Soricidae

Water shrew, Neomys fodiens, photographed by J. van der Kooij.

Belongs within: Lipotyphla.
Contains: Sorex, Blarina, Cryptotis, Soriculus, Myosorex, Crocidurinae.

The Soricidae are the shrews, the largest modern 'family' of lipotyphlans. The earliest definite Soricidae are recorded from the middle Eocene of North America, but with confirmed examples from Eurasia shortly thereafter and potential Eurasian species even earlier (Rose 2006). Crown group shrews are characterised by the possession of a double temporo-mandibular joint with neomorphic second lower joint surface and of a deep fossa accomodating the internal temporal muscle opening into the medial side of the coronoid process of the mandible, and the absence of the zygomatic arch, but these characters are not yet present in the stem subfamily Heterosoricinae (Butler 1988, Rose 2006).

The majority of crown-group shrews are divided between the subfamilies Soricinae and Crocidurinae, distinguished by features of the mandibular condyle and of P₄ (Butler 1978). Many species of Soricinae also have the teeth pigmented red by the deposition of iron in a superficial aprismatic layer over the white enamel (Rose 2006). A third subfamily, Myosoricinae, has also been recognised for a group of African genera including the mouse shrews Myosorex. A further extinct subfamily, the Limnoecinae, is known from the Middle Miocene to Middle Pliocence of North America (Van Valen 1967). Within the Soricinae, the living tribes Soricini and Neomyini are each found in both Eurasia and the Americas. The Blarinini are present in the Recent fauna only in the Americas, but are also known from the Pliocene and Pleistocene of Eurasia. The fourth tribe, the extinct Allosoricini, are known from from the Middle Miocence to the late Pliocene of Europe (Van Valen 1967). The water shrews of the genus Neomys hunt aquatic prey, and are found in temperate Eurasia.

See also: Meet the shrews (Taxon of the Week: Soricidae).

Characters (from Rose 2006): Mandible with long, pointed angular process; auditory bulla and zygomatic arch absent. Enlarged anterior incisors, lower incisor procumbent and often with serrated or multilobed crown; remaining antemolar teeth reduced; P₄ primitively unicuspid and triangular; P⁴/M₁ shear developed; dilambdodont molars. Entoconid usually large and separate from hypolophid.

