Thelyphoninae

Thelyphonus sp., copyright Arthur Anker.

Belongs within: Uropygi.

The Thelyphoninae is a group of whipscorpions found in south-east Asia and Melanesia (Rowland & Cooke 1973).

Characters (from Rowland & Cooke 1973): Tarsal segments of first leg of female usually modified (except in Minbosius manilanus and Abaliella rohdei), variously incrassate or apophysate; pedipalpal patellar apophysis of male unmodified, occasionally longer than in female; abdominal sternites II and III modified; two, four or no ommatoids present; keel present between lateral and median eyes; posterior abdominal tergites divided.

<==Thelyphoninae [Thelyphonini]
    |--Minbosius Speijer 1936 RC73
    |    `--*M. manilanus (Koch 1843) RC73
    |--Chajnus Speijer 1936 RC73
    |    `--*C. renschi Speijer 1936 RC73
    |--Glyptogluteus Rowland 1973 RC73
    |    `--*G. augustus Rowland 1973 RC73
    |--Ginosigma Speijer 1936 RC73
    |    |--*G. schimkewitschi (Tarnani 1894) RC73
    |    `--G. lombokensis Speijer 1936 RC73
    |--Tetrabalius Thorell 1888 RC73, H02
    |    |--*T. seticauda (Doleschall 1857) RC73
    |    `--T. nasutus Thorell 1889 RC73
    |--Abaliella Strand 1928 [=Abalius Kraepelin 1897 (preoc.)] RC73
    |    |--*A. rohdei (Kraepelin 1897) RC73
    |    |--A. dicranotarsalis Rowland 1973 RC73
    |    |--A. gertschi Rowland 1973 RC73
    |    |--A. manilana (Kraepelin 1901) RC73
    |    |--A. samoana (Kraepelin 1879) RC73
    |    `--A. willeyi (Pocock 1898) RC73
    `--Thelyphonus Latreille 1802 H02 [=Telyphonus (l. c.) KK84]
         |--*T. caudatus (Linnaeus 1758) RC73 [=Phalangium caudatum H02; incl. T. thorellii Tarnani 1890 T94]
         |--T. anthracinus Pocock 1894 RC73
         |--T. antillanus T94
         |--T. asperatus Thorell 1888 RC73
         |--T. assamensis Stoliczka 1869 [=Telyphonus assamensis] S69
         |--T. billitonensis Speijer 1931 RC73
         |--T. borneensis Kraepelin 1897 RC73
         |--T. burchardi Kraepelin 1912 RC73
         |--T. celebensis Kraepelin 1897 RC73
         |--T. doriae Thorell 1888 RC73
         |--T. grandis Speijer 1931 RC73
         |--T. hanseni Kraepelin 1897 RC73
         |--T. ignotus T94
         |--T. insulanus Koch & Keyserling 1884 RC73, KK84
         |--T. javanus T94
         |--T. kinabaluensis Speijer 1933 RC73
         |--T. klugi Kraepelin 1897 RC73
         |--T. lawrencei Rowland 1973 RC73
         |--T. leucurus Pocock 1898 RC73
         |--T. linganus Koch 1843 RC73
         |--T. pococki Tarnani 1901 RC73
         |--T. proscorpius T94
         |--T. rufipes [=Telyphonus rufipes] D57
         |--T. schnehageni Kraepelin 1897 RC73
         |--T. semperi Kraepelin 1897 RC73
         |--T. sepiaris Butler 1873 RC73 [incl. T. nigrescens Butler 1873 T89]
         |--T. seticauda Doleschall 1857 [=Telyphonus seticauda] D57
         |--T. spinimanus [=Telyphonus spinimanus] S69
         |--T. stimpsonii Wood 1861 [=Telyphonus stimpsonii] S69
         |--T. sucki Kraepelin 1897 RC73
         |--T. sumatranus Kraepelin 1897 RC73
         |--T. vanoorti Speijer 1936 RC73
         `--T. wayi Pocock 1900 RC73

*Type species of generic name indicated

REFERENCES

[D57] Doleschall, C. L. 1857. Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië, series 3, 3 (5–6): 399–434, plates 1–2.

[H02] Harvey, M. S. 2002. The neglected cousins: What do we know about the smaller arachnid orders? Journal of Arachnology 30 (2): 357–372.

