The Upper Carboniferous lophospirid Worthenia, from here.

Belongs within: Vetigastropoda.

The Ordovician to Middle Triassic Trochonematoidea included the gastropod families Trochonematidae and Lophospiridae. The two families shared a characteristic shell morphology: a median angulation running along or above the midline of each spire (replaced by a shallow sinus in some Trochonematidae), with further angulations above and/or below. The inner lining of the shell was nacreous. In the Lophospiridae, the aperture of the shell included a deep labral sinus or slit, but in the Trochonematidae this was replaced by an internal channel (Knight et al. 1960).

<==Trochonematoidea [Lophospiroidea, Trochonematacea]
    |--Trochonematidae [Trochonematinae] BR05
    |    |--Proturritella Koken 1889 (see below for synonymy) KC60
    |    |    `--*P. gracilis Koken 1889 KC60
    |    |--Amaurotoma Knight 1945 KC60
    |    |    `--*A. subsinuata (Meek & Worthen 1861) [=Pleurotomaria subsinuata] KC60
    |    |--Cyclobathmus Knight 1940 KC60
    |    |    `--*C. haworthi (Beede 1907) [=Trepospira haworthi] KC60
    |    |--Cyclites Knight 1940 KC60
    |    |    `--C. depressus (Beede 1907) (see below for synonymy) KC60
    |    `--Trochonema Salter 1859 BR05
    |         |  i. s.: T. etheridgei F71
    |         |         T. montgomerii (Etheridge 1896) [=Eunema montgomerii] F71
    |         |         T. nodosa Etheridge 1898 F71
    |         |--*T. (Trochonema) umbilicata (Hall 1847) [=Pleurotomaria umbilicata] KC60
    |         |--T. (Eunema Salter 1859) KC60
    |         |    `--T. (*E.) strigillatum (Salter 1859) [=*Eunema strigillatum] KC60
    |         `--T. (Trochonemopsis Meek 1872) KC60
    |              `--T. (*T.) tricarinatum Meek 1872 [=T. tricarinata] KC60
    `--Lophospiridae [Gyronematinae, Lophospirinae, Ruedemanniinae] BR05
         |--Gyronema Ulrich in Ulrich & Scoffield 1897 BR05
         |    `--*G. pulchellum (Ulrich & Scofield 1897) [=Trochonema (*Gyronema) pulchellum] KC60
         |--Ruedemannia Foerste 1914 BR05 [incl. Coronilla Perner 1907 non Beneden 1871] KC60
         |    `--*R. lirata (Ulrich in Ulrich & Scofield 1897) [=Lophospira (Seelya) lirata] KC60
         |--Trochonemella Okulitch 1935 KC60
         |    `--*T. notabilis (Ulrich in Ulrich & Scofield 1897) [=Lophospira notabilis] KC60
         |--Longstaffia Cossmann 1908 [=Tubulosa Cossmann 1908 non Schweigger 1820] KC60
         |    `--*L. tubulosa (Lindström 1884) [=Pleurotomaria tubulosa, *Tubulosa tubulosa] KC60
         |--Antitrochus Whidborne 1891 KC60
         |    `--*A. arietinus Whidborne 1891 KC60
         |--Araeonema Knight 1933 (see below for synonymy) KC60
         |    `--*A. virgatum Knight 1933 KC60
         |--Rhabdotocochlis Knight 1933 KC60
         |    `--*R. rugata Knight 1933 KC60
         |--Omphalonema Grabau 1936 KC60
         |    `--*O. multispirale Grabau 1936 KC60, W63
         |--Lophospira Whitfield 1886 BR05 (see below for synonymy)
         |    |--*L. milleri (Miller 1877) KC60 (see below for synonymy)
         |    `--L. conradana YN83
         |--Cinclidonema Knight 1945 W63
         |    |--*C. texanum Knight 1945 W63
         |    `--C. dignitas Waterhouse 1963 W63
         |--Manzanospira Batten 1989 PE02
         |    |--*M. manzanicum (Girty 1909) [=Phanerotrema manzanicum] PE02
         |    |--M. turrita Pan & Erwin 2002 PE02
         |    `--M. wordensis Batten 1989 PE02
         |--Loxoplocus Fischer 1885 KC60
         |    |--L. (Loxoplocus) solutus (Whiteaves 1884) (see below for synonymy) KC60
         |    `--L. (Donaldiella Cossmann 1903) (see below for synonymy) KC60
         |         `--L. (*D.) filosa (Donald 1902) [=*Goniospira filosa] KC60
         |--Yunnania Mansuy 1912 B79
         |    |--*Y. termieri Mansuy 1912 B79
         |    |--Y. meridionalis Mansuy 1914 B79
         |    |--Y. romanovskyi Licharew 1967 B79
         |    `--Y. semigranulosa Licharew 1967 B79
         `--Worthenia de Koninck 1883 TE01 [incl. Platyworthenia Chronic 1952 KC60]
              |--*W. tabulata (Conrad 1835) [=Turbo tabulatus] KC60
              |--W. conica Assmann 1924 TE01
              |--W. corrugata Chronic 1952 TE01
              |--W. esinensis TE01
              |--W. extendia Tong & Erwin 2001 TE01
              |--W. marmolatae Kittl 1894 TE01
              |--W. pura TE01
              `--W. subpunctata TE01

