tellina retiaria; incl. T. lintea Hutton 1873 (preoc.), T. perconfusa Iredale 1915] F27a

Pseudarcopagia marshalli Finlay 1927 [=Tellina (Arcopagia) inconspicua Marshall 1918 non T. inconspicua Broderip & Sowerby 1829] F27b

*Type species of generic name indicated

REFERENCES

[BD86] Barash, A., & Z. Danin. 1986. Further additions to the knowledge of Indo-Pacific Mollusca in the Mediterranean Sea (Lessepsian migrants). Spixiana 9 (2): 117–141.

[BW09] Bryce, C., & C. Whisson. 2009. The macromolluscs of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 177–208.

Cox, L. R., N. D. Newell et al. 1969. Treatise on Invertebrate Paleontology pt N. Bivalvia vol. 2. The Geological Society of America, Inc. and The University of Kansas.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[F27a] Finlay, H. J. 1927a. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[F27b] Finlay, H. J. 1927b. New specific names for austral Mollusca. Transactions and Proceedings of the New Zealand Institute 57: 488–533.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[GO06] Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrödl & M. K. Nishiguchi. 2006. Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons. Proceedings of the National Academy of Sciences of the USA 103 (20): 7723–7728.

[GW02] Giribet, G., & W. Wheeler. 2002. On bivalve phylogeny: a high-level analysis of the Bivalvia (Mollusca) based on combined morphology and DNA sequence data. Invertebrate Biology 121 (4): 271–324.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[HJ08] Huisman, J. M., D. S. Jones, F. E. Wells & T. Burton. 2008. Introduced marine biota in Western Australian waters. Records of the Western Australian Museum 25 (1): 1–44.

[KC11] Kocot, K. M., J. T. Cannon, C. Todt, M. R. Citarella, A. B. Kohn, A. Meyer, S. R. Santos, C. Schander, L. L. Moroz, B. Lieb & K. M. Halanych. 2011. Phylogenomics reveals deep molluscan relationships. Nature 477: 452–456.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[M62] Monniot, F. 1962. Recherches sur les graviers a Amphioxus de la région de Banyuls-sur-Mer. Vie et Milieu 13: 231–322.

[PP78] Poorman, F. L., & L. H. Poorman. 1978. Additional molluscan records from Bahía de Los Angeles, Baja California Norte. Veliger 20 (4): 369–374.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

[S98] Stilwell, J. D. 1998. Late Cretaceous Mollusca from the Chatham Islands, New Zealand. Alcheringa 22 (1): 29–85.

[TH77] Tendal, O. S., & R. R. Hessler. 1977. An introduction to the biology and systematics of Komokiacea (Textulariina, Foraminiferida). Galathea Report 14: 165–194.

Therapsida

Reconstruction of Lemurosaurus pricei, by Theropsida.


Belongs within: Synapsida.
Contains: Dicynodontia, Cynodontia.

The Therapsida is the clade of all synapsids except for the basal 'pelycosaurs'. The earliest members of the clade are known from the Late Permian (Kemp 1988).

Synapomorphies (from Kemp 1988): Lateral temporal fenestra enlarged; supratemporal lost; shallow groove (external auditory meatus) on posterior face of squamosal; septomaxilla with long posterodorsal process; pineal foramen on raised boss; reflected lamina of angular greatly enlarged as thin sheet; jaw articulation shifted forwards; mobile basipterygoid articulation absent; interpterygoid vacuity reduced; palatal teeth reduced; paroccipital process extending more laterally than ventrally; vomer transversely widened between internal nares; transverse process of pterygoid shifted forward; no ossified intercentra in trunk; ossified sternum; cleithrum reduced, scapular blade narrow, coracoid and procoracoid reduced, screw-shaped glenoid absent; ilium expanded both anteriorly and posteriorly, acetabulum circular, fourth trochanter of femur absent; phalanges reduced.