<==Soricidae
    |  i. s.: Arctisorex polaris TH03
    |         Mafia Reumer 1984 P04
    |           `--M. csarnotensis Reumer 1984 P04
    |         Ruwenzorisorex suncoides IT07
    |         Limnoecinae V67
    |           |--Angustidens Repenning 1967 V67
    |           `--Limnoecus Stirton 1930 V67
    |         Podihik Deraniyagala 1958 V67
    |--Heterosoricinae B88
    |    |--Domnina Cope 1873 [incl. Miothen Cope 1873, Protosorex Scott 1895] V67
    |    |    |--*D. gradata Cope 1873 M60
    |    |    |--‘*Miothen’ crassigenus Cope 1873 M60
    |    |    |--‘Protosorex’ crassus M60
    |    |    |--‘Miothen’ gracile Cope 1873 M60
    |    |    `--D. thompsoni Simpson 1941 M58
    |    |--Paradomina Hutchison 1966 V67
    |    |--Ingentisorex Hutchison 1966 V67
    |    `--Trimylus Roger 1885 [incl. Heterosorex Gaillard 1915] V67
    `--+--Soricinae B78
       |    |--Soricini V67
       |    |    |--Sorex MB86
       |    |    |--Crocidosorex Lavocat 1951 [incl. Oligosorex Kretzoi 1959] V67
       |    |    |--Antesorex Repenning 1967 V67
       |    |    |--Drepanosorex Kretzoi 1941 V67
       |    |    |--Microsorex Baird 1877 V67
       |    |    |    `--M. hoyi (Baird 1858) B75 [=Sorex hoyi B75; incl. M. winnemana BP87]
       |    |    |--Alluvisorex Hutchison 1966 V67
       |    |    |--Blarinella Thomas 1911 V67
       |    |    |    |--B. quadraticauda IT07
       |    |    |    `--B. wardi IT07
       |    |    |--Petenyia Kormos 1934 V67
       |    |    `--Zelceina Sulimski 1962 V67
       |    |--Blarinini V67
       |    |    |  i. s.: ‘Sorex’ dehneli Kowalski 1956 V67
       |    |    |--Blarina RM05
       |    |    |--Adeloblarina Repenning 1967 V67
       |    |    |--Cryptotis V67
       |    |    |--Paracryptotis Hibbard 1950 V67
       |    |    |--Shikamainosorex Hasegawa 1957 V67
       |    |    |--Blarinoides Sulimski 1959 V67
       |    |    |    `--B. mariae Sulimski 1959 P04
       |    |    `--Peisorex Kowalski & Li 963 V67
       |    |--Neomyini [Amblycoptinae, Anourosoricinae, Crossopinae, Hydrosoridae, Nectogalinae, Soriculi] V67
       |    |    |--Neomys Kaup 1829 (see below for synonymy) V67
       |    |    |    |--N. anomalus OC03
       |    |    |    |--N. fodiens (Pennant 1771) FK55
       |    |    |    |    |--N. f. fodiens CM07
       |    |    |    |    `--N. f. bicolor CM07
       |    |    |    `--N. schelkovnikovi IT07
       |    |    |--Nectogale Milne-Edwards 1870 V67
       |    |    |    `--N. elegans IT07
       |    |    |--Anourosorex Milne-Edwards 1870 [=Anurosorex Anderson 1875; incl. Pygmura Anderson 1875] V67
       |    |    |    `--A. squamipes M58
       |    |    |--Soriculus V67
       |    |    |--Petenyiella Kretzoi 1956 [incl. Allopachyura Kormos 1934] V67
       |    |    |--Episoriculus Ellerman & Morrison-Scott 1951 V67
       |    |    |--Asoriculus Kretzoi 1959 V67
       |    |    |    `--A. gibberodon (Petényi 1864) (see below for synonymy) P04
       |    |    |--Chodsigoa Kashchenko 1907 V67
       |    |    |--Beremendia Kormos 1934 V67
       |    |    |--Chimarrogale Anderson 1877 [incl. Crossogale Thomas 1921] V67
       |    |    |    |--C. hantu IT07
       |    |    |    |--C. himalayica IT07
       |    |    |    |--C. leander M58
       |    |    |    |--C. phaeura IT07
       |    |    |    |--C. platycephala IT07
       |    |    |    |--C. styani IT07
       |    |    |    `--C. sumatrana IT07
       |    |    |--Amblycoptus Kormos 1926 V67
       |    |    |--Hesperosorex Hibbard 1957 V67
       |    |    |--Notiosorex Baird 1877 V67
       |    |    |    |--N. crawfordi (Coues 1877) MB86
       |    |    |    |    |--N. c. crawfordi MB86
       |    |    |    |    `--N. c. evotis (Coues 1877) MB86
       |    |    |    |--N. gigas G69
       |    |    |    `--N. (Xenosorex) phillipsii Schaldach 1966 G69
       |    |    |--Deinsdorfia Heller 1963 V67
       |    |    |    `--D. kordosi Reumer 1984 P04
       |    |    `--Megasorex Hibbard 1950 V67
       |    |         `--M. gigas IT07
       |    `--Allosoricinae V67
       |         |  i. s.: ‘Sorex’ gracilidens Viret & Zapfe 1951 V67
       |         `--Allosorex Fejfar 1966 V67
       `--+--Myosorex MJ11
          `--Crocidurinae B78

Asoriculus gibberodon (Pétenyi 1864) [=Crocidura gibberodon, Episoriculus gibberodon, Soriculus gibberodon] P04

Neomys Kaup 1829 [incl. Amphisorex Duvernoy 1835, Crossopus Wagler 1832, Hydrogale Kaup 1829, Hydrosorex Duvernoy 1835, Leucorrhynchus Kaup 1829, Myosictis Pomel 1854, Pinalia Gray 1838] V67

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[B78] Butler, P. M. 1978. Insectivora and Chiroptera. In: Maglio, V. J. & H. B. S. Cooke (eds) Evolution of African Mammals pp. 56–68. Harvard University Press: Cambridge (Massachusetts).

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals pp. 117–141. Clarendon Press: Oxford.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[FK55] Felten, H., & C. König. 1955. Einige Säugetiere aus dem Zentralmassiv, Südfrankreich. Senckenbergiana Biologica 36: 267–269.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[M58] McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 113–214.

[M60] McKenna, M. C. 1960. The Geolabidinae: a new subfamily of early Cenozoic erinaceoid insectivores. University of California Publications in Geological Sciences 37 (2): 131–164.

[MJ11] Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer & W. J. Murphy. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334: 521–524.

[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.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403–491.

