CHAPPUISIIDAE Chappuis, 1940


CHAPPUISIUS Kiefer, 1938

  • Chappuisius inopinus Kiefer, 1938 [Europe; ground waters]
  • Chappuisius singeri Chappuis, 1939 [Europe: mesopsammon]


    SUPERFAMILY CLETODOIDEA T. Scott, 1905

    CLETODIDAE Scott, 1905 (sensu Por, 1986)

    The family Cletodidae T. Scott, 1905 is a benthic harpacticoid taxon with about 150 species in 38 genera (George 2020, George 2021, Walter and Boxshall 2022), including species of the Ceratonotus-group sensu Conroy-Dalton, 2001 formerly placed in Ancorabolinae Sars, 1909 (George 2020).

    Por (1986) in his review of the family Cletodidae transferred some genera of this family to a new sub-family of Canthocamptidae: Hemimesochrinae, as well as provisionally placed the genus Cletocamptus in the "Canthocamptidae incertae sedis". Later on, Gee (1999), sharing the Por's (1986) opinion, provisionally included the genus Cletocamptus in the family Canthocamptidae incertae sedis, suggestings as well that this genus "must be removed to a new family which will brobably prove to be a sister family to the Cletodidae".

    More recently, Mielke (2000) confirmed that "The situation of the genus Cletocamptus is complex and in some respects inconsistent..... This uncertainty is due to the conjecture that the two families, Canthocamptidae and Cletodidae, appear to be poly- or para-phyletic assemblages".

    The genus Pontocletodes Apostolov, 1980 was not defined by Por (1986) and it is provisionally assigned to Cletodidae. Bodin (1997) reported the same genus as "incertae sedis", adding some new genera to the family Cletodidae.

    "Uncovering the systematics of Copepoda Harpacticoida, the second-most abundant component of the meiobenthos after Nematoda, is of major importance for any further research dedicated especially to ecological and biogeographical approaches. Based on the evolution of the podogennontan first swimming leg, a new phylogenetic concept of the Ancorabolidae Sars and Cletodidae T. Scott sensu Por (Copepoda, Harpacticoida) is presented, using morphological characteristics. It confirms the polyphyletic status of the Ancorabolidae and its subfamily Ancorabolinae Sars and the paraphyletic status of the subfamily Laophontodinae Lang. Moreover, it clarifies the phylogenetic relationships of the so far assigned memb ers of the family.

    An exhaustive phylogenetic analysis was undertaken using 150 morphological characters, resulting in the establishment of a now well-justified monophylum Ancorabolidae. In that context, the Ancorabolus-lineage sensu Conroy-Dalton and Huys is elevated to sub-family rank. Furthermore, the membership of Ancorabolina George in a rearranged monophylum Laophontodinae is confirmed. Conversely, the Ceratonotus-group sensu Conroy-Dalton is transferred from the hitherto Ancorabolinae to the Cletodidae. Within these, the Ceratonotus-group and its hypothesised sister-group Cletodes Brady are combined to form a monophyletic subfamily Cletodinae T. Scott, subfam. nov. Consequently, it was necessary to restructure the Ancorabolidae, Ancorabolinae and Laophontodinae and extend the Cletodidae to include the displacement and exclusion of certain taxa. Moreover, comparison of the Ancorabolidae, Cletodidae, Laophontoidea and other Podogennonta shows that the Ancorabolidae and Cletodidae form sister-groups in a monophylum Cletodoidea Bowman and Abele, which similarly has a sister-group-relationship with the Laophontoidea T. Scott. " (from George, 2020

    "Currently the Ceratonotus group sensu Conroy-Dalton (2001) comprises nine genera: Arthuricornua Conroy-Dalton, 2001, Ceratonotus Sars, 1909, Dendropsyllus Conroy-Dalton, 2003, Dimorphipodia Lee & Huys, 2019, Dorsiceratus Drzycimski, 1967, Echinopsyllus Sars, 1909, Polyascophorus George, 1998, Pseudechinopsyllus George, 2006, and Touphapleura Conroy-Dalton, 2001 (Sars1909; Drzycimski 1967; George 1998, 2006a; Conroy-Dalton 2001; Conroy-Dalton 2003a; Lee & Huys 2019)" (from George, 2021)


    Ceratonotus group sensu Conroy-Dalton (2001)

    POROPSYLLUS George, 2021

  • Poropsyllus menzelae George, 2021

    PARATOUPHAPLEURA George, 2021

  • Paratouphapleura aaroni George, 2021

    PSEUDOPOLIASCOPHORUS George, 2021

  • Pseudopolyascophorus monoceratus (George et al., 2013)

    TAUROCERATUS George, 2021

  • Tauroceratus tauroides (George, 2006)
  • Tauroceratus concavus (Conroy-Dalton, 2003)
  • Tauroceratus steiningeri (George, 2006)
  • Tauroceratus vareschii (George, 2006)


    Pending a definitive review of the family, the genus Cletocamptus is herein provisonally placed in the Cletodidae sensu Scott, 1904; Por, 1986, which presently should include the following other genera: Acrenhydrosoma, Cletodes, Enhydrosoma, Enhydrosomella, Kollerua, Limnocletodes, Stylicletodes [Monocletodes Lang], Barbaracletodes, Australonannopus, Scintis, Strongylacron, Schizacron, Monocletodes, Intercletodes, Dyacrenhydrosoma, Paracrenhydrosoma Triathrix, Sphingothrix, Neoacrenhydrosoma (Por, 1986, Fiers, 1987; Bodin, 1997; Gee, 1998,1999) and Spinapecruris Gee, 2001.