<==Therapsida
    |  i. s.: Delphinognathus N85
    |         Lycaenops N85
    |         Microurania [Microuraniidae] TT05
    |           `--M. minima Ivakhnenko 1995 TT05
    |         Porosteognathus [Pristerognathidae] TT05
    |         Inostrancevia [Inostranceviidae] TT05
    |           `--I. uralensis Tatarinov 1974 TT05
    |         Scylacosuchus [Scylacosauridae] TT05
    |           `--S. orenburgensis TT05
    |         Moschorhinidae TT05
    |           |--Annatherapsidus petri (Amalitskii 1922) TT05
    |           `--Chthonosaurus velocidens V’yushkov 1955 TT05
    |--Biarmosuchia S03
    |    |  i. s.: Phthinosuchus K88
    |    |--Biarmosuchidae TT05
    |    |    |--Biarmosuchus S03 [incl. Biarmosaurus K88]
    |    |    `--Biarmosuchoides romanovi Tverdochlebova & Ivakhnenko 1994 TT05
    |    `--+--Hipposaurus S03
    |       `--Ictidorhinidae S03
    |            |--Lycaenodon Broom 1925 [incl. Hipposauroides Boonstra 1952] S03
    |            |    `--*L. longiceps Broom 1925 [incl. Hipposaurus rubidgei Broom 1940, *Hipposauroides rubidgei] S03
    |            `--+--Ictidorhinus martinsi S03
    |               `--Burnetiamorpha S03
    |                    |--Lemurosaurus pricei S03
    |                    `--Burnetiidae S03
    |                         |--Burnetia mirabilis S03
    |                         |--Proburnetia S03
    |                         `--Bullacephalus S03
    `--+--Eotitanosuchus [Eotitanosuchia] CL97
       `--Eutherapsida [Anomodontia] S03
            |--+--Dicynodontia S03
            |  `--Theriodontia [Therosauria] S03
            |       |--Gorgonopsia [Gennetotheria] S03
            |       `--Eutheriodontia HK01
            |            |--Cynodontia S03
            |            `--Therocephalia S03
            |                 |  i. s.: Lycosuchus HK01
            |                 |         Promoschorhynchus S03
            |                 |         Olivieria S03
            |                 |--Bauria HK01 [Bauriamorpha, Baurioidea K88]
            |                 `--Whaitsiidae K88
            `--Dinocephalia S03
                 |  i. s.: Rhopalodon TT05
                 |         Sinophoneus Cheng & Ji 1996 Cl97
                 |         Microsyodon orlovi CL97
                 |         Estemmenosuchus [Estemmenosuchidae] TT05
                 |--Stenocybus Cheng & Li 1997 [Stenocybusidae] CL97
                 |    `--*S. acidentatus Cheng & Li 1997 CL97
                 `--+--Ulemosaurus TT05 [Tapinocephalia CL97, Tapinocephalidae]
                    |    `--U. gigas TT05
                    `--Anteosauridae [Anteosauria] CL97
                         |--Titanophoneus adamanteus (Orlov 1958) CL97, TT05
                         |--Deuterosaurus jubilaei (von Nopcsa 1928) TT05
                         |--Syodon CL97
                         |--Anteosaurus CL97
                         `--Australosyodon CL97

*Type species of generic name indicated

REFERENCES

[CL97] Cheng Z. & Li J. 1997. A new genus of primitive dinocephalian - the third report on Late Permian Dashankou lower tetrapod fauna. Vertebrata PalAsiatica 35 (1): 35-43.

[HK01] Hopson, J. A., & J. W. Kitching. 2001. A probainognathian cynodont from South Africa and the phylogeny of nonmammalian cynodonts. Bulletin of the Museum of Comparative Zoology 156 (1): 5-35.

[K88] Kemp, T. S. 1988. Interrelationships of the Synapsida. In The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals (M. J. Benton, ed.) pp. 1-22. Clarendon Press: Oxford.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[S03] Sidor, C. A. 2003. The naris and palate of Lycaenodon longiceps (Therapsida: Biarmosuchia), with comments on their early evolution in the Therapsida. Journal of Paleontology 77 (5): 977-984.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27-77.

