Camaenidae
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 .
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 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 .
In this family, the number of haploid chromosomes lies between 26 and 30 (according to the values in this table).
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; 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:
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:
Genera
The following genera are accepted in the Camaenidae:
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 [ link removed ]
↑ 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 .