[RM05] Reilly, S. M., R. W. Manning, C. C. Nice & M. R. J. Forstner. 2005. Systematics of isolated populations of short-tailed shrews (Soricidae: Blarina) in Texas. Journal of Mammalogy 86 (5): 887–894.

Rose, K. D. 2006. The Beginning of the Age of Mammals. JHU Press.

[TH03] Tedford, R. H., & C. R. Harington. 2003. An Arctic mammal fauna from the Early Pliocene of North America. Nature 425: 388–390.

[V67] Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135 (5): 217–284.

Elephantidae

African forest elephant Loxodonta africana cyclotis, from here.

Belongs within: Elephantida.
Contains: Elephas.

The Elephantidae, elephants, first appeared in the late Pliocene and are distinguished from earlier proboscideans by their distinctive 'conveyor-belt' mode of tooth development. The greatly enlarged adult molars develop sequentially rather than all at once, so that only one or two molars are functional at any one time. As each functioning molar becomes worn down through use, its developing successor moves forward to take its place. In the most basal elephantids, the lower premolars were also present, and the molars retained distinct crowns; in the Elephantinae, the premolars became non-functional and the molar crowns became pressed together into a series of laminae. Except for Stegotetrabelodon and Primelephas, elephantids also lost the lower tusks of earlier proboscideans (Coppens et al. 1978).

Living elephants belong to two genera, Elephas and Loxodonta. The African elephant Loxodonta africana is distinguished from Elephas by its concave back and trunk with two terminal 'fingers'.

Synapomorphies (from Tassy & Shoshani 1988): basicranium elevated; loss of enamel band on upper tusks; molars with five or more lophids, becoming laminae.

<==Elephantidae [Stegotetrabelodontinae]
    |--Stegotetrabelodon Petrocchi 1941 TS88, CM78
    |    |--*S. syrticus Petrocchi 1941 [incl. S. lybicus Petrocchi 1941, S. sahabianus Petrocchi 1943] CM78
    |    `--S. orbus Maglio 1970 CM78
    `--+--Stegodibelodon Coppens 1972 TS88, CM78
       |    `--*S. schneideri Coppens 1972 CM78
       `--Elephantinae CM78
            |--Primelephas Maglio 1970 CM78
            |    |--*P. gomphotheroides Maglio 1970 CM78
            |    `--P. korotorensis (Coppens 1965) [=Stegodon korotorensis] CM78
            `--+--+--Elephas FS15
               |  `--Loxodonta Cuvier 1825 FS15, CM78
               |       |--*L. africana (Blumenbach 1797) (see below for synonymy) CM78
               |       |    |--L. a. africana B84
               |       |    `--L. a. cyclotis B84
               |       |--L. adaurora Maglio 1970 CM78
               |       `--L. atlantica (Pomel 1879) [=Elephas atlanticus; incl. E. pomeli Arambourg 1952] CM78
               |            |--L. a. atlantica CM78
               |            `--L. a. zulu (Scott 1907) [=Elephas (Loxodon) zulu] CM78
               `--Mammuthus Burnett 1830 FS15, AC98 (see below for synonymy)
                    |  i. s.: ‘Archidiskodon’ gromovi D07
                    |         M. jeffersoni D07
                    |         M. lamarmorae D07
                    |--M. subplanifrons (Osborn 1928) (see below for synonymy) CM78
                    `--+--M. africanavus (Arambourg 1952) [=Elephas africanavus, Loxodonta africanava] CM78
                       `--+--M. meridionalis (Nesti 1825) (see below for synonymy) CM78
                          |--+--*M. primigenius (Blumenbach 1799) CM78 (see below for synonymy)
                          |  `--M. armeniacus [=Parelephas armeniacus; incl. M. trogontherii] CM78
                          `--+--M. columbi CM78 [=Elephas primigenius columbi FP64, C77, Parelephas columbi CM78]
                             |--M. exilis Stock & Furlong 1928 FS15, AC98
                             `--M. imperator CM78 [=Archidiskodon imperator CM78, Elephas imperator FP64]

*Loxodonta africana (Blumenbach 1797) [=Elephas africanus; incl. L. atlantica angammensis Coppens 1965, L. prima Dart 1929] CM78

Mammuthus Burnett 1830 TS88, AC98 [incl. Archidiskodon Pohlig 1888 AC98, Cheirolites von Meyer 1848 AC98, Dicyclotherium Géoffroy-Saint-Hilaire 1837 AC98, Mammonteus Camper in Osborn 1924 AC98, Metarchidiskodon Osborn 1934 AC98, Paraelephas Osborn 1924 AC98]

