Khairkhaniidae

The Lower Devonian onychochilid Voskopiella barborae, from Frýda (1992).

Belongs within: Mollusca.

The Khairkhaniidae were a Lower Cambrian family of planispiral molluscs, possibly basal or stem gastropods. Parkhaev (2002) suggested that the khairkhaniid genus Protowenella was ancestral to the Upper Cambrian to Lower Devonian Onychochilidae, a family of turbospiral hyperstrophic forms, though he did not explicitly state his reasons for suggesting the connection.

Characters of Khairkhaniidae (from Parkhaev 2002): Shell planispiral or almost planispiral with up to three or four coils. Aperture almost circular with diameter significantly less than diameter of shell (Parkhaev's listing of a paired pallial caecum as a character of Khairkhaniidae should be treated with caution, as this is not something that can be observed directly from the shell).

<==Khairkhaniidae [Khairkhaniiformes]
    |--+--+--Barskovia Golubev 1976 [incl. Nekolenia Vassiljeva 1998] P02
    |  |  `--Philoxenella Vostokova 1962 P02
    |  `--+--Khairkhania Missarzhevsky 1981 P02
    |     |--Ardrossania Runnegar in Bengtson et al. 1990 P02
    |     `--Michniakia Missarzhevsky 1966 P02
    `--+--Protowenella Runnegar & Jell 1976 P02
       `--+--Xinjispira Yu & Rong 1987 P02
          `--Clisospiroidea (see below for synonymy) BR05
               |  i. s.: Mimospira FB01
               |--Clisospiridae BR05
               |    |--Atracura Horný 1964 [Atracurinae] BR05
               |    |--Trochoclisa Horný 1964 [Trochoclisinae] BR05
               |    `--Clisospirinae BR05
               |         |--Clisospira Billings 1865 BR05
               |         `--Progalerus Holzapfel 1895 [Progalerinae] BR05
               `--Onychochilidae P02
                    |--Beshtashella Missarzhevsky in Missarzhevsky & Mambetov 1981 P02
                    |--Voskopiella barborae Frýda 1992 FBL02
                    |--Onychochilus Lindström 1884 [Onychochilinae] BR05
                    |--Hyperstrophema Horný 1964 [Hyperstropheminae] BR05
                    |--Scaevogyra Whitfield 1878 [Scaevogyridae, Scaevogyrinae] BR05
                    `--Yuwenia Runnegar 1981 [incl. Kistassella Missarzhevsky 1989] P02
                         `--*Y. bentleyi Runnegar 1981 [incl. *Kistassella spiralis Missarzhevsky 1989] P02

Clisospiroidea [Clisospiracea, Dextrobranchia, Mimospirina, Onychochiliformes, Onychochiloidea, Umbraculiformii] BR05

*Type species of generic name indicated

REFERENCES

[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.

Frýda, J. 1992. Mode of life of a new onychochilid mollusc from the Lower Devonian of Bohemia. Journal of Paleontology 66 (2): 200-205.

[FB01] Frýda, J., & R. B. Blodgett. 2001. The oldest known heterobranch gastropod, Kuskokwimia gen. nov., from the Early Devonian of west-central Alaska, with notes on the early phylogeny of higher gastropods. Vĕstnik Českého Geologického Ústavu 76 (1): 39-53.

[FBL02] Frýda, J., R. B. Blodgett & A. C. Lenz. 2002. New Early Devonian gastropods from the families Crassimarginatidae (new family) and Scoliostomatidae (new family), Royal Creek area, Yukon Territory, Canada. Journal of Paleontology 76 (2): 246-255.

[P02] Parkhaev, P. Yu. 2002. Phylogenesis and the system of the Cambrian univalved mollusks. Paleontologicheskii Zhurnal 2002 (1): 27-39 (transl. Paleontological Journal 36 (1): 25-36).

Amphigastropoda

Steinkern of Pelagiella, from Landing et al. (2002).

Belongs within: Helcionelliformes.
Contains: Bellerophontidae.

The name 'Amphigastropoda' has been used for an assemblage of planispiral or cap-shaped, mostly Palaeozoic molluscs. Primarily centered around the Bellerophontoidea, other groups that have been included in the Amphigastropoda by various authors include the Pelagielliformes, Cyrtolitidae and Tryblidiidae. The grouping was originally proposed on the basis that these animals were gastropod-like but untorted; however, studies of muscle scars the genus Bellerophon itself indicate that it, at least, underwent torsion, and the Amphigastropoda in the broad sense is probably a polyphyletic assemblage (Peel 1972).

