Camaenidae

{{Short description|Family of gastropods}} {{Use dmy dates|date=December 2022}} {{Automatic taxobox | image = Terrestrial molluscs from Laos (10.3897-zookeys.834.28800) Figure 58.jpg | image_caption = | taxon = Camaenidae | authority = Pilsbry, 1895<ref>Pilsbry H. A. (1895). ''Manual of Conchology, structural and systematic, with illustrations of the species. Second series: Pulmonata.'' Helicidae – Volume VII. '''9'''(33a): [https://archive.org/stream/manualofconcholo09tryorich#page/n41/mode/1up xxxii].</ref> | synonyms_ref = | synonyms = * Bradybaenidae Pilsbry, 1934 (1898) * Crassispirinae McLean, 1971 * Zonulispirinae McLean, 1971 | type_genus = ''Camaena'' | type_genus_authority = Albers, 1850 | subdivision_ranks = Genera | subdivision = See text }}

'''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.<ref>Cuezzo M. G. (2003). "Phylogenetic analysis of the Camaenidae (Mollusca: Stylommatophora) with special emphasis on the American taxa". ''Zoological Journal of the Linnean Society'' '''138'''(4): 449-476. {{doi|10.1046/j.1096-3642.2003.00061.x}}.</ref>

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.<ref>Wade, C.M., Hudelot, C., Davison, A., Naggs, F., Mordan, P.B. Molecular phylogeny of the helicoid land snails (Pulmonata: Stylommatophora: Helicoidea), with special emphasis on the Camaenidae. ''Journal of Molluscan Studies'' '''73''': 411-415.</ref><ref>Bouchet, P., Rocroi, J.P. Classification and Nomenclator of gastropod families. ''Malacologia'' '''47''': 1-397.</ref>

This molecular study also implies that the Bradybaeninae, previously 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 stimulatory organ. There are no synapomorphies uniting this diverse family. The Australasian group is a closely related to the Bradybaeninae.<ref>Scott B. 1996. ''Phylogenetic relationships of the Camaenidae (Pulmonata: Stylommatophora: Helicoidea)''. Journal of Molluscan Studies, 62: 65-73. [https://archive.today/20120713181230/http://mollus.oxfordjournals.org/cgi/content/abstract/62/1/65 Abstract]</ref>

In this family, the number of haploid chromosomes lies between 26 and 30 (according to the values in this table).<ref name="Barker">Barker G. M.: Gastropods on Land: ''Phylogeny, Diversity and Adaptive Morphology''. in Barker G. M. (ed.): ''[https://books.google.com/books?id=WlvX-9Wt0toC&hl The biology of terrestrial molluscs]''. CABI Publishing, Oxon, UK, 2001, {{ISBN|0-85199-318-4}}. 1-146, cited pages: 139 and 142.</ref>

== Taxonomy == 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;<ref>Kobelt W. (1902). ''Systematisches Conchilien-Cabinet'', ed. 2, Bd. 1, Abt 13, Theil 2: 1033.</ref> 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<ref>Iredale T. (1938). The Australian Zoologist '''9'''(2): 112.</ref> *subfamily Sinumeloninae Solem, 1992<ref>Solem A. (1992). Records of the South Australian Museum, Monograph series '''2''': 161.</ref>

A different taxonomy of the Caemenidae was used by Schileyko (1998–2003).<ref>Schileyko A. A. (1998-2003). Treatise on Recent terrestrial pulmonate molluscs. ''Ruthenica'' supplement 2.</ref>

The new taxonomy of the gastropods, published in 2017 and accepted by WoRMS, gives the following subfamilies:<ref name="Bouchet 2017">[http://www.bioone.org/doi/10.4002/040.061.0201 Revised Classification, Nomenclator and Typification of Gastropod and Monoplacophoran Families; Malacologia 61(1-2):1-526. 2017]</ref> * 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] As of March 2023, WoRMS accepts four subfamilies:<ref name = worms>{{Cite web |title=WoRMS - World Register of Marine Species - ''Camaenidae'' Pilsbry, 1895 |url=https://www.marinespecies.org/aphia.php?p=taxdetails&id=762708 |access-date=2023-03-15 |website=www.marinespecies.org}}</ref>

* Bradybaeninae Pilsbry, 1895 * Camaeninae Iredale, 1937 * Hadrinae Iredale, 1937 * Helicostylinae Ihering, 1909 (1890)