Mammuthus meridionalis (Nesti 1825) [=Elephas meridionalis, Archidiskodon meridionalis; incl. E. moghrebiensis Arambourg 1970] CM78

*Mammuthus primigenius (Blumenbach 1799) CM78 [=Elephas primigenius FP64, Mammonteus primigenius CM78]

Mammuthus subplanifrons (Osborn 1928) [=Archidiskodon subplanifrons; incl. A. andrewsi Dart 1929, A. planifrons nyanzae MacInnes 1942, Mammuthus (Archidiskodon) scotti Meiring 1955] CM78

*Type species of generic name indicated

REFERENCES

[AC98] Alcover, J. A., X. Campillo, M. Macias & A. Sans. 1998. Mammal species of the world: additional data on insular mammals. American Museum Novitates 3248: 1–29.

[B84] Barnes, R. F. W. 1984. Elephants. In: Macdonald, D. (ed.) All the World’s Animals: Hoofed Mammals pp. 12–21. Torstar Books: New York.

[C76] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[CM78] Coppens, Y., V. J. Maglio, C. T. Madden & M. Beden. 1978. Proboscidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 336–367. Harvard University Press: Cambridge (Massachusetts).

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

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[TS88] Tassy, P., & J. Shoshani. 1988. The Tethytheria: elephants and their relatives. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 283–315. Clarendon Press: Oxford.

Last updated: 15 August 2018.

Elephantida

Life-size reconstructions of Stegodon, copyright Vjdchauhan.

Belongs within: Proboscidea.
Contains: Elephantidae.

The proboscidean clade Elephantida unites the paraphyletic 'gomphotheres' with the living Elephantidae: essentially, all proboscideans closer to modern elephants than the mastodons (Mammutidae). As recognised here, the Elephantida are first recorded in the Miocene of Africa, subsequently spreading to Eurasia and the Americas. Subgroups include the Amebelodontidae of the Miocene and Pliocene which had a distinctive elongate lower jaw ending in shovel-shaped tusks.

Synapomorphies (from Tassy & Shoshani 1988): Lacrimal foramen generally not opened in the orbit at maturity; palatine with reduced spina nasalis posterior; external nasal opening enlarged laterally; enlarged tritolophid on M_1-2; cement on crowns of teeth; deep bicipital groove on humerus and higher trochanter; carpus and metacarpus less cursorial; lunar with enlarged articular surface for ulna.

<==Elephantida [Amebelodontinae, Gomphotheriidae, Gomphotheriinae, Gomphotherioidea]
    |--Choerolophodon Schlessinger 1917 TS88, CM78
    |    `--C. palaeindicus SKR04
    |--Amebelodontidae TS88
    |    |--Archaeobelodon TS88
    |    |--Serbelodon TS88
    |    |--Gnathabelodon TS88
    |    |--Amebelodon Barbour 1927 D07
    |    |    |--A. britti D07
    |    |    `--A. floridanus D07
    |    `--Platybelodon Borissiak 1928 TS88, CM78
    |         |--*P. danovi CM78
    |         |--P. filholi CM78
    |         `--P. grangeri CM78
    `--+--+--Stegomastodon TS88
       |  |    |--S. platensis FS15
       |  |    `--S. waringi FS15
       |  `--Cuvieronius Osborn 1923 FS15, D07 [Cuvieroninae]
       |       |--C. hyodon D07
       |       `--C. tropicus D07
       `--+--Elephantoidea SW06
          |    |--Elephantidae SW06
          |    `--Stegodontidae SW06
          |         |--Stegolophodon Petrocchi 1943 TS88, CM78
          |         `--Stegodon TS88
          |              |  i. s.: S. ganesa KS91
          |              |         S. insignis KS91
          |              |         S. kaisensis CM78
          |              |--S. florensis FS15
          |              `--+--S. orientalis FS15
          |                 `--S. trigonocephalus FS15
          `--+--Tetralophodon Falconer & Cautley 1857 TS88, CM78
             |    |--T. exoletus DW04
             |    |--T. longirostris CM78
             |    `--‘Mastodon’ mirificus C77
             `--Anancus Aymard 1855 [incl. Pentalophodon Falconer & MacInnes 1942; Anancinae] CM78
                  |--*A. arvernensis [=Mastodon arvernensis] CM78
                  |--A. jebtebsus D07
                  |--A. kenyensis (MacInnes 1942) [=Pentalophodon sivalensis kenyensis] CM78
                  |--A. osiris Arambourg 1945 CM78
                  `--A. petrocchii Coppens 1965 [=A. (Pentalophodon) petrocchii] CM78