[KK84] Koch, L., & E. Keyserling. 1884-1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

[RC73] Rowland, J. M., & J. A. L. Cooke. 1973. Systematics of the arachnid order Uropygida (=Thelyphonida). Journal of Arachnology 1: 55–71.

[S69] Stoliczka, F. 1869. Contribution to the knowledge of Indian Arachnoidea. Journal of the Asiatic Society of Bengal 38 pt 2 (4): 201–251, plates 18–20.

[T89] Thorell, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XXI.—Aracnidi Artrogastri Birmani raccolti da L. Fea nel 1885–1887. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 521–729.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt. 4 (4): 1–63.

Conciliterga

Reconstruction of Kuamaia lata, from Hou and Bergström (1997).

Belongs within: Trilobitomorpha.

The Conciliterga are a group of soft-bodied arthropods related to trilobites known from the Lower and Middle Cambrian Burgess Shale and Chengjiang faunas.

Synapomorphies (from Cotton & Braddy 2004, not including Australimicola): Dorsal bulge present in exoskeleton accomodating drop-shaped ventral eyes; anteromedian margin of cephalon notched, accomodating strongly sclerotised plate; distal lobe of exopod large, teardrop-shaped, with long attachment to proximal lobe; joints between posterior tergites functional, anterior ones variably fused. (Running data matrix from Legg et al. (2013) through TNT identifies only one synapomorphy for Conciliterga including Australimicola: mediolateral spines present.)

Conciliterga [Helmetiida]
    |--Australimicola LSE13
    `--+--+--Skioldia LSE13
       |  `--+--Saperion glumaceum Hou et al. 1991 LSE13, CB04
       |     `--Tegopelte Simonetta & Delle Cave 1975 LSE13, CB04
       |          `--T. gigas Simonetta & Delle Cave 1975 CB04
       `--+--Nathorstia Walcott 1912 B95
          `--Helmetiidae CB04
               |--Helmetia Walcott 1918 CB04
               |    `--H. expansa Walcott 1918 CB04
               |--Rhombicalvaria Hou 1987 CB04, B95
               |    `--R. acantha Hou 1987 CB04
               `--Kuamaia Hou 1987 CB04, B95
                    |--K. lata Hou 1987 CB04
                    `--K. muricata Hou & Bergström 1997 CB04

*Type species of generic name indicated

REFERENCES

[B95] Bousfield, E. L. 1995. A contribution to the natural classification of Lower and Middle Cambrian arthropods: Food-gathering and feeding mechanisms. Amphipacifica 2: 3–34.

[CB04] Cotton, T. J., & S. J. Braddy. 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh: Earth Sciences 94: 169–193.

[LSE13] Legg, D. A., M. D. Sutton & G. D. Edgecombe. 2013. Arthropod fossil data increase congruence of morphological and molecular phylogenies. Nature Communications 4 (2485): 1–7.

Lejopyge

Lejopyge laevigata, from Angelin (1851–1878).

Belongs within: Agnostida.

Lejopyge is a genus of agnostidans known from the Middle and Upper Cambrian of Eurasia (Harrington et al. 1959).

Characters (from Harrington et al. 1959): Axial furrows of both shields obsolete except near base of glabella and at upper end of pygidial axis; genae smooth or rugose; surface of pygidium smooth.

<==Lejopyge Hawle & Corda 1847 HLC03
    |--*L. laevigata (Dalman 1828) [=Battus laevigatus] HLC03
    |--L. acantha Robison 1984 HLC03
    |--L. armata (Linnarsson 1869) (see below for synonymy) HLC03
    |--L. barrandei (Hicks 1872) HLC03
    |--L. calva Robison 1964 HLC03
    |--L. elegans (Tullberg 1880) HLC03
    |--L. hybridus (Brøgger 1878) HLC03
    |--L. multifora Öpik 1979 HLC03
    |--L. seminula (Whitehouse 1939) HLC03
    `--L. sinensis Lu & Lin in Peng 1987 HLC03

Lejopyge armata (Linnarsson 1869) [=Agnostus laevigatus var. armatus, L. laevigata armata, L. laevigatus armatus (l. c.); incl. L. laevigata perrugata Westergård 1946, L. armata trigonospinosa Yang 1982, L. zhejiangensis Qiu in Qiu, Lu et al. 1983] HLC03

*Type species of generic name indicated

REFERENCES

Harrington, H. J., G. Henningsmoen et al. 1959. Systematic descriptions. In Treatise on Invertebrate Paleontology pt O. Arthropoda 1 pp. O170–O540. Geological Society of America and University of Kansas Press.

