Camaenidae

Camaenidae is a family of air-breathing land snails, terrestrial pulmonate gastropod mollusks in the superfamily Helicoidea, the typical snails and their allies. This is one of the most diverse families in the clade Stylommatophora.

These snails occur in a wide variety of habitats in the tropics of Eastern Asia and Australasia.

A large American group, which is mainly represented by species from the Caribbean, has, until recently, also been subsumed under the Camaenidae. However, latest molecular phylogenetic studies showed that these species represent a different family, the Pleurodontidae.

This molecular study also implies that the Bradybaeninae, previously being treated as a distinct family within the Helicoidea, is a junior synonym of the Camaenidae.

Anatomy

Camaenid shells are often quite large (25–50 mm), but a number of species also have small shells (<5 mm). Shells reveal a remarkable diversity in shape and colour, which is partly linked with their lifestyle. For instance, arboreal species tend to have large and conical shells, whereas terrestrial species often have rather flat shells. The shells of some taxa can be vividly coloured, showing banding or other conspicuous patterns, but others are plain and uniform.

This family is defined by a missing diverticulum and a missing stimulatory organ. It is suggested that the family Camaenidae as currently delimited is a polyphyletic taxon. There are no synapomorphies uniting this diverse family. The American group is closely related to the families Helicidae and Helminthoglyptidae, while the Australasian group is a closely related to the Bradybaenidae.

In order to retain the Camaenidae as a monophyletic clade, the Neotropical Pleurodontidae will need to be removed as an independent family, and the Bradybaenidae will need to be included. This taxonomic decision is currently pending a formal suggestion and wider acceptance among systematists, however.

In this family, the number of haploid chromosomes lies between 26 and 30 (according to the values in this table).Barker G. M.: Gastropods on Land: ''Phylogeny, Diversity and Adaptive Morphology''. in Barker G. M. (ed.):
The biology of terrestrial molluscs
'. CABI Publishing, Oxon, UK, 2001, . 1-146, cited pages: 139 and 142.

Subfamilies

Division into subfamilies has been suggested, however, given the unresolved relationships on the family level, the subfamilial treatments must be considered hypothetical. They do not reflect the results of comprehensive phylogenetic analyses, and are not corroborated by current molecular data.

The following three subfamilies have been recognized in the taxonomy of Bouchet & Rocroi (2005) (as based on a suggestion of Alan Solem)
*subfamily Camaeninae Pilbry, 1895 – synonyms: Amphidrominae Kobelt, 1902; Hadridae Iredale, 1937; Xanthomelontidae Iredale, 1937; Chloritidae Iredale, 1938; Papuinidae Iredale, 1938; Calyciidae Iredale, 1941; Planispiridae Iredale, 1941; Cristovalinae Schileyko, 2003
*subfamily Rhagadinae Iredale, 1938
*subfamily Sinumeloninae Solem, 1992

A different taxonomy of the Caemenidae was used by Schileyko (1998–2003).

The new taxonomy of the gastropods, published in 2017 and accepted by WoRMS, gives the following subfamilies:Revised Classification, Nomenclator and Typification of Gastropod and Monoplacophoran Families; Malacologia 61(1-2):1-526. 2017
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* Camaeninae Pilsbry, 1895 Amphidrominae Kobelt, 1902* Bradybaeninae Pilsbry, 1934 (1898)
** tribe Bradybaenini Pilsbry, 1934 (1898) Eulotidae Möllendorff, 1898; = Fruticicolinae Kobelt, 1904; = Buliminopsinae Hoffmann, 1928** tribe Aegistini Kuroda & Habe, 1949
** tribe Euhadrini Habe, Okutani & Nishiwaki, 1994
* Hadrinae Iredale, 1937 [= Xanthomelontidae Iredale, 1937; = Rhagadidae Iredale, 1938; = Chloritidae Iredale, 1938; = Papuinidae Iredale, 1938; = Calyciidae Iredale, 1941; = Planispiridae Iredale, 1941; = Sinumeloninae Solem, 1992; = Cristovalinae Schileyko, 2003]
* Helicostylinae Ihering, 1909378 [= Pfeifferiinae Gray, 1855; = Cochlostylidae Möllendorff, 1890]

Genera

Currently, the following genera are accepted within the family Camaenidae:Köhler F. (2010). "Three new species and two new genera of land snails from the Bonaparte Archipelago in the Kimberley, Western Australia (Pulmonata, Camaenidae)". ''Molluscan Research'' 30(1): 1-16.