Elephantida incertae sedis:
  Afrochoerodon kisumuensis SKR04
  Paratetralophodon TS88
  Rhynchotherium TS88
  Eubelodon TS88
  Haplomastodon TS88
  Notiomastodon TS88
  Gomphotherium Burmeister 1837 TS88, CM78 (see below for synonymy)
    |--*G. angustidens (Cuvier 1806) (see below for synonymy) CM78
    |    |--G. a. angustidens CM78
    |    `--G. a. pontileviensis CM78
    |--G. ngorora Maglio 1974 CM78
    |--‘Moeritherium’ pandionis Pilgrim 1912 [=Trilophodon pandionis] CB78
    |--G. productum CM78
    `--G. pygmaeus (Depéret 1897) (see below for synonymy) CM78

Gomphotherium Burmeister 1837 TS88, CM78 [incl. Bunolophodon Vacek 1877 CM78, Serridentinus Osborn 1923 CM78, Trilophodon Falconer & Cautley 1857 CM78]

*Gomphotherium angustidens (Cuvier 1806) [=Mastodon angustidens; incl. Trilophodon angustidens kisumuensis MacInnes 1942, M. angustidens var. libyca Fourtau 1918, Gomphotherium angustidens libycus, Protanancus macinnesi Arambourg 1945, Mastodon spenceri Fourtau 1918] CM78

Gomphotherium pygmaeus (Depéret 1897) [=Mastodon angustidens pygmaeus, Phiomia pygmaeus, Trilophodon olisponensis pygmaeus] CM78

*Type species of generic name indicated

REFERENCES

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[CB78] Coppens, Y., & M. Beden 1978. Moeritherioidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 333–335. Harvard University Press: Cambridge (Massachusetts).

[CM78] Coppens, Y., V. J. Maglio, C. T. Madden & M. Beden. 1978. Proboscidea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 336–367. Harvard University Press: Cambridge (Massachusetts).

[DW04] Deng T., Wang X., Ni X. & Liu L. 2004. Sequence of the Cenozoic mammalian faunas of the Linxia Basin in Gansu, China. Acta Geologica Sinica (English Edition) 78 (1): 8–14.

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

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[KS91] Kennedy, K. A. R., A. Sonakia, J. Chiment & K. K. Verma. 1991. Is the Narmada hominid an Indian Homo erectus? American Journal of Physical Anthropology 86: 475–496.

[SKR04] Sanders, W. J., J. Kappelman & D. T. Rasmussen. 2004. New large-bodied mammals from the late Oligocene site of Chilga, Ethiopia. Acta Palaeontologica Polonica 49 (3): 365–392.

[SW06] Shoshani, J., R. C. Walter, M. Abraha, S. Berhe, P. Tassy, W. J. Sanders, G. H. Marchant, Y. Libsekal, T. Ghirmai & D. Zinner. 2006. A proboscidean from the late Oligocene of Eritrea, a “missing link” between early Elephantiformes and Elephantimorpha, and biogeographical implications. Proceedings of the National Academy of Sciences of the USA 103 (46): 17296–17301.

[TS88] Tassy, P., & J. Shoshani. 1988. The Tethytheria: elephants and their relatives. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 283–315. Clarendon Press: Oxford.

Last updated: 15 August 2018.

Dicynodontia

Mounted skeleton of Lystrosaurus murrayi in the Muséum national d'histoire naturelle, Paris, photographed by FunkMonk.

Belongs within: Eutherapsida.
Contains: Kannemeyeriiformes.

The Dicynodontia are a group of herbivorous synapsids known from the the Late Permian to the Cretaceous, though they did not survive past the Late Triassic outside Australia. At their peak, they were the world's dominant herbivores. One genus, Lystrosaurus, included the most abundant terrestrial vertebrates in the earliest Triassic immediately following the end-Permian mass extinction. The name 'dicynodont' refers to the pair of large tusks that were generally the only teeth in their beaked jaws.

Synapomorphies (from Kemp 1988): Zygomatic arch dorsally displaced; dentaries fused at symphysis; articular surface of lower jaw sloping posteroventrally; horn developed on the jaws; intramandibular fenestra present; premaxillary secondary palate present.

