Camaenidae

Family of gastropods

Camaenidae

A live individual of Meridolum gulosum
Scientific classification
Domain:  | Eukaryota
Kingdom:  | Animalia
Phylum:  | Mollusca
Class:  | Gastropoda
Subclass:  | Heterobranchia
Order:  | Stylommatophora
Superfamily:  | Helicoidea
Family:  | Camaenidae
Pilsbry, 1895[1]
Type genus
Camaena

Albers, 1850

Genera

See text

Synonyms

* Bradybaenidae Pilsbry, 1934 (1898)
* Crassispirinae McLean, 1971
* Zonulispirinae McLean, 1971

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[edit]

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.[5]

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).[6]

## Subfamilies[edit]

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]

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

## Genera[edit]

Currently, the following genera are accepted within the family Camaenidae:[13][14]

Taxa with main occurrence in South-East Asia

* Camaena Albers, 1850 – type genus of the family Camaenidae
* Amphicoelina Haas, 1933[15][16]
* Amphidromus Albers, 1850
* Bouchetcamaena Thach, 2018 (debated synonym of Trichochloritis Pilsbry, 1891)
* Chloritis Beck, 1837[17]
* Entadella Páll-Gergely & Hunyadi, 2016
* Ganesella W. T. Blanford, 1863
* Globotrochus F. Haas, 1935
* Jiaoliaous Zhang, 2024 [18]
* Karaftohelix Pilsbry, 1927
* Moellendorffia
* Neocepolis Pilsbry, 1891
* Obba
* Parachloritis Ehrmann, 1912
* Pfeifferia Gray, 1853
* 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[19]
* Aetholitis Stanisic, 2010[20]
* Amplirhagada Iredale, 1933[21]
* Arnemelassa Iredale, 1938[22]
* Aslintesta Solem, 1992[23]
* Australocosmica Köhler, 2011[24]
* Austrocomaena Stanisic, 2010[20]
* Austrochloritis Pilsbry, 1890[25]
* Basedowena Iredale, 1937[26]
* Baudinella Thiele, 1931[27]
* Bentosites Iredale, 1933[21]
* Billordia Stanisic, 2010[20]
* Boriogenia Stanisic, 2010[20]
* 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[28]
* 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[20]
* 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[20]
* Kandochloritis Shea & Griffiths, 2010[20]
* Kendrickia Solem, 1985
* Kimberleydiscus Köhler, 2010[13] – with one species Kimberleydiscus fasciatus Köhler, 2010[13]
* Kimberleymelon Köhler, 2010[13] – with one species Kimberleymelon tealei Köhler, 2010[13]
* Kimberleytrachia Köhler, 2011[29]
* Kimboraga Iredale, 1939
* Lacustrelix Iredale, 1933
* Lamprellia Stanisic, 2010[20]
* Lynfergusonia Stanisic, 2010[20]
* Marylnessa Stanisic, 2010[20]
* Meliobba Iredale, 1940
* Melostrachia Solem, 1979
* Meridistis Stanisic, 2010[20]
* Meridolum Iredale, 1942
* Mesodontrachia Solem, 1985
* Micromelon Solem, 1992
* Minimelon Solem, 1993
* Molema Köhler, 2011[29]
* Montanomelon Solem, 1993
* Monteithosites Stanisic, 1996
* Moretonistes Stanisic, 2010[20]
* 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[20]
* Petraconcha Clark, 2009[30]
* Plectorhagada Iredale, 1933
* Pleuroxia Ancey, 1887
* Pommerhelix Clark, 2009[30]
* Ponderconcha Clark, 2009[30]
* 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[31]
* 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[20]
* Stanisicia Clark, 2009[30]
* Strepsitaurus Solem, 1997
* Tatemelon Solem, 1993
* Temporena Iredale, 1933
* Thersites Pfeiffer, 1855
* Tolgachloritis Iredale, 1933
* Toombatrachia Stanisic, 2010[20]
* 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[32]

