Warty slug Onchidium verruculatum, copyright Cory Pittman.

Belongs within: Heterobranchia.

The Systellommatophora are a group of slug-like gastropods including the marine Onchidiidae and the terrestrial Veronicelloidea. Members of this group have two pairs of contractile tentacles on the head, with the eyes at the tip of the upper pair, and have the anus and renal opening at the posterior end of the body (Burch & Pearce 1990). The dorsum is covered by a leathery covering, the notum. The Veronicelloidea are divided between the herbivorous Veronicellidae and the carnivorous Rathouisiidae, the latter family being characterised by narrow dagger-like radular teeth (Stanisic et al. 2010).

<==Systellommatophora (see below for synonymy)BR17
    |--Onchidiidae (see below for synonymy)BR05
    |    |--Scaphis Labbé 1934 non Eschw. 1824 (ICBN) [Scaphidae]BR05
    |    |--Peronia Fleming 1822 [Peroniadae, Peroniidae]BR05
    |    |    `--*P. peronii (Cuvier 1804) [=Onchidium peronii]BR17
    |    |--Watsoniella Hoffmann 1928BR17 [=Hoffmannola Strand 1932BR05; Hoffmannolidae, Hoffmannoloidea]
    |    |    `--*W. lesliei (Stearns 1892) [=Onchidium lesliei, *Hoffmannola lesliei]BR17
    |    |--Peronina Plate 1893 [Peroninidae]BR05
    |    |    `--*P. alta Plate 1893BR17
    |    |--Oncis Plate 1893BR17 [=Platevindex Baker 1938BR05; Platevindecidae]
    |    |    `--*O. coriaceus (Semper 1880) [=Onchidium coriaceum, *Platevindex coriaceus]BR17
    |    |--Quoya Labbé 1934BR17 [=Quoyella Starobogatov 1976BR05; Quoyellidae]
    |    |    `--*Q. indica Labbé 1934 [=*Quoyella indica]BR17
    |    |--Onchidina Semper 1882 [Onchidinidae]BR05
    |    |    `--*O. australis Semper 1882BR17
    |    |--Arctonchis Dall 1905O27
    |    |    `--*A. borealis (Dall 1871) [=Onchidella borealis]O27
    |    |--Onchidella Gray 1850BR05 (see below for synonymy)
    |    |    |--*O. nigricans (Quoy & Gaimard 1832) [=Onchidium nigricans]BR17
    |    |    |--O. binneyi Stearns 1893PP78
    |    |    |--O. campbelli Filhol 1880P61
    |    |    |--O. celticaP61
    |    |    |--O. flavescens Wissel 1904P61
    |    |    |--O. irrorata (Gould 1852)P61
    |    |    |--O. obscura Plate 1893P61
    |    |    `--O. patelloides (Quoy & Gaimard 1832)P61
    |    `--Onchidium Buchanan 1800 [=Oncidium (l. c.) nec Sw. 1800 (ICZN) nec Nees 1823 (ICZN); Onchidiinae]BR05
    |         |--*O. typhae Buchanan 1800BR17
    |         |--O. buetschlii Stantchinsky 1907H09
    |         |--O. carpenteriC64
    |         |--O. daemelli Semper 1882W78
    |         |--O. fungiforme Stantchinsky 1907H09
    |         |--O. meriakrii Stantchinsky 1907H09
    |         |--O. pallidipes Tapparone Canefri 1889 [=Oncidium pallidipes]TC89
    |         |--O. pallipesTC89
    |         |--O. punctatum Quoy & Gaimard 1832H09
    |         |--O. tigrinumTC89
    |         |--O. tumidum Semper 1880H09
    |         `--O. verruculatum Cuvier 1804H09
    `--Veronicelloidea [Mesogoneata, Veronicellacea]SS10
         |--Rathouisiidae [Rathouisiadae]SS10
         |    |--Rathouisia Heude 1884BR05
         |    |    `--*R. sinensis (Heude 1882) [=Vaginulus sinensis]BR17
         |    |--Prisma Simroth 1891SS10
         |    |    `--P. prismatica (Tapparone-Canefri 1883)SS10
         |    `--Atopos Simroth 1891SS10
         |         |--*A. semperi Simroth 1891SS10
         |         `--A. australis (Heynemann 1876)SS10
         `--Veronicellidae [Imeriniinae, Sarasinulinae, Vaginuloidea, Veronicellida, Veronicelliformes]SS10
              |--Imerinia Cockerell 1891BR05
              |    `--*I. grandidieri (Crosse & Fischer 1871) [=Vaginula grandidieri]BR17
              |--Veronicella Blainville 1817 [Veronicellinae]BR05
              |    |--*V. laevis Blainville 1817BR17
              |    `--V. floridanaR64
              |--Semperula Grimpe & Hoffmann 1925 [Semperulinae]BR05
              |    |--*S. idae (Semper 1885) [=Vaginula idae]BR17
              |    `--S. insularis Thomé 1983JG19
              |--Sarasinula Grimpe & Hoffmann 1924SS10
              |    `--*S. plebeia (Fischer 1868)BR17 [=Vaginula plebeiaBR17; incl. V. hedleyi Simroth 1889SS10]
              |--Leonardia Tapparone Canefri 1889TC89
              |    `--*L. nevilliana Tapparone Canefri 1889TC89
              |--Angustipes Colosi 1922BP90
              |    `--A. ameghiniBP90
              |--Diplosolenodes Thomé 1975BP90
              |    `--D. occidentaleBP90
              |--Pseudoveronicella Germain 1908 [Pseudoveronicellinae]BR05
              |    |--*P. gravieri (Germain 1908) [=Veronicella gravieri]BR17
              |    `--P. zootoca (Hoffmann 1927)TMT96
              |--Vaginulus Férussac 1821 [Anadenia, Euadenia, Mesotremata, Vaginulacea, Vaginulidae, Vaginulinae]BR05
              |    |--*V. taunaisii Férussac 1821BR17
              |    |--V. natalensisG79b
              |    |--V. (Latipes) occidentalis (Guilding 1825)J68
              |    `--V. sloaniiG79a
              |    |--V. andersoniana Tapparone Canefri 1889TC89
              |    |--V. birmanica Theobald 1864TC89
              |    `--V. proxima Tapparone Canefri 1889TC89
              |--Laevicaulis Simroth 1913 (see below for synonymy)SS10
              |    |--*L. comorensis (Fischer 1883) [=Vaginula comorensis]SS10
              |    |--L. alte (Férussac 1821) [incl. Vaginula leydigi Simroth 1889]SS10
              |    |--‘Vaginula’ frauenfeldi Semper 1885 [=*Meisenheimeria frauenfeldi]BR17
              |    `--L. schnitzleriPB27
              `--Leidyula Baker 1925BP90
                   |--L. floridanaBP90
                   |--L. kraussiBP90
                   |--L. moreletiBP90
                   `--L. sloaneiBP90

