Lyocyclus solutus, from Thiele (1925).

Belongs within: Gastropoda.
Contains: Coelostylinidae, Pseudozygopleuridae, Goniospiridae, Zygopleuridae, Vermetidae, Pseudomelaniidae, Epitoniidae, Littorinina, Stromboidea, Capulidae, Hipponicidae, Calyptraeidae, Naticidae, Cypraeoidea, Tonnoidea, Neogastropoda.

The Hypsogastropoda are a major clade of caenogastropods possessing an exophallic penis positioned behind the right cephalic tentacle (Simone 2011). They are primarily marine though several lineages have diversified in fresh water. Basal relationships within the hypsogastropods remain poorly established. However, a clade Latrogastropoda including the neogastropods and related taxa may be recognised by the presence of a pair of retractor muscles in association with the snout, an enlarged mantle border that covers the head when active, and a sharp pointed apex on the marginal radular teeth. Latrogastropoda may be divided between the Stromboidea and Rhynchogastropoda with the latter clade united by the presence of a retractible proboscis. A clade containing the Capuloidea, Hipponicidae and Calyptraeoidea contains species with a hairy periostracum and nuchal flaps on both sides of the head. The remaining Rhynchogastropoda, forming the clade Adenogastropoda, show a tendency towards predatory behaviour with a simplified stomach and modified foregut. The Siphonogastropoda, including the Cypraeoidea, Tonnoidea and Neogastropoda, share a fusiform shell, determinate growth, and a well-developed siphonal canal. Finally, the Tonnoidea and Neogastropoda are united by synapomorphies including a terminal nucleus on the operculum and development of the incurrent siphon into an exploratory structure (Simone 2011).

The Late Palaeozoic high-spired Pseudozygopleuroidea represent possible early precursors of the Hypsogastropoda.

Synapomorphies (from Simone 2011): Nephridial gland present. Loss of pair of auxiliary muscle (m12) of horizontal muscle (m6). Exophallic penis present, situated behind and at right of right cephalic tentacle, with respective duct. Development of albumen and capsule glands at pallial oviduct. Each statocyst with single statolith.

<==Hypsogastropoda (see below for synonymy)
    |--+--Pseudomelaniidae PC08
    |  `--Epitoniidae PC08
    `--+--‘Amauropsis’ rossiana Smith 1907 S11
       `--+--Littorinina GS75
          `--Latrogastropoda [Neomesogastropoda, Strombogastropoda, Troschelina] BD12
               |  i. s.: Eunatica BD12
               |           |--E. papilla (Gmelin 1791) BD12
               |           `--E. telaaraneae (Melville 1901) [=Natica (Eunatica) telaaraneae] M01
               |--Stromboidea S11
               `--Rhynchogastropoda S11
                    |--+--Capuloidea BD12
                    |  |    |--Capulidae S11
                    |  |    |--Gyrotropis Gabb 1877 BR17 [Gyrotropidae BD12]
                    |  |    |    `--*G. squamosa Gabb 1877 BR17 [=Trichotropis squamosa BD12]
                    |  |    `--Haloceras Dall 1889 BR05 [Haloceratidae BR17]
                    |  |         `--*H. cingulata (Verrill 1884) [=Cithna cingulata] BR17
                    |  `--+--Hipponicidae S11
                    |     `--Calyptraeoidea [Calyptaeiformes, Calyptraeiformi, Calyptraeiformii, Calyptraeoidei] BD12
                    |          |  i. s.: Thylacus Conrad 1860 BD12
                    |          |           `--*T. cretaceus Conrad 1860 BD12
                    |          |--Calyptraeidae S11
                    |          `--Lyocyclus Thiele 1925 BR05 [Lyocyclidae GS75, Lyocyclinae]
                    |               `--*L. solutus Thiele 1925 BR17
                    `--Adenogastropoda [Epinephridia, Phaneropneumana, Phaneropneumona, Simrothina] S11
                         |--Naticidae S11
                         `--Siphonogastropoda [Angyostomata, Siphonata] S11
                             |--Cypraeoidea ZLK11
                             `--Peogastropoda S11
                                  |--Tonnoidea ZLK11
                                  `--Neogastropoda ZLK11