Araeonema Knight 1933 [incl. Palaeoturbina Wenz 1938, Turbina deKoninck 1881 non Brown in Herrmannsen 1847] KC60

Cyclites depressus (Beede 1907) [=Wortheniopsis depressa; incl. Pleurotomaria multilineata Girty 1908, *Cyclites multilineatus] KC60

Lophospira Whitfield 1886 BR05 [incl. Ptychonema Perner 1903 KC60, Schizolopha Ulrich in Ulrich & Scofield 1897 KC60]

*Lophospira milleri (Miller 1877) KC60 [=Murchisonia milleri KC60, Loxoplocus (Lophospira) milleri KC60, M. bicincta Hall 1847 non M’Coy 1844 KC60, Lophospira bicincta YN83]

Loxoplocus (Donaldiella Cossmann 1903) [=Goniospira Donald 1902 non Cossmann 1896; incl. Pagodispira Grabau 1922] KC60

Loxoplocus (Loxoplocus) solutus (Whiteaves 1884) [=Murchisonia soluta; incl. M. tropidophora Whiteaves 1884, *Loxoplocus tropidophorus] KC60

Proturritella Koken 1889 [incl. Gonionema Koken 1896, Nematotrochus Koken 1925, Pseudeunema Cossmann 1899] KC60

*Type species of generic name indicated


[B79] Batten, R. L. 1979. Gastropods from Perak, Malaysia. Part 2. The trochids, patellids and neritids. American Museum Novitates 2685: 1-26.

[BR05] Bouchet, P., J.-P. Rocroi, J. Frýda, B. Hausdorf, W. Ponder, Á. Valdés & A. Warén. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1-2): 1-397.

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

[KC60] Knight, J. B., L. R. Cox, A. M. Keen, R. L. Batten, E. L. Yochelson & R. Robertson. 1960. Gastropoda: systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia, pp. I169-I331. Geological Society of America, and University of Kansas Press.

[PE02] Pan, H.-Z., & D. H. Erwin. 2002. Gastropods from the Permian of Guangxi and Yunnan Provinces, south China. Journal of Paleontology 76 (Memoir 56): 1-49.

[TE01] Tong, J., & D. H. Erwin. 2001. Triassic gastropods of the southern Qinling Mountains, China. Smithsonian Contributions to Paleobiology 92: 1-47.

[W63] Waterhouse, J. B. 1963. Permian gastropods of New Zealand. Part 4—Platyceratacea, Anomphalacea, Neritacea, and correlations. New Zealand Journal of Geology and Geophysics 6 (5): 817-842.

[YN83] Yu W. & Ning H. 1983. Middle Ordovician gastropods from Ruoqiang, Xinjiang. Acta Palaeontologica Sinica 22 (2): 199-203.

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