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, but a number of species also have small shells. 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.

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)
  • 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.
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:

Genera

The following genera are accepted in the Camaenidae:
Genera not placed in a subfamily:Indocamaena Thach, 2023Kenyirus Clements & Tan, 2012Ogeramua Christensen, 2017Philbouchetia Thach, 2020Sinocamaena M. Wu, 2024
Subfamily Bradybaeninae Pilsbry, 1934 :
Subfamily Camaeninae:Albersia H. Adams, 1865Amphicoelina Zilch, 1960Amphidromus Albers, 1850Anceyoconcha S. Tumpeesuwan & C. Tumpeesuwan, 2020Beddomea G. Nevill, 1878Bellatrachia Schileyko, 2018Bouchetcamaena Thach, 2018Burmochloritis Godwin-Austen, 1920Camaena Albers, 1850Camaenella Pilsbry, 1893Cryptaegis W. F. Clapp, 1923Dentichloritis Páll-Gergely & Neubert, 2019Discoconcha I. Rensch, 1935Disteustoma Iredale, 1941Dorcasidea Iredale, 1941Entadella Páll-Gergely & Hunyadi, 2016Ganesella W. T. Blanford, 1863Gemitelix Iredale, 1941Globotrochus F. Haas, 1935Kendallena Iredale, 1941Mecyntera Iredale, 1941Moellendorffia Ancey, 1887Moellendorffiella Pilsbry, 1905Neocepolis Pilsbry, 1891Neotrachia Schileyko, 2018Obba H. Beck, 1837Obbiberus F. Haas, 1935Oreobba Pilsbry, 1894Pseudobba Möllendorff, 1891Pseudopartula L. Pfeiffer, 1856Pseudostegodera M. Wu & Z.-Y. Chen, 2021Pseudotrachia Schileyko, 2018Rhytidoconcha I. Rensch, 1933Stegodera E. von Martens, 1876Trachia E. von Martens, 1860Trachychloritis F. Haas, 1934Tradeustoma Iredale, 1941Trichelix Ancey, 1887Trichochloritis Pilsbry, 1891Vulnus Sykes, 1904
Subfamily Hadrinae:Adclarkia J. Stanisic, 1996Aetholitis J. Stanisic, 2010Amplirhagada Iredale, 1933Ariophantopsis B. Rensch, 1930Arnemelassa Köhler & Bouchet, 2020Arnhemtrachia Köhler & Criscione, 2013Aslintesta Solem, 1992Australocosmica Köhler, 2011Austrocamaena J. Stanisic, 2010Austrochloritis Pilsbry, 1891Basedowena Iredale, 1937Baudinella Thiele, 1931Bentosites Iredale, 1933Billordia J. Stanisic, 2010Boriogenia J. Stanisic, 2010Brigaladra L. Stanisic, 2022Calvigenia Iredale, 1938Calycia H. Adams, 1865Canefriula Iredale, 1941Caperantrum Solem, 1997Cardiotrachia Criscione & Köhler, 2014Carinotrachia Solem, 1985Catellotrachia Iredale, 1933Chloritis H. Beck, 1837Chloritisanax Iredale, 1933Chloritobadistes Iredale, 1933Claudettea Iredale, 1941Coliolus Tapparone Canefri, 1886Contramelon Iredale, 1937Cooperconcha Solem, 1992Crikey J. Stanisic, 2009Cristigibba Tapparone Canefri, 1883Cristilabrum Solem, 1981Crookshanksia J. Stanisic, 2010Crystallopsis Ancey, 1887Cupedora Iredale, 1937Damochlora Iredale, 1938Denhamiana J. Stanisic, 2013Dirutrachia Iredale, 1937Discomelon Iredale, 1938Divellomelon Iredale, 1933Euryladra L. Stanisic, 2022Eurytrachia J. Stanisic, 2010Exiligada Iredale, 1939Falspleuroxia Solem, 1997Figuladra Köhler & Bouchet, 2020Forcartia Clench & R. D. Turner, 1963Forrestena J. Stanisic, 2010Galadistes Iredale, 1938Globorhagada Iredale, 1933Gloreugenia Iredale, 1933Glyptorhagada Pilsbry, 1890Gnarosophia Iredale, 1933Granulomelon Iredale, 1933Hadra E. von Martens, 1860Jacksonena Iredale, 1937Jimbouria J. Stanisic, 2010Kandoschloritis Shea & O. L. Griffiths, 