[HLC03] Hong, P. S., J. G. Lee & D. K. Choi. 2003. Trilobites from the Lejopyge armata zone (upper Middle Cambrian) of the Machari Formation, Yongwol Group, Korea. Journal of Paleontology 77 (5): 895–907.

Lisaniidae

Cranidium of Eoshengia, from here.

Belongs within: Ptychopariida.

The Lisaniidae are a group of small trilobites known from the upper Middle Cambrian of Eurasia (Hong et al. 2003).

Characters (from Hong et al. 2003): Small, convex; glabella large, subquadrate; frontal area short, flat to convex.

<==Lisaniidae
    |--Lisania Walcott 1911 [incl. Aojia Kobayashi 1935] HLC03
    |--Redlichaspis Kobayashi 1935 [=Lisaniella Chang 1963] HLC03
    |--Shengia Xiang in Egorova et al. 1963 HLC03
    |--Platylisania Zhang & Jell 1987 HLC03
    `--Eoshengia Yang 1978 HLC03
         |--*E. subquadrata Yang 1978 HLC03
         |--E. constricta Zhang 1981 HLC03
         |--E. dongqinlingia Yan in Yang et al. 1991 HLC03
         |--E. gaoqiaogouensis Yang & Liu in Yang et al. 1991 HLC03
         |--E. jiudiantangensis Yang 1978 HLC03
         |--E. paragenalata Yang 1978 HLC03
         |--E. paratenuis Yang in Yang et al. 1991 HLC03
         |--E. parva Yang in Yang et al. 1991 HLC03
         |--E. quadrata Yang 1978 HLC03
         |--E. rigida Yan in Yang et al. 1991 HLC03
         |--E. shorteglabella Yang & Liu in Yang et al. 1991 HLC03
         |--E. spinosa Yang 1978 HLC03
         |--E. sudani Jell & Hughes 1997 HLC03
         |--E. tenuis Zhang 1981 HLC03
         |--E. xinjiangensis Zhang 1981 HLC03
         `--E. youshuiensis (Yang 1978) HLC03

*Type species of generic name indicated

REFERENCES

[HLC03] Hong, P. S., J. G. Lee & D. K. Choi. 2003. Trilobites from the Lejopyge armata zone (upper Middle Cambrian) of the Machari Formation, Yongwol Group, Korea. Journal of Paleontology 77 (5): 895–907.

Cyclolorenzella

Holaspid cranidia of Cyclolorenzella sp., from Hong et al. (2003).

Belongs within: Ptychopariida.

Cyclolorenzella is a genus of trilobites known from the upper Middle Cambrian of Asia. Members of this genus have a convex preglabellar boss and small palpebral lobes at glabellar mid-length (Hong et al. 2003).

<==Cyclolorenzella Kobayashi 1960 [Diceratocephalidae] HLC03
    |--*C. quadrata (Kobayashi 1935) [=Lorenzella quadrata] HLC03
    |--C. acalle (Walcott 1905) HLC03
    |--C. caijiapingensis Yang 1978 HLC03
    |--C. convexa (Resser & Endo 1937) HLC03
    |--C. hebeiensis Wittke & Zhu in Zhu & Wittke 1989 HLC03
    |    |--C. h. hebeiensis HLC03
    |    `--C. h. tangshanensis Wittke & Zhu in Zhu & Wittke 1989 HLC03
    |--C. humilis Zhang in Qiu et al. 1983 HLC03
    |--C. kushanensis (Chu 1959) HLC03
    |--C. latisulcata Zhang in Qiu et al. 1983 HLC03
    |--C. longispinosa Wittke & Zhu in Zhu & Wittke 1989 HLC03
    |--C. parabola (Lu 1957) HLC03
    |--C. paraconvexa Yang 1978 HLC03
    |--C. pustulosa (Chu 1959) HLC03
    |--C. regularis (Walcott 1906) HLC03
    |--C. subcylindrica (Chu 1959) HLC03
    |--C. tonkinenensis (Mansuy 1916) HLC03
    |--C. tuma Yang 1978 HLC03
    |--C. uniforma Wittke & Zhu in Zhu & Wittke 1989 HLC03
    `--C. yentaiensis (Chu 1959) HLC03

*Type species of generic name indicated

REFERENCES

[HLC03] Hong, P. S., J. G. Lee & D. K. Choi. 2003. Trilobites from the Lejopyge armata zone (upper Middle Cambrian) of the Machari Formation, Yongwol Group, Korea. Journal of Paleontology 77 (5): 895–907.

