Family of gastropods
Camaenidae is afamily of air-breathingland snails ,terrestrial pulmonate gastropod mollusks in the superfamilyHelicoidea , the typical snails and their allies. This is one of the most diverse families in theclade Stylommatophora .
These snails occur in a wide variety ofhabitats in thetropics ofEastern Asia andAustralasia .[ 2]
A large American group, which is mainly represented by species from theCaribbean , has, until recently, also been subsumed under the Camaenidae. However, latest molecular phylogenetic studies showed that these species represent a different family, thePleurodontidae .[ 3] [ 4]
This molecular study also implies that theBradybaeninae , previously treated as a distinct family within theHelicoidea , is a juniorsynonym of the Camaenidae.
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 nosynapomorphies uniting this diverse family. The Australasian group is a closely related to theBradybaeninae .[ 5]
In this family, the number ofhaploid chromosomes lies between 26 and 30 (according to the values in this table).[ 6]
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 threesubfamilies have been recognized inthe taxonomy of Bouchet & Rocroi (2005) (as based on a suggestion of Alan Solem)
subfamily Camaeninae Pilbry, 1895 – synonyms: Amphidrominae Kobelt, 1902;[ 7] subfamilyRhagadinae Iredale, 1938[ 8] subfamilySinumeloninae 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 byWoRMS , 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]
The following genera are accepted in the Camaenidae:[ 12]
Genera not placed in a subfamily:
SubfamilyBradybaeninae Pilsbry , 1934 (1898)
TribeAegistini 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 TribeEuhadrini Habe, Okutani & Nishiwaki, 1994 TribeBradybaenini 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: SubfamilyCamaeninae :
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 SubfamilyHadrinae :
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 SubfamilyHelicostylinae :
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 ^ Pilsbry H. A. (1895).Manual of Conchology , structural and systematic, with illustrations of the species. Second series: Pulmonata.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 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 Zoologist9 (2): 112. ^ Solem A. (1992). Records of the South Australian Museum, Monograph series2 : 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 ^a b "WoRMS - World Register of Marine Species -Camaenidae Pilsbry, 1895" .www.marinespecies.org . Retrieved15 March 2023 .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 .
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