===Genera=== The following genera are accepted in the Camaenidae:<ref name=worms />

Genera not placed in a subfamily: * ''Indocamaena'' Thach, 2023 * ''Kenyirus'' Clements & Tan, 2012 * ''Ogeramua'' Christensen, 2017 * ''Philbouchetia'' Thach, 2020 * ''Sinocamaena'' M. Wu, 2024

Subfamily Bradybaeninae Pilsbry, 1934 (1898): {{div col|colwidth=22em}} *Tribe Aegistini Kuroda & Habe, 1949 **''Aegista'' Albers, 1850 **''Aegistohadra'' M. Wu, 2004 **''Coelorus'' Pilsbry, 1900 **''Dolicheulota'' Pilsbry, 1901 **''Guamampa'' Schileyko, 1997 **''Landouria'' Godwin-Austen, 1918 **''Lepidopisum'' Kuroda & Habe, 1958 **''Mandarina'' Pilsbry, 1895 **''Mastigeulota'' Pilsbry, 1895 **''Miyakoia'' Minato, 1980 **''Neoaegista'' Azuma, 1955 **''Neochloritis'' Minato, 1982 **''Nesiohelix'' Kuroda & Emura, 1943 **''Nipponochloritis'' Habe, 1955 **''Pancala'' Kuroda & Habe, 1949 **''Plecteulota'' Möllendorff, 1892 **''Plectotropis'' E. von Martens, 1860 **''Pseudaspasita'' Möllendorff, 1901 **''Satsuma'' A. Adams, 1868 **''Takasagohadra'' Kuroda, 1941 **''Torobaena'' F. Haas, 1935 **''Tricheulota'' Pilsbry, 1895 **''Trishoplita'' Jacobi, 1898 **''Yakuchloritis'' Habe, 1955

*Tribe Euhadrini Habe, Okutani & Nishiwaki, 1994 **''Euhadra'' Pilsbry, 1890

*Tribe Bradybaenini Pilsbry, 1934 **''Acusta'' E. von Martens, 1860 **''Ainohelix'' Kuroda & Taki, 1933 **''Apatetes'' Gude, 1914 **''Armandiella'' Ancey, 1901 **''Bradybaena'' H. Beck, 1837 **''Buliminidius'' Heude, 1890 **''Cathaica'' Moellendorff, 1884 **''Chosenelix'' Pilsbry, 1927 **''Coccoglypta'' Pilsbry, 1895 **''Ezohelix'' Kuroda & Emura, 1938 **''Fruticicola'' Held, 1838 **''Grabauia'' Yen, 1935 **''Karaftohelix'' Pilsbry, 1927 **''Kugitangia'' Schileyko, Pazilov & Abdulazizova, 2017 **''Laeocathaica'' Moellendorff, 1899 **''Metodontia'' Möllendorff, 1886 **''Mikiria'' Godwin-Austen, 1918 **''Neofruticicola'' Schileyko, Pazilov & Abdulazizova, 2020 **''Neseulota'' Ehrmann, 1911 **''Paraegista'' Kuroda & Azuma, 1951 **''Phaeohelix'' Kuroda & Habe, 1949 **''Platypetasus'' Pilsbry, 1895 **''Pliocathaica'' Andreae, 1900 **''Ponsadenia'' Schileyko, 1978 **''Pseudiberus'' Ancey, 1887 **''Pseudobuliminus'' Gredler, 1886 **''Rudens'' Heude, 1890 **''Secusana'' Gredler, 1894 **''Semibuliminus'' Möllendorff, 1899 **''Stenogyropsis'' Möllendorff, 1899 **''Stilpnodiscus'' Möllendorff, 1899 **''Trichobradybaena'' M. Wu & J.-Y. Guo, 2003 **''Trichocathaica'' Gude, 1919

*Genera not placed to tribe: **''Eueuhadra'' M. Wu, 2004 **''Jiaoliaous'' G. Zhang, 2024 **''Sinochloritis'' M. Wu & Z.-Y. Chen, 2019 **''Sinorachis'' M. Wu & Z.-Y. Chen, 2019 **''Traumatophora'' Ancey, 1887 {{div col end}}

