Camaenidae Last updated July 12, 2025  Family of gastropods
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 . [ 2]  
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 . [ 3]   [ 4]  
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 . [ 5]  
In this family, the number of haploid  chromosomes  lies between 26 and 30 (according to the values in this table). [ 6]  
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; [ 7]   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 [ 8]   subfamily Sinumeloninae  Solem, 1992 [ 9]   A different taxonomy of the Caemenidae was used by Schileyko (1998–2003). [ 10]  
The new taxonomy of the gastropods, published in 2017 and accepted by WoRMS , gives the following subfamilies: [ 11]  
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: [ 12]  
Genera The following genera are accepted in the Camaenidae: [ 12]  
Genera not placed in a subfamily:
Subfamily Bradybaeninae   Pilsbry , 1934 (1898) :
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  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:  Subfamily Camaeninae :
 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 Subfamily Hadrinae :
 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 Subfamily Helicostylinae :
 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 References  ↑    Pilsbry H. A.  (1895).  Manual of Conchology , structural and systematic, with illustrations of the species. Second series: Pulmonata.  Helicidae – Volume VII.  9 (33a): xxxii .  ↑   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 .  ↑   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.  ↑   Bouchet, P., Rocroi, J.P. Classification and Nomenclator of gastropod families.  Malacologia   47 : 1-397.  ↑   Scott B. 1996. Phylogenetic relationships of the Camaenidae (Pulmonata: Stylommatophora: Helicoidea) . Journal of Molluscan Studies, 62: 65-73. Abstract    ↑   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, ISBN     0-85199-318-4 . 1-146, cited pages: 139 and 142.  ↑    Kobelt W.  (1902). Systematisches Conchilien-Cabinet , ed. 2, Bd. 1, Abt 13, Theil 2: 1033.  ↑   Iredale T. (1938). The Australian Zoologist 9 (2): 112.  ↑   Solem A. (1992). Records of the South Australian Museum, Monograph series 2 : 161.  ↑   Schileyko A. A. (1998-2003). Treatise on Recent terrestrial pulmonate molluscs. Ruthenica  supplement 2.  ↑    Revised Classification, Nomenclator and Typification of Gastropod and Monoplacophoran Families; Malacologia 61(1-2):1-526. 2017    1  2    "WoRMS - World Register of Marine Species - Camaenidae  Pilsbry, 1895" . www.marinespecies.org . Retrieved 15 March  2023 . 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 . This page is based on this 
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