## References[edit]

1. ^ Pilsbry H. A. (1895). Manual of Conchology, structural and systematic, with illustrations of the species. Second series: Pulmonata. Helicidae – Volume VII. 9(33a): xxxii.
2. ^ 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.
3. ^ 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.
4. ^ Bouchet, P., Rocroi, J.P. Classification and Nomenclator of gastropod families. Malacologia 47: 1-397.
5. ^ Scott B. 1996. Phylogenetic relationships of the Camaenidae (Pulmonata: Stylommatophora: Helicoidea). Journal of Molluscan Studies, 62: 65-73. Abstract
6. ^ 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.
7. ^ Kobelt W. (1902). Systematisches Conchilien-Cabinet, ed. 2, Bd. 1, Abt 13, Theil 2: 1033.
8. ^ Iredale T. (1938). The Australian Zoologist 9(2): 112.
9. ^ Solem A. (1992). Records of the South Australian Museum, Monograph series 2: 161.
10. ^ Schileyko A. A. (1998-2003). Treatise on Recent terrestrial pulmonate molluscs. Ruthenica supplement 2.
11. ^ Revised Classification, Nomenclator and Typification of Gastropod and Monoplacophoran Families; Malacologia 61(1-2):1-526. 2017
12. ^ "WoRMS - World Register of Marine Species - Camaenidae Pilsbry, 1895". www.marinespecies.org. Retrieved 15 March 2023.
13. ^ a b c d e Köhler F. (2010). "Three new species and two new genera of land snails from the Bonaparte Archipelago in the Kimberley, Western Australia (Pulmonata, Camaenidae)". Molluscan Research 30(1): 1-16.
14. ^ ITIS
15. ^ Haas, F. (1933). "Zur Systematik der chinesischen "Helicodonten"". Archiv für Molluskenkunde. 65 (4/5): 230–231.
16. ^ Páll-Gergely B. & Asami T. (2014). "Additional information on the distribution, anatomy and systematics of living and fossil Chinese Plectopylidae (Gastropoda: Pulmonata)". Genus 25(3): 527-564. PDF.
17. ^ Maassen W. J. M. (2009). "Remarks on the genus Chloritis in Sulawesi, Indonesia, with the descriptions of two new species (Gastropoda: Pulmonata: Camaenidae)". Zoologische Mededelingen 83 HTM Archived 16 January 2014 at the Wayback Machine.
18. ^ Zhang, Guoyi; Naggs, Fred; Peter, Andrus; Christopher M, Wade (2024). "Phylogenetic insights into the terrestrial snails Helicoidei (Gastropoda: Stylommatophora) with special emphasis on the Camaenidae". Zoological Journal of the Linnean Society. doi:10.1093/zoolinnean/zlae027.
19. ^ Stanisic, J. 1996. New land snails from boggomoss environments in the Dawson Valley, southeastern Queensland (Eupulmonata: Charopidae and Camaenidae). Memoirs of the Queensland Museum 39: 343-354
20. ^ a b c d e f g h i j k l m n o Stanisic, J., Shea, M., Potter, D. and Griffiths, O. 2010. Australian Land Snails – Volume 1: A field guide to eastern Australian species. Bioculture Press, Mauritius, 591 pp.
21. ^ a b Iredale, T. 1933. Systematic notes on Australian land shells. Records of the Australian Museum 19: 37-59
22. ^ Iredale, T. 1938. A basic list of the land Mollusca of Australia. Pt III. Australian Zoologist 9: 83-124
23. ^ Solem, A. 1992. Camaenid land snails from southern and eastern South Australia, excluding Kangaroo Island. Pt 1. Systematics, distribution and variation. Records of the South Australian Museum Monograph Series 2: 1-338
24. ^ Köhler, F. 2011. Australocosmica, a new genus of land snails from the Kimberley, Western Australia (Eupulmonata, Camaenidae). Malacologia, 53(2): 199−216
25. ^ Pilsbry, H.A. 1890. in Tryon, G.W. & Pilsbry, H.A. Manual of Conchology. Philadelphia : Conchological Section, Academy of Natural Sciences Ser. 2 Vol. 6 324 pp.
26. ^ Iredale, T. 1937. An annotated check list of the land shells of South and Central Australia. South Australian Naturalist 18: 6-59
27. ^ Thiele, J. 1931. Handbuch der Systematischen Weichtierkunde. Jena : Gustav Fischer pp. 377-778
28. ^ Stanisic J. (24 August) 2009. Crikey steveirwini gen. et sp. nov. from montane habitats in the Wet Tropics of northeastern Queensland, Australia (Gastropoda: Eupulmonata: Camaenidae). Zootaxa 2206: 62–68. abstract.
29. ^ a b Köhler, F. 2011. The camaenid species of the Kimberley Islands, Western Australia (Stylommatophora: Helicoidea). Malacologia, 54(1-2): 203–406
30. ^ a b c d Clark, S. 2009. A review of the land snail genus Meridolum (Gastropoda: Camaenidae) from central New South Wales, Australia. Molluscan Research, 29:61-120.
31. ^ Zhang, W.-H. & Shea, M. (2008). A new genus and species of land snail of the family Camaenidae from New South Wales. Molluscan Research, 28: 123-132.
32. ^ Köhler, F. & Shea, M. (2012). Youwanjela, a new genus of land snail from the Kimberley, Western Australia (Eupulmonata, Camaenidae). Zoosystematics and Evolution, 88: 25-31.

## Further reading[edit]

* 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.

## External links[edit]

* Media related to Camaenidae at Wikimedia Commons
* "Family Camaenidae". Australian government. Archived from the original on 13 October 2012.
* "Camaenid Land Snails from Western and Central Australia (Mollusca: Pulmonata: Camaenidae) | Western Australian Museum". Western Australian Museum. Retrieved 13 May 2023.

Taxon identifiers

Camaenidae|

* Wikidata: Q927331
* Wikispecies: Camaenidae
* AFD: Camaenidae
* BOLD: 1167
* CoL: 7NG9L
* EoL: 2631
* GBIF: 5495
* iNaturalist: 85654
* IRMNG: 114509
* ITIS: 77709
* NCBI: 83226
* Open Tree of Life: 725733
* Paleobiology Database: 333755
* WoRMS: 762708

Authority control databases: National |

* Israel

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