Synapsida

Mounted skeleton of Edaphosaurus at the Field Museum of Natural History, Chicago, photographed by Jason Cross.


Belongs within: Reptiliomorpha.
Contains: Therapsida.

The Synapsida is the total group for mammals—that is, mammals and all fossil animals more closely related to mammals than to reptiles. The most basal synapsids, the Permian 'pelycosaurs', were still fairly 'reptilian' in their general appearance, and include such well-known exemplars as the carnivorous Dimetrodon and the herbivorous Edaphosaurus.

Synapomorphies (from Kemp 1988): Temporal fenestra bounded above by squamosal and postorbital; occipital plate broad and tilted forwards; post-temporal fenestra small; postparietal single and median; septomaxilla with broad base and dorsal process; medial process of jugal sutured to pterygoid, ectopterygoid primitively present.

<==Synapsida [Edaphosauria, Ophiacodontia, Pelycosauria, Sphenacodontia, Theromorpha, Theropsida]
    |  i. s.: Varanosaurus GKR88
    |         Protoclepsydrops LR95
    |         Mycterosaurus longiceps M06
    |         Archaeothyris florensis M06
    |         Oedaleops GKR88
    |--+--Eothyris LR95 [Eothyrididae K88]
    |  `--Caseidae K88
    `--+--Varanopseidae K88
       |    |--Basicranodon fortsillensis M06
       |    |--Archaeovenator hamiltonensis D-SMS06
       |    |--Mesenosaurus romeri D-SMS06
       |    |--Elliotsmithia longiceps MS01
       |    |--Varanops brevirostris K88
       |    `--Aerosaurus GKR88
       `--+--Ophiacodon GKR88 [Ophiacodontidae K88]
          `--+--Edaphosaurus N85 [Edaphosauridae K88]
             |    `--E. poconias N85
             `--Sphenacodontidae K88
                  |  i. s.: Secodontosaurus BH04
                  |--Haptodus K88
                  `--+--Therapsida K88
                     `--Dimetrodon Cope 1878 BH04
                          |--D. grandis N85
                          |--D. limbatus BH04
                          |--D. milleri BH04
                          |--D. natalis BH04
                          |--D. occidentalis BH04
                          `--D. teutonis Berman et al. 2001 BH04

*Type species of generic name indicated

REFERENCES

[BH04] Berman, D. S., A. C. Henrici, S. S. Sumida & T. Martens. 2004. New remains of Dimetrodon teutonis (Synapsida: Sphenacodontidae) from the Lower Permian of Germany. Annals of Carnegie Museum 73 (2): 48-56.

[D-SMS06] Dias-da-Silva, S., S. P. Modesto & C. L. Schultz. 2006. New material of Procolophon (Parareptilia: Procolophonoidea) from the Lower Triassic of Brazil, with remarks on the ages of the Sanga do Cabral and Buena Vista formations of South America. Canadian Journal of Earth Sciences 43: 1685-1693.

[GKR88] Gauthier, J. A., A. G. Kluge & T. Rowe. 1988. The early evolution of the Amniota. In The Phylogeny and Classification of the Tetrapods, vol. 1. Amphibians, Reptiles, Birds (M. J. Benton, ed.) pp. 103-155. Clarendon Press: Oxford.

[K88] Kemp, T. S. 1988. Interrelationships of the Synapsida. In The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals (M. J. Benton, ed.) pp. 1-22. Clarendon Press: Oxford.

[LR95] Laurin, M., & R. R. Reisz. 1995. A reevaluation of early amniote phylogeny. Zoological Journal of the Linnean Society 113: 165-223.

[M06] Modesto, S. P. 2006. The cranial skeleton of the Early Permian aquatic reptile Mesosaurus tenuidens: implications for relationships and palaeobiology. Zoological Journal of the Linnean Society 146: 345-368.

[MS01] Modesto, S. P., & R. M. H. Smith. 2001. A new Late Permian captorhinid reptile: a first record from the South African Karoo. Journal of Vertebrate Paleontology 21 (3): 405-409.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.