<==Dicynodontia
    |--Kombuisia M00
    |--Myosaurus M00
    `--Pristerodontia M00
         |  i. s.: Odontocyclops M00
         |--Lystrosaurus Cope 1870 M00, D07 [Lystrosauridae]
         |    |--L. curvatus D07
         |    |--L. declivis D07
         |    |--L. georgi D07
         |    |--L. hedini D07
         |    |--L. mccaigi D07
         |    |--L. murrayi M00
         |    |--L. oviceps D07
         |    |--L. platycopes D07
         |    |--L. rajurkari D07
         |    |--L. robustus D07
         |    |--L. shichanggouensis D07
         |    `--L. weidenrichi D07
         `--+--+--Dinanomodon M00
            |  `--Kannemeyeriiformes M00
            `--Dicynodon Owen 1845 M00, D07 [Dicynodontidae]
                 |--D. bolorhinus D07
                 |--D. grimbeeki D07
                 |--D. lacerticeps M00
                 |--D. leontops D07
                 |--D. lissops D07
                 |--D. osborni D07
                 |--D. plateceps D07
                 |--D. trautscholdi D07
                 |--D. trigonocephalus D07
                 |--D. turpior D07
                 `--D. whaitsi D07

Dicynodontia incertae sedis:
  Cistecephalus Owen 1876 D07
    `--C. microrhinus D07
  Emydoses M00
  Otsheria K88
  Venyukovioidea TT05
    |  i. s.: Suminia getmanovi Ivakhnenko 1994 TT05
    `--Venyukoviidae TT05
         |--Ulemica efremovi Ivakhnenko 1996 TT05
         `--Venjukovia K88
  Aulacocephalodon peavoti N85
  Diictodon Owen 1876 D07
    `--E. feliceps D07

*Type species of generic name indicated

REFERENCES

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

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

[M00] Maisch, M. W. 2000. Observations on Karoo and Gondwana vertebrates. Part 2: A new skull-reconstruction of Stahleckeria potens von Huene, 1935 (Dicynodontia, Middle Triassic) and a reconstruction of kannemeyeriiform phylogeny. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 220 (1): 127–152.

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

[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.

Last updated: 24 August 2018.

Elephas

Asian elephant Elephas maximus photographed by Manoj Sindagi.

Belongs within: Elephantidae.

Though represented in the modern fauna only by the Asian elephant Elephas maximus, the genus Elephas is first recorded in the middle Pliocene of eastern and southern Africa, spreading from there into the Middle East by the later Pliocene (Coppens et al. 1978). The genus remained widespread in the Old World until it became extinct in Africa and Europe in the late Pleistocene.

Characters (from Coppens et al. 1978): Skull high, forehead fore-shortened. Premaxillary tusk sockets generally widely separated, tusks usually gently curved in a single plane. Frontoparietal surface flat to concave; upper borders of the temporal fossae forming sharp, raised ridges. Mandible corpus strongly convex laterally, condyles transversely elongated and directed upward and inward.

<==Elephas Linnaeus 1758 [incl. Omoloxodon Deraniyagala 1955, Palaeoloxodon Matsumoto 1929, Pilgrimia Osborn 1924] CM78
    |--E. ekorensis Maglio 1970 CM78
    |--+--E. iolensis Pomel 1895 (see below for synonymy) CM78
    |  |--E. recki Dietrich 1916 (see below for synonymy) CM78
    |  `--+--E. falconeri CM78
    |     `--E. namadicus CM78 [=Paleoloxodon namadicus DW04]
    `--+--+--E. celebensis CM78
       |  `--+--E. planifrons [=Archidiskodon planifrons] CM78
       |     `--E. platycephalus CM78
       `--+--E. hysudricus CM78
          `--+--*E. maximus Linnaeus 1758 CM78
             |    |--E. m. maximus B84
             |    |--E. m. bengalensis B84
             |    |--E. m. hirsutus B84
             |    `--E. m. sumatrensis B84
             `--E. hysudrindicus CM78

Elephas iolensis Pomel 1895 [incl. Pilgrimia archidiskodontoides Haughton 1932, Archidiskodon broomi Osborn 1928, A. hanekomi Dart 1929, Palaeoloxodon kuhni Dart 1929, A. sheppardi Dart 1927, Pilgrimia subantiqua Haughton 1932, A. transvaalensis Dart 1927, Pi. wilmani Dart 1920, Pi. yorki Dart 1929] CM78

Elephas recki Dietrich 1916 [=E. (Archidiskodon) recki, Loxodonta (Pilgrimia) antiqua recki, *Omoloxodon recki; incl. Archidiskodon exoptatus Dietrich 1942, E. exoptatus] CM78

*Type species of generic name indicated

==References==

[B84] Barnes, R. F. W. 1984. Elephants. In All the World’s Animals: Hoofed Mammals (D. Macdonald, ed.) pp. 12-21. Torstar Books: New York.