Phacopida

Eccoptochile mariana, from the American Museum of Natural History.

Belongs within: Ptychopariida.
Contains: Sphaerexochinae, Pilekiinae, Cheirurinae.

The Phacopida are a group of trilobites known from the Ordovician to the Devonian. Most members of this group are distinguished by the possession of proparian facial sutures, with the suture emerging in front of the genal angle. They are divided between the Phacopina, with a broadly anteriorly expanded glabella and schizochroal eyes, and the Cheirurina, with four pairs of glabellar furrows, small holochroal eyes and a spiny pygidium (Prothero 1998).

<==Phacopida
    |--Phacopina P98
    |    |--Greenops P98
    |    `--Phacops P98
    |         |--P. crosslei Etheridge & Mitchell 1895 F71
    |         |--P. latifrons H04
    |         |--P. latigenalis Etheridge & Mitchell 1895 F71
    |         |--P. longicaudatus M87
    |         |--P. macdonaldi Fletcher 1950 F71
    |         |--P. rana P98
    |         `--P. serratus F71
    `--Cheiruroidea [Cheirurina] P98
         |--Pliomeridae PVP85
         |    |  i. s.: Emsurella Rosova 1960 PVP85
         |    |           `--E. humilla Rosova 1960 PVP85
         |    |         Pseudocybele nasuta FO99, J79
         |    |         Pliomerellus Chugaeva 1973 PVP85
         |    `--Protopliomeropinae PVP85
         |         |--Protopliomerops Kobayashi 1934 PVP85
         |         `--Rossaspis Harrington 1957 PVP85
         |              `--R. superciliosa J79
         `--Cheiruridae PVP85
              |  i. s.: Krattaspis Öpik 1937 PVP85
              |--Sphaerexochinae PVP85
              |--+--Pilekiinae PVP85
              |  `--Cheirurinae PVP85
              `--Eccoptochilinae PVP85
                   |--Foulonia Přibyl & Vaněk in Přibyl, Vaněk & Pek 1985 PVP85
                   |    `--*F. peregrina (Dean 1966) [=Ceraurinella peregrinus] PVP85
                   |--Eccoptochiloides Prantl & Přibyl 1947 PVP85
                   |    |--E. scuticauda (Barrande 1846) [incl. *E. tumescens (Barrande 1852)] PVP85
                   |    |--E. henryi Přibyl, Vaněk & Pek 1985 PVP85
                   |    `--E. octacantha (Angelin 1854) PVP85
                   |--Placoparina Whittard 1940 PVP85
                   |    `--*P. sedgwickii (McCoy 1849) PVP85
                   |         |--P. s. sedgwickii PVP85
                   |         `--P. s. shelvensis Hughes 1969 PVP85
                   `--Eccoptochile Hawle & Corda 1847 PVP85
                        |--*E. clavigera (Beyrich 1845) PVP85
                        |--E. almadenensis Romano 1980 PVP85
                        |--E. impedita Hammann 1972 PVP85
                        |--E. mariana (Verneuil & Barrande 1855) PVP85
                        |--E. perlata Hawle & Corda 1847 PVP85
                        |--E. quillieri (Tromelin in Guillier 1873) PVP85
                        `--E. scrobiculata (Angelin 1854) PVP85

*Type species of generic name indicated

REFERENCES

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

[FO99] Fortey, R. A. & R. M. Owens. 1999. Feeding habits in trilobites. Palaeontology 42 (3): 429–465.

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

[J79] Jaanusson, V. 1979. Ordovician. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A136–A166. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[M87] Mitchell, J. 1887. Notes on the geology of Bowning, N.S.W. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1193–1204.