    CLETODES Brady, 1872

    Synonym: Thieliella Schriever, 1984a (part.) (after Schriever, 1986); Liljeborgia Claus, 1866 (synonym)

  • Cletodes abyssicola Scott T. & Scott A., 1901 accepted as Mesocletodes abyssicola (Scott T. & Scott A., 1901)
  • Cletodes armata Scott T., 1903 accepted as Echinocletodes armatus (Scott T., 1903)
  • Cletodes barnishi Wells, 1967
  • Cletodes buchholtzi Boeck, 1873 accepted as Strongylacron buchholtzi (Boeck, 1873)
  • Cletodes brucei Lang, 1948
  • Cletodes bodini (Dinet, 1974) [Europe; marine]
  • Cletodes carthaginiensis Monard, 1935 [north Africa, Tunisia; marine]
  • Cletodes confusum Gomez, 2000 [USA; lagoons]
  • Cletodes curvirostris Scott T., 1894 accepted as Spinapecruris curvirostris (Scott T., 1894)
  • Cletodes dentatus Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Cletodes dissimilis Willey, 1935 [Bermuda; ?]
  • Cletodes dorae Por, 1979 [Red sea]
  • Cletodes endopodita (Schriever, 1984) [north Atlantic ocean]
  • Cletodes hartmannae Lang, 1965 [USA; sea waters]
  • Cletodes hirsutipes Scott T., 1897
  • Cletodes irrasa Scott T. & Scott A., 1894 accepted as Mesocletodes irrasus (Scott T. & Scott A., 1894)
  • Cletodes lata Scott T., 1892 accepted as Eurycletodes (Oligocletodes) latus (Scott T., 1892)
  • Cletodes laticauda Boeck, 1873 accepted as Eurycletodes (Eurycletodes) laticauda (Boeck, 1873)
  • Cletodes latipes Scott A., 1909 accepted as Enhydrosoma latipes (Scott A., 1909)
  • Cletodes latirostris Drzycimiski, 1967 [Norway; sea waters]
  • Cletodes leptostylis Sars G.O., 1920 accepted as Stylicletodes leptostylis (Sars G.O., 1920)
  • Cletodes limicola limicola Brady, 1872 [north Europe, Algerie, Indian ocean; sea waters]
  • Cletodes limicola tunisiensis Monard, 1935 [Tunisia; marine]
  • Cletodes linearis (Claus, 1866)
  • Cletodes longicaudatus (Boeck, 1872) [Palaearctic: sea waters] (=Stylicletodes longicaudatus (Brady & Robertson, 1876)
  • Cletodes longifurca Lang, 1948 [north Europe; sea waters]
  • Cletodes macrura Fiers, 1991 [USA; coastal waters]
  • Cletodes meyerorum George & Muller, 2013 [SE Atlantic]
  • Cletodes millerorum Hamond, 1973 [Australia; sea waters]
  • Cletodes monensis Thompson I.C., 1893 accepted as Mesocletodes monensis (Thompson I.C., 1893)
  • Cletodes neglecta Scott T., 1903
  • Cletodes nordatlantica (Schriever, 1982) accepted as Monocletodes varians (Scott T., 1903) accepted as Metahuntemannia varians (Scott T., 1903)
  • Cletodes numidicus Monard, 1935 accepted as Stylicletodes numidicus (Monard, 1935)
  • Cletodes pectinata Brady & Robertson, 1876
  • Cletodes perplexa Scott T., 1899 accepted as Cletodes perplexus (Scott T., 1899) accepted as Enhydrosoma perplexa (Scott T., 1899) accepted as Acrenhydrosoma perplexa (Scott T., 1899) accepted as Acrenhydrosoma perplexum (Scott T., 1899)
  • Cletodes perplexus (Scott T., 1899) accepted as Enhydrosoma perplexa (Scott T., 1899) accepted as Acrenhydrosoma perplexa (Scott T., 1899) accepted as Acrenhydrosoma perplexum (Scott T., 1899)
  • Cletodes pontica (Krichagin, 1877)
  • Cletodes propinqua Brady & Robertson, 1876 accepted as Enhydrosoma propinquum (Brady, 1880)
  • Cletodes pseudodissimilis Coull, 1971 [USA; meiobenthic]
  • Cletodes pseudodissimilisoris Gomez, 2000 [USA; lagoons]
  • Cletodes pusillus Sars, 1920 [north Europe; sea waters]
  • Cletodes reducta (Schriever, 1982)
  • Cletodes reductus Moore, 1978 [France; sea waters]
  • Cletodes reyssi Soyer, 1964 [France; coastal waters]
  • Cletodes sarsi Scott T., 1905 accepted as Enhydrosoma sarsi (Scott T., 1905)
  • Cletodes setosus Marinov & Apostolov, 1985 [spanish Sahara; coastal waters]
  • Cletodes similis Scott T., 1895 accepted as Eurycletodes (Oligocletodes) similis (Scott T., 1895)
  • Cletodes smirnovi Bodin, 1970 [France; sea intertidal]
  • Cletodes spinulipes Por, 1967 [Elat, Red sea; marine]
  • Cletodes stylicaudatus Willey, 1935 accepted as Stylicletodes stylicaudatus (Willey, 1935)
  • Cletodes subnigra Brady & Robertson, 1876
  • Cletodes tenuipes T. Scott, 1896 [north Europe; sea waters]
  • Cletodes tenuiremis Scott T., 1893 accepted as Itunella tenuiremis (Scott T., 1893)
  • Cletodes tuberculatus Fiers, 1991 [USA; coastal waters]
  • Cletodes varians Scott T., 1903 (syn.Monocletodes varians (Scott T., 1903); Metahuntemannia varians (Scott T., 1903)
  • Cletodes walvisi Dinet, 1974 [Europe; marine]
  • Cletodes yotabis Por, 1967 [Elat, Red sea; marine ]