[CM78] Coppens, Y., V. J. Maglio, C. T. Madden & M. Beden. 1978. Proboscidea. In Evolution of African Mammals (V. J. Maglio & H. B. S. Cooke, eds) pp. 336-367. Harvard University Press: Cambridge (Massachusetts).

[DW04] Deng T., Wang X., Ni X. & Liu L. 2004. Sequence of the Cenozoic mammalian faunas of the Linxia Basin in Gansu, China. Acta Geologica Sinica (English Edition) 78 (1): 8-14.

Andreaeopsida

Andreaea rupestris, photographed by Michael Lüth.

Belongs within: Bryophyta.

As recognised here, the Andreaeopsida comprise a small assemblage of basal mosses in which the spores are released from the capsule via dehiscence through longitudinal slits, rather than via an operculum as in other mosses (Smith & Davison 1993). The monophyly of the Andreaeopsida is uncertain, with many authors regarding its members as a paraphyletic grade dividing the basal Sphagnopsida from the remaining mosses. The position of Takakia, a distinctive moss genus found in eastern Asia and north-west North America, has been particularly contentious, but it has been linked to Andreaeobryum by features such as the presence of secondary protonemata and of massed beaked mucilage-producing papillae. Takakia is otherwise distinguished by erect shoots arising from a stolon, terete leaves with oil droplets and slime hairs, and naked lateral gametangia. Species of Andreaeaceae are epilithic: Andreaea typically grows on granite or other acidic rocks while Andreaeobryum macrosporum grows on limestone (Flora of North America).

<==Andreaeopsida
    |--Takakia [Takakiaceae, Takakiales, Takakiidae] FHH01
    |    `--T. ceratophylla FHH01
    `--Andreaeaceae [Andreaeales, Andreaeidae] FHH01
         |--Andreaeobryum macrosporum FHH01
         |--Acroschisma wilsoni D03
         `--Andreaea Hedw. 1801 SK02
              |  i. s.: A. acutifolia Hooker & Wilson 1844 SK02
              |           |--A. a. ssp. acutifolia SK02
              |           `--A. a. ssp. acuminata (Mitt.) Vitt 1980 SK02 [=A. petrophila var. acuminata D24]
              |         A. amblyophylla Müll.Hal. ex Broth. 1895 SK02
              |         A. appendiculata D03
              |         A. asperula Mitt. 1860 [incl. A. muelleri (nom. inv.)] SK02
              |         A. australis Mitt. 1856 SK02
              |         A. erubescens Müll.Hal. 1898 SK02
              |           |--A. e. var. erubescens SK02
              |           `--A. e. var. nigrita Müll.Hal. 1898 SK02
              |         A. eximia Müll.Hal. 1898 SK02
              |         A. flabellata Müll.Hal. 1883 SK02
              |         A. flexuosa R.Br.bis 1893 SK02
              |         A. gainii Cardot 1911 SK02
              |         A. heinemannii Hampe & Müll.Hal. 1846 SK02
              |         A. julicaulis Müll.Hal. 1898 SK02
              |         A. laxifolia D03
              |         A. marginata D03
              |         A. microvaginata Müll.Hal. 1898 SK02
              |         A. montana Mitt. 1859 SK02
              |         A. mutabilis Hooker & Wilson 1844 SK02 (see below for synonymy)
              |         A. nitida Hooker & Wilson 1844 SK02
              |         A. pseudo-alpina D03
              |         A. subulata Harv. ex Hooker 1840 (see below for synonymy) SK02
              |           |--A. s. var. subulata D03
              |           `--A. s. var. perichaetialis D03
              |         A. tasmanica Rodway 1916 SK02
              |         A. tenera Müll.Hal. 1898 SK02
              |         A. thedenii H04
              |--A. (subg. Chasmocalyx) nivalis D24
              |    |--A. n. var. nivalis D24
              |    `--A. n. var. fuscescens D24
              `--A. subg. Eu-Andraea D24
                   |--A. alpina Hedw. 1801 D24, SK02 [=Jungermannia alpina D24]
                   |    |--A. a. var. alpina D24
                   |    `--A. a. var. flavicans D24
                   |--A. crassinervia D24
                   |    |--A. c. var. crassinervia D24
                   |    `--A. c. var. huntii D24
                   |--A. obovata D24
                   |--A. rothii D24
                   |    |--A. r. var. rothii D24
                   |    |--A. r. var. falcata D24
                   |    |--A. r. var. grimsulana D24
                   |    `--A. r. var. hamata D24
                   `--A. rupestris Hedw. 1801 [incl. A. petrophila] SK02
                        |--A. r. var. rupestris D24
                        |--A. ‘petrophila’ var. alpestris D24
                        |--A. ‘petrophila’ var. flaccida D24
                        |--A. ‘petrophila’ var. gracilis D24
                        |--A. ‘petrophila’ var. sparsifolia D24
                        `--A. ‘petrophila’ var. sylvicola D24