Taxa with main occurrence in South-East Asia
* '' Camaena'' Albers, 1850 – type genus of the family Camaenidae
* '' Amphicoelina'' Haas, 1933Páll-Gergely B. & Asami T. (2014). "Additional information on the distribution, anatomy and systematics of living and fossil Chinese Plectopylidae (Gastropoda: Pulmonata)". ''Genus'' 25(3): 527-564
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* ''Amphidromus'' Albers, 1850
* '' Bouchetcamaena'' Thach, 2018 (debated synonym of ''Trichochloritis'' Pilsbry, 1891)
* '' Chloritis'' Beck, 1837
* '' Entadella'' Páll-Gergely & Hunyadi, 2016
* '' Ganesella'' W. T. Blanford, 1863
* '' Globotrochus'' F. Haas, 1935
* '' Karaftohelix'' Pilsbry, 1927
* '' Moellendorffia''
* ''Neocepolis'' Pilsbry, 1891
* '' Obba''
* '' Parachloritis'' Ehrmann, 1912
* '' Pfeifferia'' Gray, 1853
* '' Philbouchetia'' Thach, 2020
* ''Plectotropis'' E. von Martens, 1860
* ''Pseudobuliminus'' Gredler, 1886
* '' Pseudopartula'' L. Pfeiffer, 1856
* ''Satsuma''
* '' Sinorachis'' M. Wu & Z.-Y. Chen, 2019
* '' Stegodera''
* '' Trachia''
* '' Trachystyla''
* ''Trichochloritis'' Pilsbry, 1891

Taxa with main occurrence in Papua New Guinea to Solomon Islands
* '' Albersia''
* '' Calycia'' H. Adams, 1865
* '' Canefriula''
* Chloraea Albers, 1850
* ''Cryptaegis''
* '' Crystallopsis''
* '' Forcartia''
* '' Mecyntera''
* '' Megalacron'' I. Rensch, 1934
* '' Meliobba''
* '' Papuanella''
* '' Papuexul'' Iredale, 1933
* ''Papuina'' Martens, 1860
* '' Papustyla''
* ''Planispira'' Beck, 1837
* '' Quirosena'' Iredale, 1941
* '' Rhynchotrochus'' Möllendorff, 1895
* '' Rhytidoconcha''
* '' Smeatonia''

Australian genera In Australia, the Camaenidae comprise 131 currently recognized genera, most of which are endemic to the continent.

* '' Adclarkia'' Stanisic, 1996
* '' Aetholitis'' Stanisic, 2010Stanisic, J., Shea, M., Potter, D. and Griffiths, O. 2010. Australian Land Snails – Volume 1: A field guide to eastern Australian species. Bioculture Press, Mauritius, 591 pp.
* '' Amplirhagada'' Iredale, 1933Iredale, T. 1933. Systematic notes on Australian land shells. Records of the Australian Museum 19: 37-59
* '' Arnemelassa'' Iredale, 1938
* '' Aslintesta'' Solem, 1992
* '' Australocosmica'' Köhler, 2011
* '' Austrocomaena'' Stanisic, 2010
* '' Austrochloritis'' Pilsbry, 1890
* ''Basedowena'' Iredale, 1937
* ''Baudinella'' Thiele, 1931
* '' Bentosites'' Iredale, 1933
* '' Billordia'' Stanisic, 2010
* '' Boriogenia'' Stanisic, 2010
* '' Calvigenia'' Iredale, 1938
* '' Caperantrum'' Solem, 1992

* '' Carinotrachia'' Solem, 1985
* '' Chloritisanax'' Iredale, 1933
* '' Contramelon'' Iredale, 1937
* '' Cooperconcha'' Solem, 1997
* ''Crikey'' Stanisic, 2009 – with one species ''Crikey steveirwini'' Stanisic, 2009Stanisic J. (24 August) 2009. ''Crikey steveirwini gen. et sp. nov. from montane habitats in the Wet Tropics of northeastern Queensland, Australia (Gastropoda: Eupulmonata: Camaenidae)''. Zootaxa 2206: 62–68
abstract