[PVP85] Přibyl, A., J. Vaněk & I. Pek. 1985. Phylogeny and taxonomy of family Cheiruridae (Trilobita). Acta Universitatis Palackianae Olomucensis Facultas Rerum Naturalium Geographica-Geologica XXIV 83: 107–193.

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

Olenacea

Olenus gibbosus, from here.

Belongs within: Ptychopariida.

The Olenacea are a group of trilobites known from the Middle Cambrian to the Upper Ordovician (Harrington et al. 1959).

Characters (from Harrington et al. 1959): Exoskeleton subovate to elongate, opisthoparian (exceptionally proparian), micropygous to (rarely)subisopygous, with rounded or spinose genal angles. Cephalon with narrow border, small to medium-sized eyes and in most forms distinct eye ridges; glabella tapering forward, with simple, sigmoidal, or bifurcate lateral furrows (if present); preglabellar field present or absent; librigenae separated by median suture or fused together. Hypostoma probably free. Thorax with 9 to 24 segments. Pygidium with or without marginal spines.

<==Olenacea
    |--Elviniinae [Elviniidae] S80
    |    |--Wentsuia Sun 1935 S80
    |    |    |--*W. granulosa Sun 1935 S80
    |    |    |--W. iota Shergold 1980 S80
    |    |    `--‘Elrathia’ munda Resser & Endo in Endo & Resser 1937 S80
    |    `--Dunderbergia S80
    |         |--*D. nitida (Hall & Whitfield 1877) S80
    |         |--D. bigranulosa Palmer 1960 S80
    |         |--D. brevispina Palmer 1965 S80
    |         `--D. simplex Rasetti 1961 S80
    |--Pterocephaliidae S80
    |    |--Pterocephaliinae S80
    |    |    |--Sigmocheilus S80
    |    |    `--Pterocephalia S80
    |    |--Changshaniinae S80
    |    |    |--Changshania conica S80
    |    |    `--Parachangshania Chien 1958 S80
    |    |         `--‘Maladioidella’ elongata Endo 1944 S80
    |    `--Eugonocare Whitehouse 1939 [Aphelaspidinae] S80
    |         |--*E. tessellatum Whitehouse 1939 S80
    |         |--E. quadratum Henderson 1976 S80
    |         `--E. whitehousei Henderson 1976 S80
    `--Olenidae S80
         |  i. s.: Hedinia regalis Troedsson 1937 S80
         |         Porterfieldia punctata FO99
         |         Peltura scarabaeoides FO99
         |--Leptoplastus Angelin 1854 [Leptoplastinae] S80
         |    |--*L. stenotus Angelin 1854 S80
         |    `--L. rhaphidophorus Angelin 1854 S80
         `--Oleninae S80
              |--Plicatolina Shaw 1951 S80
              |    |--*P. kindlei Shaw 1951 S80
              |    |--P. lucida Lazarenko 1966 S80
              |    |--P. perlata lazarenko 1966 S80
              |    |--P. quadrata Pokrovskaya 1966 S80
              |    |--P. scalpta Harrington & Leanza 1957 S80
              |    `--P. yakutica Pokrovskaya 1966 S80
              `--Olenus LN05
                   |--O. attenuatus (Boeck 1838) LN05
                   |--O. dentatus (Westergård 1922) LN05
                   |--O. gibbosus (Wahlenberg 1821) LN05
                   `--+--O. transversus LN05
                      `--+--O. truncatus (Brünnich 1781) LN05
                         `--O. wahlenbergi (Wahlenberg 1821) LN05

*Type species of generic name indicated

REFERENCES

[FO99] Fortey, R. A. & R. M. Owens. 1999. Feeding habits in trilobites. Palaeontology 42 (3): 429–465.

Harrington, H. J., G. Henningsmoen et al. 1959. Systematic descriptions. In Treatise on Invertebrate Paleontology pt O. Arthropoda 1 pp. O170–O540. Geological Society of America and University of Kansas Press.

[LN05] Lauridsen, B. W., & A. T. Nielsen. 2005. The Upper Cambrian trilobite Olenus at Andrarum, Sweden: a case of iterative evolution? Palaeontology 48 (5): 1041–1056.

[S80] Shergold, J. H. 1980. Late Cambrian trilobites from the Chatsworth Limestone, western Queensland. Bureau of Mineral Resources, Geology and Geophysics—Bulletin 186: 1–111.