Subfamily Camaeninae: {{div col|colwidth=22em}} * ''Albersia'' H. Adams, 1865 * ''Amphicoelina'' Zilch, 1960 * ''Amphidromus'' Albers, 1850 * ''Anceyoconcha'' S. Tumpeesuwan & C. Tumpeesuwan, 2020 * ''Beddomea'' G. Nevill, 1878 * ''Bellatrachia'' Schileyko, 2018 * ''Bouchetcamaena'' Thach, 2018 * ''Burmochloritis'' Godwin-Austen, 1920 * ''Camaena'' Albers, 1850 * ''Camaenella'' Pilsbry, 1893 * ''Cryptaegis'' W. F. Clapp, 1923 * ''Dentichloritis'' Páll-Gergely & Neubert, 2019 * ''Discoconcha'' I. Rensch, 1935 * ''Disteustoma'' Iredale, 1941 * ''Dorcasidea'' Iredale, 1941 * ''Entadella'' Páll-Gergely & Hunyadi, 2016 * ''Ganesella'' W. T. Blanford, 1863 * ''Gemitelix'' Iredale, 1941 * ''Globotrochus'' F. Haas, 1935 * ''Kendallena'' Iredale, 1941 * ''Mecyntera'' Iredale, 1941 * ''Moellendorffia'' Ancey, 1887 * ''Moellendorffiella'' Pilsbry, 1905 * ''Neocepolis'' Pilsbry, 1891 * ''Neotrachia'' Schileyko, 2018 * ''Obba'' H. Beck, 1837 * ''Obbiberus'' F. Haas, 1935 * ''Oreobba'' Pilsbry, 1894 * ''Pseudobba'' Möllendorff, 1891 * ''Pseudopartula'' L. Pfeiffer, 1856 * ''Pseudostegodera'' M. Wu & Z.-Y. Chen, 2021 * ''Pseudotrachia'' Schileyko, 2018 * ''Rhytidoconcha'' I. Rensch, 1933 * ''Stegodera'' E. von Martens, 1876 * ''Trachia'' E. von Martens, 1860 * ''Trachychloritis'' F. Haas, 1934 * ''Tradeustoma'' Iredale, 1941 * ''Trichelix'' Ancey, 1887 * ''Trichochloritis'' Pilsbry, 1891 * ''Vulnus'' Sykes, 1904 {{div col end}}