Amphigastropoda
    |--Cyrtolitida BR05
    |--Tryblidiidae T59 [Tryblidiida BR05]
    |    |--Helcionopsis Ulrich & Schofield 1897 T59
    |    |    `--*H. striata T59
    |    |--Pilina Koken 1925 T59
    |    |    `--*P. unguis [=Tryblidium unguis] T59
    |    |--Tryblidium Lindström 1880 T59
    |    |    |--*T. reticulatum T59
    |    |    `--T. nycteis (Billings 1862) [=Metoptoma nycteis] T59
    |    `--Vallatotheca Foerste 1914 T59
    |         |--*V. manitoulini T59
    |         `--V. elegantula (Chapman 1916) (see below for synonymy) T59
    |--Pelagielliformes [Orthostrophina, Pelagiellacea, Pelagiellida, Pelagielloidea] BR05
    |    |--Pelagiellidae BR05
    |    |    |--Costipelagiella Horny 1964 P02
    |    |    |--Tianzhushanospira Yu 1979 P02
    |    |    |--Proeccyliopterus Kobayashi 1939 [Proeccyliopteridae] BR05
    |    |    |--Protoscaevogyra Kobayashi 1939 [Protoscaevogyridae] BR05
    |    |    |--Tannuspira Missarzhevsky 1989 P02
    |    |    |    `--T. magnifica Missarzhevsky 1989 P02
    |    |    `--Pelagiella Matthew 1895 BR05
    |    |         |--P. adunca Missarzhevsky 1966 P02
    |    |         |--P. lorenzi Kobayashi 1939 P02
    |    |         `--P. madianensis Zhou & Xiao 1984 P02
    |    `--Aldanellidae BR05
    |         |--Paraaldanella Golubev 1976 P02
    |         |--Nomgoliella Missarzhevsky 1981 P02
    |         |    |--N. australiensis P02
    |         |    |--N. rotunda P02
    |         |    `--N. sinistrovolubilis Missarzhevsky 1981 P02
    |         `--Aldanella Vostokova 1962 BR05
    |              |--A. crassa Missarzhevsky 1969 P02
    |              `--+--A. utchurica Missarzhevsky 1969 P02
    |                 `--+--A. rozanovi Missarzhevsky 1966 P02
    |                    `--A. operosa Missarzhevsky 1966 P02
    `--Bellerophontoidea [Bellerophontacea, Bellerophontida, Bellerophontina] BR05
         |--Bellerophontidae BR05
         |--Bucanella Meek 1871 [=Bucaniella Fischer 1885; Bucanellinae, Bucaniellinae] BR05
         |--Tremanotus Hall 1867 [=Trematonotus Fischer 1885; Tremanotidae, Tremanotinae, Trematonotidae] BR05
         |--Tropidodiscus Meek & Worthen 1866 [Tropidodiscidae, Tropidodiscinae] BR05
         |--Pterothecidae BR05
         |    |--Pterotheca Salter 1852 [Pterothecinae] BR05
         |    |--Carinaropsis Hall 1847 [Carinaropsidae, Carinaropsinae] BR05
         |    `--Pedasiola Spriesterbach 1919 [Pedasiolinae] BR05
         |--Sinuitidae [Sinuitoidea] BR05
         |    |--Aiptospira Wang 1980 [Aiptospirinae] BR05
         |    |--Hispanosinuites Frýda & Gutierrez-Marco 1996 [Hispanosinuitinae] BR05
         |    `--Sinuitinae BR05
         |         |--Sinuites Koken 1896 BR05
         |         `--Protowarthia Ulrich & Scofield 1897 [Protowarthiidae] BR05
         |--Euphemitidae BR05
         |    |--Paleuphemites Horný 1962 [Paleuphemitinae] BR05
         |    `--Euphemitinae BR05
         |         |--Euphemites Warthin 1930 BR05
         |         `--Warthia Waagen 1880 W63
         |              |--*W. brevisinuata Waagen 1880 W63
         |              |--W. americana Girty 1909 W63
         |              |--W. micromphala (Morris 1845) (see below for synonymy) W63
         |              |--W. perspecta Fletcher 1958 W63
         |              `--W. stricta (Dana 1847) (see below for synonymy) W63
         `--Bucaniidae BR05
              |--Salpingostoma Roemer 1876 (see below for synonymy) BR05
              |--Undulabucania Wahlman 1992 [Undulabucaniinae] BR05
              |--Plectonotinae BR05
              |    |--Plectonotus Clarke 1899 [Plectonotini] BR05
              |    `--Boucotonotus Frýda & Manda 1997 [Boucotonotini] BR05
              `--Bucaniinae [Bucaniides] BR05
                   |--Grandostoma Horný 1962 [Grandostomatidae, Grandostomatinae] BR05
                   `--Bucania Hall 1847 BR05
                        |--B. angustafasciata Waagen 1880 W63
                        `--B. kattaensis Waagen 1880 [incl. B. k. var. carinata Wanner 1942] W63

