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 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).

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: As of March 2023, WoRMS accepts four 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]


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

Genera
Currently, the following genera are accepted within the family Camaenidae:

Taxa with main occurrence in South-East Asia
 * Camaena Albers, 1850 – type genus of the family Camaenidae
 * Amphicoelina Haas, 1933
 * 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
 * Jiaoliaous Zhang, 2024
 * Karaftohelix Pilsbry, 1927
 * Moellendorffia
 * Neocepolis Pilsbry, 1891
 * Obba
 * Parachloritis Ehrmann, 1912
 * Pfeifferia Gray, 1853
 * Platypetasus
 * 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, 2010
 * Amplirhagada Iredale, 1933
 * 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, 2009
 * 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, 2011
 * 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, 2009
 * 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, 2008
 * 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, 2012