2010Kendrickia Solem, 1985Kimberleydiscus Köhler, 2010Kimberleymelon Köhler, 2010Kimberleytrachia Köhler, 2011Kimboraga Iredale, 1939Kymatobaudinia Criscione & Köhler, 2013Lacustrelix Iredale, 1937Lamprellia J. Stanisic, 2010Letitia Iredale, 1941Lorelliana J. Stanisic, 2016Lynfergusonia J. Stanisic, 2010Marilynessa J. Stanisic, 2010Megalacron I. Rensch, 1934Meliobba Iredale, 1940Melostrachia Iredale, 1938Meridistes J. Stanisic, 2010Meridolum Iredale, 1933Mesodontrachia Solem, 1985Micromelon Solem, 1992Molema Köhler, 2011Montanomelon Solem, 1993Monteithosites J. Stanisic, 1996Moretonistes J. Stanisic, 2010Mouldingia Solem, 1984Mussonena Iredale, 1938Nannochlora Criscione & Köhler, 2015Nannochloritis Köhler & Bouchet, 2020Nanotrachia Köhler & Criscione, 2013Neveritis Iredale, 1938Ningbingia Solem, 1981Noctepuna Iredale, 1933Nodulabium Criscione & Köhler, 2013Obsteugenia Iredale, 1933Offachloritis Iredale, 1933Ordtrachia Solem, 1984Ototrachia Criscione & Köhler, 2013Pallidelix Iredale, 1933Papuexul Iredale, 1933Papuina E. von Martens, 1860Papuolus Schileyko, 2003Papustyla Pilsbry, 1893Parachloritis Ehrmann, 1912Parglogenia Iredale, 1938Patrubella Iredale, 1938Perioinsolita J. Stanisic & Potter, 2010Petraconcha S. A. Clark, 2009Planispira H. Beck, 1837Plectorhagada Iredale, 1933Pleuroxia Ancey, 1887Pommerhelix S. A. Clark, 2009Ponderconcha S. A. Clark, 2009Posorites Iredale, 1933Promonturconchum Solem, 1997Protolinitis J. Stanisic, 2010Prototrachia Solem, 1984Prymnbriareus Solem, 1981Pseudcupedora Solem, 1992Pseudodistes J. Stanisic & Potter, 2010Pseudomesodontrachia Criscione & Köhler, 2013Pseudopapuina F. Haas, 1934Quirosena Iredale, 1941Quistrachia Iredale, 1939Rachita Criscione & Köhler, 2014Ramogenia Iredale, 1938Retroterra Solem, 1985Rhagada E. von Martens, 1860Rhynchotrochus Möllendorff, 1895Sauroconcha W.-H. Zhang & Shea, 2008Setobaudinia Iredale, 1933Setocallosa Criscione & Köhler, 2015Sinumelon Iredale, 1930Solmogada Iredale, 1941Sphaerospira Mörch, 1867Spurlingia Iredale, 1933Squamagenia J. Stanisic & Potter, 2010Stanisicia S. A. Clark, 2009Steorra J. Stanisic, 2010Strepsitaurus Solem, 1997Succochlea Criscione & Köhler, 2014Sulcobasis Tapparone Canefri, 1883Tatemelon Solem, 1993Temporena Iredale, 1933Thersites L. Pfeiffer, 1855Tolgachloritis Iredale, 1933Toombatrachia J. Stanisic, 2010Torresitrachia Iredale, 1939Trachiopsis Pilsbry, 1893Trachygenia J. Stanisic, 2010Trozena Iredale, 1938Turgenitubulus Solem, 1981Ventopelita Iredale, 1933Vidumelon Iredale, 1933Vincentrachia Criscione & Köhler, 2013Wahgia Clench & R. D. Turner, 1959Westraltrachia Iredale, 1933Xanthomelon E. von Martens, 1860Xeromelon Criscione & Köhler, 2016Youwanjela Köhler & Shea, 2012Zyghelix Iredale, 1937
Subfamily Helicostylinae:Anixa Pilsbry, 1895Calocochlea W. Hartmann, 1843Chloraea Albers, 1850Chrysallis Albers, 1850Cochlodryas E. von Martens, 1860Cochlostyla A. Férussac, 1821Dolichostyla Pilsbry, 1896Helicobulinus Broderip, 1841Helicostyla A. Férussac, 1821Hypselostyla L. Pfeiffer, 1868Leytia Pilsbry, 1892Mesanella Clench & R. D. Turner, 1952Orthostylus H. Beck, 1837Orustia Mörch, 1852Pachya Albers, 1850Pachysphaera Pilsbry, 1892Pfeifferia J. E. Gray, 1853Phengus Albers, 1850Phoenicobius Mörch, 1852Pyrochilus Pilsbry, 1893Trachystyla Pilsbry, 1892