Subfamily Hadrinae: {{div col|colwidth=22em}} * ''Adclarkia'' J. Stanisic, 1996 * ''Aetholitis'' J. Stanisic, 2010 * ''Amplirhagada'' Iredale, 1933 * ''Ariophantopsis'' B. Rensch, 1930 * ''Arnemelassa'' Köhler & Bouchet, 2020 * ''Arnhemtrachia'' Köhler & Criscione, 2013 * ''Aslintesta'' Solem, 1992 * ''Australocosmica'' Köhler, 2011 * ''Austrocamaena'' J. Stanisic, 2010 * ''Austrochloritis'' Pilsbry, 1891 * ''Basedowena'' Iredale, 1937 * ''Baudinella'' Thiele, 1931 * ''Bentosites'' Iredale, 1933 * ''Billordia'' J. Stanisic, 2010 * ''Boriogenia'' J. Stanisic, 2010 * ''Brigaladra'' L. Stanisic, 2022 * ''Calvigenia'' Iredale, 1938 * ''Calycia'' H. Adams, 1865 * ''Canefriula'' Iredale, 1941 * ''Caperantrum'' Solem, 1997 * ''Cardiotrachia'' Criscione & Köhler, 2014 * ''Carinotrachia'' Solem, 1985 * ''Catellotrachia'' Iredale, 1933 * ''Chloritis'' H. Beck, 1837 * ''Chloritisanax'' Iredale, 1933 * ''Chloritobadistes'' Iredale, 1933 * ''Claudettea'' Iredale, 1941 * ''Coliolus'' Tapparone Canefri, 1886 * ''Contramelon'' Iredale, 1937 * ''Cooperconcha'' Solem, 1992 * ''Crikey'' J. Stanisic, 2009 * ''Cristigibba'' Tapparone Canefri, 1883 * ''Cristilabrum'' Solem, 1981 * ''Crookshanksia'' J. Stanisic, 2010 * ''Crystallopsis'' Ancey, 1887 * ''Cupedora'' Iredale, 1937 * ''Damochlora'' Iredale, 1938 * ''Denhamiana'' J. Stanisic, 2013 * ''Dirutrachia'' Iredale, 1937 * ''Discomelon'' Iredale, 1938 * ''Divellomelon'' Iredale, 1933 * ''Euryladra'' L. Stanisic, 2022 * ''Eurytrachia'' J. Stanisic, 2010 * ''Exiligada'' Iredale, 1939 * ''Falspleuroxia'' Solem, 1997 * ''Figuladra'' Köhler & Bouchet, 2020 * ''Forcartia'' Clench & R. D. Turner, 1963 * ''Forrestena'' J. Stanisic, 2010 * ''Galadistes'' Iredale, 1938 * ''Globorhagada'' Iredale, 1933 * ''Gloreugenia'' Iredale, 1933 * ''Glyptorhagada'' Pilsbry, 1890 * ''Gnarosophia'' Iredale, 1933 * ''Granulomelon'' Iredale, 1933 * ''Hadra'' E. von Martens, 1860 * ''Jacksonena'' Iredale, 1937 * ''Jimbouria'' J. Stanisic, 2010 * ''Kandoschloritis'' Shea & O. L. Griffiths, 2010 * ''Kendrickia'' Solem, 1985 * ''Kimberleydiscus'' Köhler, 2010 * ''Kimberleymelon'' Köhler, 2010 * ''Kimberleytrachia'' Köhler, 2011 * ''Kimboraga'' Iredale, 1939 * ''Kymatobaudinia'' Criscione & Köhler, 2013 * ''Lacustrelix'' Iredale, 1937 * ''Lamprellia'' J. Stanisic, 2010 * ''Letitia'' Iredale, 1941 * ''Lorelliana'' J. Stanisic, 2016 * ''Lynfergusonia'' J. Stanisic, 2010 * ''Marilynessa'' J. Stanisic, 2010 * ''Megalacron'' I. Rensch, 1934 * ''Meliobba'' Iredale, 1940 * ''Melostrachia'' Iredale, 1938 * ''Meridistes'' J. Stanisic, 2010 * ''Meridolum'' Iredale, 1933 * ''Mesodontrachia'' Solem, 1985 * ''Micromelon'' Solem, 1992 * ''Molema'' Köhler, 2011 * ''Montanomelon'' Solem, 1993 * ''Monteithosites'' J. Stanisic, 1996 * ''Moretonistes'' J. Stanisic, 2010 * ''Mouldingia'' Solem, 1984 * ''Mussonena'' Iredale, 1938 * ''Nannochlora'' Criscione & Köhler, 2015 * ''Nannochloritis'' Köhler & Bouchet, 2020 * ''Nanotrachia'' Köhler & Criscione, 2013 * ''Neveritis'' Iredale, 1938 * ''Ningbingia'' Solem, 1981 * ''Noctepuna'' Iredale, 1933 * ''Nodulabium'' Criscione & Köhler, 2013 * ''Obsteugenia'' Iredale, 1933 * ''Offachloritis'' Iredale, 1933 * ''Ordtrachia'' Solem, 1984 * ''Ototrachia'' Criscione & Köhler, 2013 * ''Pallidelix'' Iredale, 1933 * ''Papuexul'' Iredale, 1933 * ''Papuina'' E. von Martens, 1860 * ''Papuolus'' Schileyko, 2003 * ''Papustyla'' Pilsbry, 1893 * ''Parachloritis'' Ehrmann, 1912 * ''Parglogenia'' Iredale, 1938 * ''Patrubella'' Iredale, 1938 * ''Perioinsolita'' J. Stanisic & Potter, 2010 * ''Petraconcha'' S. A. Clark, 2009 * ''Planispira'' H. Beck, 1837 * ''Plectorhagada'' Iredale, 1933 * ''Pleuroxia'' Ancey, 1887 * ''Pommerhelix'' S. A. Clark, 2009 * ''Ponderconcha'' S. A. Clark, 2009 * ''Posorites'' Iredale, 1933 * ''Promonturconchum'' Solem, 1997 * ''Protolinitis'' J. Stanisic, 2010 * ''Prototrachia'' Solem, 1984 * ''Prymnbriareus'' Solem, 1981 * ''Pseudcupedora'' Solem, 1992 * ''Pseudodistes'' J. Stanisic & Potter, 2010 * ''Pseudomesodontrachia'' Criscione & Köhler, 2013 * ''Pseudopapuina'' F. Haas, 1934 * ''Quirosena'' Iredale, 1941 * ''Quistrachia'' Iredale, 1939 * ''Rachita'' Criscione & Köhler, 2014 * ''Ramogenia'' Iredale, 1938 * ''Retroterra'' Solem, 1985 * ''Rhagada'' E. von Martens, 1860 * ''Rhynchotrochus'' Möllendorff, 1895 * ''Sauroconcha'' W.-H. Zhang & Shea, 2008 * ''Setobaudinia'' Iredale, 1933 * ''Setocallosa'' Criscione & Köhler, 2015 * ''Sinumelon'' Iredale, 1930 * ''Solmogada'' Iredale, 1941 * ''Sphaerospira'' Mörch, 1867 * ''Spurlingia'' Iredale, 1933 * ''Squamagenia'' J. Stanisic & Potter, 2010 * ''Stanisicia'' S. A. Clark, 2009 * ''Steorra'' J. Stanisic, 2010 * ''Strepsitaurus'' Solem, 1997 * ''Succochlea'' Criscione & Köhler, 2014 * ''Sulcobasis'' Tapparone Canefri, 1883 * ''Tatemelon'' Solem, 1993 * ''Temporena'' Iredale, 1933 * ''Thersites'' L. Pfeiffer, 1855 * ''Tolgachloritis'' Iredale, 1933 * ''Toombatrachia'' J. Stanisic, 2010 * ''Torresitrachia'' Iredale, 1939 * ''Trachiopsis'' Pilsbry, 1893 * ''Trachygenia'' J. Stanisic, 2010 * ''Trozena'' Iredale, 1938 * ''Turgenitubulus'' Solem, 1981 * ''Ventopelita'' Iredale, 1933 * ''Vidumelon'' Iredale, 1933 * ''Vincentrachia'' Criscione & Köhler, 2013 * ''Wahgia'' Clench & R. D. Turner, 1959 * ''Westraltrachia'' Iredale, 1933 * ''Xanthomelon'' E. von Martens, 1860 * ''Xeromelon'' Criscione & Köhler, 2016 * ''Youwanjela'' Köhler & Shea, 2012 * ''Zyghelix'' Iredale, 1937 {{div col end}}