Laevicaulis Simroth 1913 [incl. Eleutherocaulis Simroth 1913, Meisenheimeria Grimpe & Hoffmann 1924; Meisenheimeriinae]SS10

Onchidella Gray 1850BR05 [=Oncidiella Crosse & Fischer 1878BR05; incl. Occidentella Hoffmann 1929KM17; Onchidellidae, Onchidelloidea, Oncidiellidae]

Onchidiidae [Abranchiatae, Amphipneustea, Dendrobranchiatae, Onchidia, Onchidiacea, Onchidiida, Onchidiiformes, Onchidiiformii, Onchidioidei, Onchidioinei, Onchodoridae, Oncidiacea, Oncidiadae, Opisthotremata, Silicodermatae]BR05

Systellommatophora [Clasthurethra, Digonophora, Ditremata, Gymnomorpha, Gymnophila, Mesommatophora, Soleolifera, Teletremata]BR17

*Type species of generic name indicated


[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.

[BP90] Burch, J. B., & T. A. Pearce. 1990. Terrestrial Gastropoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 201–309. John Wiley & Sones: New York.

[C64] Carpenter, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 33: 517–686.

[G79a] Gibbons, J. S. 1879a. Notes on the habits and distribution, &c., of certain W. Indian Pulmonifera. Journal of Conchology 2: 129–134.

[G79b] Gibbons, J. S. 1879b. Descriptions of two new species of land shells, and remarks on others collected on the East African coast. Journal of Conchology 2: 138–145.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[J68] Jacobson, M. K. 1968. On a collection of terrestrial mollusks from Nicaragua. Nautilus 81 (4): 114–119.

[JG19] James, D. J., P. T. Green, W. F. Humphreys & J. C. Z. Woinarski. 2019. Endemic species of Christmas Island, Indian Ocean. Records of the Western Australian Museum 34 (2): 55–114.

[KM17] Korshunova, T., A. Martynov, T. Bakken, J. Evertsen, K. Fletcher, I. W. Mudianta, H. Saito, K. Lundin, M. Schrödl & B. Picton. 2017. Corrigenda: Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda). ZooKeys 725: 139–141.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 1. Stanford University Press: Stanford University (California).

[PB27] Pilsbry, H. A., & J. Bequaert. 1927. The aquatic mollusks of the Belgian Congo, with a geographical and ecological account of Congo malacology. Bulletin of the American Museum of Natural History 53 (2): 69–602, pls 10–77.

[PP78] Poorman, F. L., & L. H. Poorman. 1978. Additional molluscan records from Bahía de Los Angeles, Baja California Norte. Veliger 20 (4): 369–374.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[R64] Ross, L. T. 1964. The land mollusks of Siesta Key, Sarasota County, Florida. Nautilus 78 (2): 50–52.

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

[TC89] Tapparone Canefri, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XVIII.—Molluschi terrestri e d’acqua dolce. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 295–359.

[TMT96] Tillier, S., M. Masselot & A. Tillier. 1996. Phylogenetic relationships of the pulmonate gastropods from rRNA sequences, and tempo and age of the stylommatophoran radiation. In: Taylor, J. D. (ed.) Origin and Evolutionary Radiation of the Mollusca pp. 267–284. Oxford University Press: Oxford.

[W78] Wells, F. E. 1978. Zonation of marine gastropods on a rocky intertidal shore in the Admiralty Gulf, Western Australia, with emphasis on the genus Nerita. Veliger 20 (3): 279–287.

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