Hypsogastropoda incertae sedis:
  Coelostylinidae BR05
  Pseudozygopleuroidea [Zygopleuroidea] BR17
    |--Pseudozygopleuridae PC08
    |--Goniospiridae BR17
    |--Zygopleuridae BR17
    |--Pommerozygiidae BR17
    |    |--Pommerozygia Gründel 1998 BR05
    |    |    `--*P. neckeritzensis Gründel 1998 BR17
    |    `--Costazygia Gründel 1999 G01
    |         `--C. bilzi Gründel 1999 G01
    `--Protorculidae BR17
         |--Protorcula Kittl 1892 BR05
         |    `--*P. subpunctata (Münster 1841) (see below for synonymy) B91
         `--Battenizyga Nützel & Erwin 2002 NE02
              `--*B. eotriassica (Batten & Stokes 1986) [=Anoptychia eotriassica] NE02
  Vermetoidea BR17
    |--Vermetidae BR17
    `--Sakarahella Bandel 2006 [Sakarahellidae] BR17
         `--*S. angulata Bandel 2006 BR17

Hypsogastropoda [Acropththalma, Ancistroglossata, Caecoidei, Cerithiopsoidei, Circulopsoidei, Ctenoglossa, Cyclomyaria, Echinospiracea, Echinospirida, Haemiglossata, Hamiglossa, Hydrobioidea, Littorinacea, Littorinata, Littoriniformes, Littorinimorpha, Littorinoidei, Melanellida, Metamesogastropoda, Nudipeda, Peloridae, Pleurembolica, Ptenoglossa, Rissoacea, Rissooidei, Scaphoconchoidea, Semiproboscidifera, Truncatellacea, Tubulibranchiata, Vermivora]

*Protorcula subpunctata (Münster 1841) [=Turritella subpunctata, Undularia (*Protorcula) subpunctata] B91

*Type species of generic name indicated


[B91] Bandel, K. 1991. Über triassische "Loxonematoidea" und ihre Beziehungen zu rezenten und paläozoischen Schnecken. Paläontologische Zeitschrift 65 (3–4): 239–268.

[BD12] Bandel, K., & D. T. Dockery III. 2012. Protoconch characters of Late Cretaceous Latrogastropoda (Neogastropoda and Neomesogastropoda) as an aid in the reconstruction of the phylogeny of the Neogastropoda. Freiberger Forschungshefte C542 (psf 20): 93–128.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[GS75] Golikov, A. N., & Y. I. Starobogatov. 1975. Systematics of prosobranch gastropods. Malacologia 15 (1): 185–232.

[G01] Gründel, J. 2001. Neritimorpha und weitere Caenogastropoda (Gastropoda) aus dem Dogger Norddeutschlands und des nordwestlichen Polens. Berliner Geowissenschaftliche Abhandlungen, Reihe E 36: 45–99.

[M01] Melville, J. C. 1901. A few further remarks upon the Erythrean molluscan fauna, with descriptions of seven species from Aden, in the collection of Commander E. R. Shopland, R.I.M. Annals and Magazine of Natural History, series 7, 7: 550–556.

[NE02] Nützel, A. & D. H. Erwin. 2002. Battenizyga, a new Early Triassic gastropod genus with a discussion of the caenogastropod evolution at the Permian/Triassic boundary. Paläontologische Zeitschrift 76 (1): 21–27.

[PC08] Ponder, W. F., D. J. Colgan, J. M. Healy, A. Nützel, L. R. L. Simone & E. E. Strong. 2008. Caenogastropoda. In: Ponder, W. F., & D. L. Lindberg (eds) Phylogeny and Evolution of the Mollusca pp. 331–383. University of California Press: Berkeley.

[S11] Simone, L. R. L. 2011. Phylogeny of the Caenogastropoda (Mollusca), based on comparative morphology. Arquivos de Zoologia 42 (4): 161–323.

[ZLK11] Zou, S., Q. Li & L. Kong. 2011. Additional gene data and increased sampling give new insights into the phylogenetic relationships of Neogastropoda, within the caenogastropod phylogenetic framework. Molecular Phylogenetics and Evolution 61: 425–435.

Last updated: 14 June 2021.

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