Subfamily Helicostylinae: {{div col|colwidth=22em}} * ''Anixa'' Pilsbry, 1895 * ''Calocochlea'' W. Hartmann, 1843 * ''Chloraea'' Albers, 1850 * ''Chrysallis'' Albers, 1850 * ''Cochlodryas'' E. von Martens, 1860 * ''Cochlostyla'' A. Férussac, 1821 * ''Dolichostyla'' Pilsbry, 1896 * ''Helicobulinus'' Broderip, 1841 * ''Helicostyla'' A. Férussac, 1821 * ''Hypselostyla'' L. Pfeiffer, 1868 * ''Leytia'' Pilsbry, 1892 * ''Mesanella'' Clench & R. D. Turner, 1952 * ''Orthostylus'' H. Beck, 1837 * ''Orustia'' Mörch, 1852 * ''Pachya'' Albers, 1850 * ''Pachysphaera'' Pilsbry, 1892 * ''Pfeifferia'' J. E. Gray, 1853 * ''Phengus'' Albers, 1850 * ''Phoenicobius'' Mörch, 1852 * ''Pyrochilus'' Pilsbry, 1893 * ''Trachystyla'' Pilsbry, 1892 {{div col end}}

==References== {{Reflist}}

== 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. {{doi|10.1093/mollus/eym030}}.

==External links== *{{Commons category-inline}} *{{cite web|url=http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/CAMAENIDAE |title=Family Camaenidae|publisher=Australian government|archive-url=https://web.archive.org/web/20121013192108/http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/CAMAENIDAE |archive-date=13 October 2012 }} *{{Cite web |title=Camaenid Land Snails from Western and Central Australia (Mollusca: Pulmonata: Camaenidae) {{!}} Western Australian Museum |url=http://museum.wa.gov.au/research/records-supplements/records/camaenid-land-snails-western-and-central-australia-mollusca-p-0 |access-date=2023-05-13 |website=Western Australian Museum |language=en}} {{Scholia|position=left}}

Category:Camaenidae Category:Helicoidea Category:Gastropod families Category:Taxa named by Henry Augustus Pilsbry