* '' Cristigibba'' Tapparone-Canefri, 1883
* '' Cristilabrum'' Solem, 1981
* '' Crookshanksia'' Stanisic, 2010
* '' Cupedora'' Iredale, 1933
* '' Damochlora'' Iredale, 1938
* '' Dirutrachia'' Iredale, 1937
* '' Discomelon'' Iredale, 1938
* '' Divellomelon'' Iredale, 1933
* '' Eurytrachia'' Stanisic, 2010
* '' Exiligada'' Iredale, 1939
* '' Eximiorhagada'' Iredale, 1938
* '' Falspleuroxia'' Solem, 1997
* '' Figuladra'' Iredale, 1938
* '' Forrestena'' Stanisic, 2010
* '' Galadistes'' Iredale, 1938
* '' Gloreugenia'' Iredale, 1933
* '' Globorhagada'' Iredale, 1933
* '' Glyptorhagada'' Pilsbry, 1890
* '' Gnarosophia'' Iredale, 1933
* '' Granulomelon'' Iredale, 1933
* '' Hadra'' Albers, 1860
* ''Jacksonena'' Iredale, 1937
* '' Jimbouria'' Stanisic, 2010
* '' Kandochloritis'' Shea & Griffiths, 2010
* '' Kendrickia'' Solem, 1985
* ''Kimberleydiscus'' Köhler, 2010 – with one species '' Kimberleydiscus fasciatus'' Köhler, 2010
* '' Kimberleymelon'' Köhler, 2010 – with one species ''Kimberleymelon tealei'' Köhler, 2010
* '' Kimberleytrachia'' Köhler, 2011Köhler, F. 2011. The camaenid species of the Kimberley Islands, Western Australia (Stylommatophora: Helicoidea). Malacologia, 54(1-2): 203–406
* '' Kimboraga'' Iredale, 1939
* '' Lacustrelix'' Iredale, 1933
* '' Lamprellia'' Stanisic, 2010
* '' Lynfergusonia'' Stanisic, 2010
* '' Marylnessa'' Stanisic, 2010
* '' Meliobba'' Iredale, 1940
* '' Melostrachia'' Solem, 1979
* '' Meridistis'' Stanisic, 2010
* '' Meridolum'' Iredale, 1942
* '' Mesodontrachia'' Solem, 1985
* '' Micromelon'' Solem, 1992
* '' Minimelon'' Solem, 1993
* '' Molema'' Köhler, 2011
* '' Montanomelon'' Solem, 1993
* '' Monteithosites'' Stanisic, 1996
* '' Moretonistes'' Stanisic, 2010
* ''Mouldingia'' Solem, 1984
* '' Mussonena'' Iredale, 1938
* '' Nannochloritis'' Iredale, 1938
* '' Neveritis'' Iredale, 1938
* '' Ningbingia'' Solem, 1981
* ''Noctepuna'' Iredale, 1933
* '' Obstengenia'' Iredale, 1933
* '' Offachloritis'' Iredale, 1933
* '' Ordtrachia'' Solem, 1984
* '' Pallidelix'' Iredale, 1933
* '' Papuexul'' Iredale, 1933
* '' Parglogenia'' Iredale, 1938
* '' Patrubella'' Iredale, 1938
* '' Perioinsolita'' Stanisic & Potter, 2010
* '' Petraconcha'' Clark, 2009Clark, S. 2009. A review of the land snail genus Meridolum (Gastropoda: Camaenidae) from central New South Wales, Australia. ''Molluscan Research'', 29:61-120.
* '' Plectorhagada'' Iredale, 1933
* ''Pleuroxia'' Ancey, 1887
* '' Pommerhelix'' Clark, 2009
* '' Ponderconcha'' Clark, 2009
* '' Posorites'' Iredale, 1933
* '' Promonturconchum'' Solem, 1997
* '' Protolinitis'' Stanisic, 2010
* ''Prototrachia'' Solem, 1984
* '' Prymnbriareus'' Solem, 1981
* '' Pseudcupedora'' Solem, 1992
* '' Quistrachia'' Iredale, 1939
* '' Ramogenia'' Iredale, 1938
* '' Rhagada'' Albers, 1860 – type genus of the subfamily Rhagadinae
* '' Retroterra'' Solem, 1985
* '' Sauroconcha'' Zhang & Shea, 2008Zhang, W.-H. & Shea, M. (2008). A new genus and species of land snail of the family Camaenidae from New South Wales. ''Molluscan Research'', 28: 123-132.
* '' Semotrachia'' Iredale, 1933
* '' Setobaudinia'' Iredale, 1933
* '' Sinumelon'' Iredale, 1930 – type genus of the subfamily Sinumeloninae
* '' Sphaerospira'' Mörch, 1867
* '' Spurlingia'' Iredale, 1933
* '' Squamagenia'' Stanisic & Potter, 2010
* '' Stanisicia'' Clark, 2009
* '' Strepsitaurus'' Solem, 1997
* '' Tatemelon'' Solem, 1993
* '' Temporena'' Iredale, 1933
* ''Thersites'' Pfeiffer, 1855
* '' Tolgachloritis'' Iredale, 1933
* '' Toombatrachia'' Stanisic, 2010
* '' Torresitrachia'' Iredale, 1939
* '' Trachiopsis'' Pilsbry, 1893
* '' Trozena'' Iredale, 1938
* '' Turgenitubulus'' Solem, 1981
* '' Varohadra'' Iredale, 1933
* '' Ventopelita'' Iredale, 1943
* '' Vidumelon'' Iredale, 1933
* '' Westraltrachia'' Iredale, 1933
* '' Xanthomelon'' Martens, 1860
* '' Youwanjela'' Köhler & Shea, 2012Köhler, F. & Shea, M. (2012). ''Youwanjela'', a new genus of land snail from the Kimberley, Western Australia (Eupulmonata, Camaenidae). Zoosystematics and Evolution, 88: 25-31.

References

Further reading

* Köhler F. (2009) "Phylogeny and evolution of the Camaenidae in north-western Australia: A model case for the study of speciation and radiation". In: McDoughall C. & Hall N. (Eds.) ''Molluscs 2009: Program and abstracts. Malacological Society of Australasia'', Brisbane, p. 55.
* Wade C. M., Hudelot C., Davison A. Naggs, F. & Mordan P. B. (2007). "Molecular phylogeny of the helicoid land snails (Pulmonata: Stylommatophora: Helicoidea), with special emphasis on the Camaenidae". ''Journal of Molluscan Studies'' 73(4): 411–415. .

External links

Australian Camaenidae

{{Taxonbar, from=Q927331

Helicoidea
Gastropod genera