Salpingostoma Roemer 1876 [Salpingostomatidae, Salpingostomatides, Salpingostomatinae, Salpingostominae] BR05

Vallatotheca elegantula (Chapman 1916) [=Helcionopsis elegantulum; incl. Tryblidium (Metoptoma) nycteis Cresswell 1893 non Metoptoma nycteis Billings 1862, Capulus nycteis, Helcionopsis nycteis] T59

Warthia micromphala (Morris 1845) [=Bellerophon micromphalus, B. (Warthia) micromphala, Goniatites (Agathiceras) micromphalus, G. (Prolecanites) micromphalus; incl. B. undulatus Dana 1847] W63

Warthia stricta (Dana 1847) [=Bellerophon strictus, Goniatites strictus; incl. B. contractus Sowerby in Jukes 1847 (n. n.)] W63

*Type species of generic name indicated

REFERENCES

[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.

Landing, E., G. Geyer & K. E. Bartowski. 2002. Latest Early Cambrian small shelly fossils, trilobites, and Hatch Hill dysaerobic interval on the Québec continental slope. Journal of Paleontology 76 (2): 287-305.

[P02] Parkhaev, P. Yu. 2002. Phylogenesis and the system of the Cambrian univalved mollusks. Paleontologicheskii Zhurnal 2002 (1): 27-39 (transl. Paleontological Journal 36 (1): 25-36).

Peel, J. S. 1972. Observations on some Lower Palaeozoic tremanotiform Bellerophontacea (Gastropoda) from North America. Palaeontology 15 (3): 412-422.

[T59] Talent, J. A. 1958-1959. Vallatotheca (Monoplacophora) in the Lower Devonian of Victoria. Mining and Geological Journal 6 (3): 3-4.

[W63] Waterhouse, J. B. 1963. Permian gastropods of New Zealand. Part 1 – Bellerophontacea and Euomphalacea. New Zealand Journal of Geology and Geophysics 6 (1): 88-112.

Bellerophontidae

Various views of Bellerophon yochelsoni, from Pan & Erwin (2002).

Belongs within: Amphigastropoda.

The Bellerophontidae were an Ordovician to Triassic family of coiled molluscs. They were superficially similar to small nautiloids with a deep sinus or slit at the aperture on the outer side of the coil. The umbilicus was narrow or absent, and the shell was ornamented by prominent growth lines (Knight et al. 1960). Most authors have considered the bellerophontids as basal (probably stem-)gastropods, though there is some debate on whether they underwent torsion like modern gastropods.

<==Bellerophontidae
    |  i. s.: Stachella PE94
    |--Bucanopsis Ulrich 1897 [Bucanopsinae] BR05
    |--Cymbularia Koken 1896 [Cymbulariidae, Cymbulariinae] BR05
    |--Bellerophontinae BR05
    |    |--Liljevallospira Knight 1945 [Liljevallospiridae] BR05
    |    `--Bellerophon Montfort 1808 BR05
    |         |--B. heterocanalicutus Pan & Yu 1996 PE02
    |         |--B. (Bellerophon) yochelsoni Pan & Erwin 2002 PE02
    |         `--B. zhesiensis DX84
    `--Knightitinae [Knightitidae] BR05
         |--Knightites Moore 1941 BR05
         `--Retispira Knight 1945 W63, PE02
              |--*R. bellireticulata Knight 1945 PE02 [=Knightites (*Retispira) bellireticulata W63]
              |--R. matanensis Pan & Yu 1993 PE02
              |--R. multicancellata Yu in Wang & Xi 1980 PE02
              |--R. sinensis Pan & Erwin 2002 PE02
              |--R. strictocanaliculata Yu in Wang & Xi 1980 PE02
              |--R. strictura Yu in Wang & Xi 1980 PE02
              `--R. texana Yochelson 1960 PE02

*Type species of generic name indicated

REFERENCES

[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.

[DX84] Ding Y., Xia G., Duan C., Li W., Liu X. & Liang Z. 1984. Study on the early Permian stratigraphy and fauna in Zhesi district, Nei Mongol Zizhiqu (Inner Mongolia). Bulletin Tianjin Institure Geol. Min. Res. 10.

Knight, J. B., L. R. Cox, A. M. Keen, R. L. Batten, E. L. Yochelson & R. Robertson. 1960. Gastropoda—Systematic description. 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-I310. Geological Society of America, Inc. and University of Kansas Press.

[PE94] Pan H.-Z. & D. H. Erwin. 1994. Gastropod diversity patterns in south China during the Chihsia-Ladinian and their mass extinction. Palaeoworld 4: 249-262.

[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.

[W63] Waterhouse, J. B. 1963. Permian gastropods of New Zealand. Part 1 – Bellerophontacea and Euomphalacea. New Zealand Journal of Geology and Geophysics 6 (1): 88-112.