Trigonotarbida

Belongs within: Euchelicerata.

The Trigonotarbida are a group of fossil arachnids known from the late Silurian to early Permian (Dunlop & Selden 2004). Recent analyses have agreed on a relationship between the trigonotarbids and the modern tetrapulmonates, including spiders and whip scorpions.

Characters (Dunlop & Selden 2004): Opisthosoma with nine dorsal tergites divided into median and lateral plates; tergites 2 and 3 commonly fused into single macrotergite; two pairs of book-lungs present.

<==Trigonotarbida [Anthracomarti]
    |--PalaeocharinidaeDS04
    |    |--Gilboarachne griersoni Shear et al. 1987S93
    |    |--Aculeatarbus depressus Shear et al. 1987S93
    |    |--Palaeocharinus Hirst 1923LA23, DS04
    |    |    |--P. calmani Hirst 1923S93
    |    |    |--P. kidstoni Hirst 1923S93
    |    |    |--P. rhyniensis Hirst 1923S93 [incl. P. hornei Hirst 1923GD14, S93]
    |    |    `--P. scourfieldi Hirst 1923S93
    |    `--GelasinotarbusS93
    |         |--G. bifidus Shear et al. 1987S93
    |         |--G. bonamoae Shear et al. 1987S93
    |         |--G. heptops Shear et al. 1987S93
    |         `--G. reticulatus Shear et al. 1987S93
    `--+--EophrynidaeDS04
       |    |--Adelocaris peruvianus Packard 1889F04
       |    |--Pocononia whitei (Ewing 1930)S93
       |    |--Gondwanarachne argentinensis Pinto & Hünicken 1980S93
       |    |--Petrovicia Fritsch 1904F04
       |    |    `--*P. proditoria Fritsch 1904F04
       |    |--Hemikreischeria Fritsch 1904F04
       |    |    `--*H. thevenini Fritsch 1904 [=Kreischeria geinitzii Brongniart 1901 (n. n.)]F04
       |    |--StenotrogulusF04 [incl. Cyclotrogulus Fritsch 1904RDS03]
       |    |    |--S. salmii [=Eophrynus salmii]F04
       |    |    `--‘Eophrynus’ sturii Hasse 1890 [=Cyclotrogulus sturii]F04
       |    |--Eophrynus Woodward 1871LA23, RDS03
       |    |    |--E. prestvicii (Buckland 1837)GD14, H02 [=Curculioides prestviciiF04]
       |    |    `--E. scharfi Scharf 1924RDS03
       |    `--Hemiphrynus [Hemiphrynidae, Pleuraraneae]S93
       |         |--H. hofmanni Frič 1901S93
       |         `--H. longipes Frič 1901S93
       `--Anthracomartidae [Anthracomatida]DR02
            |--Maiocercus Pocock 1911DR02
            |--Coryphomartus Petrunkevitch 1913DR02
            |--Cleptomartus Petrunkevitch 1949DR02
            |--Cryptomartus Petrunkevitch 1949DR02
            |--Oomartus Petrunkevitch 1953DR02
            |--Pleomartus Petrunkevitch 1945DR02
            |    `--P. palatinus (Ammon 1901)S93
            |--Brachylycosa Fritsch 1904DR02
            |    `--*B. carcinoides [=Arthrolycosa carcinoides]F04
            |--Vratislavia Fritsch 1904F04
            |    `--*V. silesiaca (Röm. 1879) [=Architarbus silesiacus]F04
            |--Brachypyge Woodward 1878DR02
            |    |--B. carbonis Woodward 1878 [=Eophrynus carbonis]F04
            |    `--B. celtica Pocock 1902F04
            |--Promygale Fritsch 1901 [Promygalidae]DR02
            |    |--P. bohemicaF04
            |    |--P. elegansF04
            |    `--P. rotundataF04
            `--Anthracomartus Karsch 1882LA23, DR02
                 |--*A. voelkelianus Karsch 1882DR02
                 |--A. affinis Kušta 1885F04
                 |--A. buchi (Goldenberg 1873) (n. d.)DR02
                 |--A. granulatus Fritsch 1904DR02
                 |--A. hageni (Goldenberg 1873) (n. d.)DR02
                 |--A. hindiGD14
                 |--A. krejcii Kušta 1883F04
                 |--A. minor Kušta 1885F04
                 |--A. palatinus Ammon 1900F04
                 |--A. socius Kušta 1888F04
                 `--A. trilobitus Scudder 1885F04

Trigonotarbida incertae sedis:
  Palaeotarbus jerami (Dunlop 1996)H02
  Lissomartus Petrunkevitch 1949 [Lissomartidae]RDS03
  KreischeriaF04 [KreischeriidaeDS04]
    `--K. wiedeiF04
  AnthracosironidaeDS04
    |--Arianrhoda Dunlop & Selden 2004DS04
    |    `--*A. bennetti Dunlop & Selden 2004DS04
    `--Anthracosiro Pocock 1903DS04
         |--A. fritschi Pocock 1903F04
         `--A. woodwardi Pocock 1903F04
  TrigonotarbidaeDS04
    |--TrigonotarbusDR02
    |    |--T. arnoldi Petrunkevitch 1955DR02
    |    `--T. johnsoni Pocock 1911S93
    `--Archaeomartus Størmer 1970S93, DS04
         |--A. levis Størmer 1970S93
         `--A. tuberculatus Størmer 1970S93
  AphantomartidaeDS04
    |--Alkenia Størmer 1970DS04
    |    `--A. mirabilis Størmer 1970DS04
    `--Aphantomartus Pocock 1911RDS03
         |--*A. areolatus Pocock 1911RDS03
         |--A. ilfeldicus (Scharf 1924) [=Eophrynus ilfeldicus]RDS03
         `--A. pustulatus (Scudder 1884)RDS03

*Type species of generic name indicated

References

[B86] Bernini, F. 1986. Current ideas on the phylogeny and the adaptive radiations of Acarida. Bolletino di Zoologia 53: 279–313.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

[DR02] Dunlop, J. A., & R. Rößler. 2002. The trigonotarbid arachnid Anthracomartus voelkelianus (Anthracomartidae). Journal of Arachnology 30 (2): 211–218.

[DS04] Dunlop, J. A., & P. A. Selden. 2004. A trigonotarbid arachnid from the Lower Devonian of Tredomen, Wales. Palaeontology 47 (6): 1469–1476.

[F04] Fritsch, A. 1904. Palaeozoische Arachniden. Selestverlag: Prague.

[GD14] Garwood, R. J., & J. Dunlop. 2014. Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders. PeerJ 2: e641.

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

[RDS03] Rössler, R., J. A. Dunlop & J. W. Schneider. 2003. A redescription of some poorly known Rotliegend arachnids from the Lower Permian (Asselian) of the Ilfeld and Thuringian Forest Basins, Germany. Paläontologische Zeitschrift 77 (2): 417–427.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.