Nomina nuda: Andreaea loricata Dusén 1903 D03
             Andreaea pulvinatula Dusén 1903 D03
             Andreaea purpurascens Dusén 1903 D03
             Andreaea vermicularis Dusén 1903 D03

Andreaea mutabilis Hooker & Wilson 1844 SK02 [incl. A. amblyophylla var. attenuata SK02, A. amblyophylla var. bullata SK02, A. petrophila var. homomalla D24, SK02]

Andreaea subulata Harv. ex Hooker 1840 [incl. A. pseudosubulata, A. subulata var. rigida, A. subulatissima] SK02

*Type species of generic name indicated

REFERENCES

[D24] Dixon, H. N. 1924. The Student's Handbook of British Mosses 3^rd ed. V. V. Sumfield: Eastbourne.

[D03] Dusén, P. 1903. Patagonian and Fuegian mosses. In: Scott, W. B. (ed.) Reports of the Princeton University Expeditions to Patagonia, 1896–1899 vol. 8. Botany pp. 63–126. The University: Princeton (New Jersey).

[FHH01] Frey, W., M. Hofmann & H. H. Hilger. 2001. The gametophyte-sporophyte junction: unequivocal hints for two evolutionary lines of archegoniate land plants. Flora 196: 431–445.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

Smith, D. K., & P. G. Davison. 1993. Antheridia and sporophytes in Takakia ceratophylla (Mitt.) Grolle: evidence for reclassification among the mosses. Journal of the Hattori Botanical Laboratory 73: 263-271.

[SK02] Streimann, H., & N. Klazenga. 2002. Catalogue of Australian Mosses. Flora of Australia Supplementary Series 17. Australian Biological Resources Study: Canberra.

Last updated: 11 January 2018.

Mystacocarida

Drawing of Derocheilocaris typica, from Michigan Science Art.

Belongs within: Pancrustacea.

The Mystacocarida are a group of minute (~0.45 to 1.01 mm), elongate, cylindrical crustaceans with morphologically relatively unspecialised limbs (Zinn 1986). All known mystacocarids are meiobenthonic, living in the interstitial spaces between sand grains in marine intertidal and subtidal areas. The relationships of mystacocarids to other crustaceans are contentious; they possess a number of apparently primitive features that may be truly so, or may represent neotenic features retained from the naupliar larval stage (Zinn 1986).

<==Mystacocarida
    |--Ctenocheilocaris Renaud-Mornant 1976 Z86
    |    |--C. claudiae Renaud-Mornant 1976 Z86
    |    `--C. galvarini (Dahl 1952) [=Derocheilocaris galvarini] Z86
    `--Derocheilocaris Pennak & Zinn 1943 [Derocheilocarididae] Z86
         |--D. algoensis McLachlan & Grindley 1974 Z86
         |--D. angolensis Hessler 1972 Z86
         |--D. delamarei Hessler 1972 Z86
         |--D. ingens Hessler 1969 Z86
         |--D. remanei Delamare Deboutteville & Chappuis 1951 Z86
         |    |--D. r. remanei Z86
         |    |--D. r. akhzivi Masry & Por 1970 Z86
         |    |--D. r. biscayensis Delamare Deboutteville 1953 Z86
         |    `--D. r. katesi Noodt 1954 Z86
         |--D. tehiyae Masry & Por 1970 Z86
         `--D. typica Pennak & Zinn 1943 Z86

REFERENCES

[Z86] Zinn, D. J. 1986. Mystacocarida. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 385-388. E. J. Brill / Dr. W. Backhuys: Leiden.