    KEY TO SPECIES


    ENHYDROSOMA Boeck, 1872 (SYN. Cletodes n. g. Brady, 1872)

    "Enhydrosoma Boeck, 1872 is one of the biggest genera in the family Cletodidae T. Scott, 1905. Gee [1] partially reviewed the taxonomic characters of the genus Enhydrosoma in the Cletodidae. Enhydrosoma species have been moved to other genera or newly erected genera within Cletodidae on several occasions [1–5]. The genus has diverse congeners, more than 53 valid species and several authors suggests a necessity for the revision of the genus since it is polyphyletic in nature" (from: Park et al., 2020)

    The genus Enhydrosoma has been a taxonomic debate since its erection by Boeck (1873). Various attempts (Sars, 1909; Lang, 1936; 1948; 1965; Thistle, 1980; Gee, 1994; Kim et al., 2014; Karanovic et al., 2015) had been taken to construct a monophyletic group but this task still could not be accomplished because of the insufficient descriptions and figures that lack important characters such as the form and setation of the mouthparts, and even periopods on some occasions (Sonmez et al,. 2018)

  • Enhydrosoma apimelon Karanovic et al., 2015
  • Enhydrosoma barnishi (Wells, 1967) (=Schizacron barnishi (Wells, 1967)
  • Enhydrosoma baruchi Coull, 1975 [USA; estuarine waters]
  • Enhydrosoma bifurcarostratum Shen & Tai, 1965 Syn. Schizacron bifurcarostratus (Shen & Tai, 1965)
  • Enhydrosoma birsteini Borutsky, 1971 (= Kollerua birsteini (Borutzky, 1971)
  • Enhydrosoma breviarticulatum Shen & Tai, 1964 (?Kollerua breviarticulatum (Shen & Tai, 1964)
  • Enhydrosoma brevipodum Gomez, 2004 [?: ?] Recently transferred to a new genus Geehyndrosoma Kim & Lee 2014
  • Enhydrosoma buchholtzi (Boeck, 1872) [north Europe, north America; marine] (=Strongylacron buchholtzi (Boeck, 1873)
  • Enhydrosoma caeni Raibaut, 1965 [Thau basin; sea waters]
  • Enhydrosoma cananeiae Jakobi, 1955 [Brazil; brackish costal waters] *
  • Enhydrosoma casoae Gómez, 2003
  • Enhydrosoma coreana Kim et al., 2014 [Korea]
  • Enhydrosoma curticauda Boeck, 1872 [Europe, Italy; sea waters]
  • Enhydrosoma curvatum Brady, 1880 accepted as Rhizothrix curvata Brady, 1880
  • Enhydrosoma curvirostris (T. Scott, 1894) [Europe; psammon] **
  • Enhydrosoma franklini Thistle, 1980 (syn. Enhydrosomella franklini (Thistle, 1980)
  • Enhydrosoma gariene Gurney, 1930 [England; brackish waters]
  • Enhydrosoma gerlachi Jakobi, 1955 [Brazil; brackish costal waters] *
  • Enhydrosoma gracile Scott T., 1903 (=Rhizothrix gracilis (Scott T., 1903)
  • Enhydrosoma guaratubae Jakobi, 1955 [Brazil; brackish costal waters] *
  • Enhydrosoma herrerai Bell & Kern, 1983 [USA; estuarine waters]
  • Enhydrosoma hopkinsi Lang, 1965 [USA; sea waters]
  • Enhydrosoma kosmetron Karanovic et al., 2015
  • Enhydrosoma illgi Chappuis, 1958 (=Orthopsyllus illgi (Chappuis, 1958)
  • Enhydrosoma intermedia Chislenko, 1978 [Japan; sea waters] Recently transferred toa new genus Geehyndrosoma Kim & Lee 2014
  • Enhydrosoma ivetteae Jakobi, 1955 [Brazil; brackish costal waters] *
  • Enhydrosoma kuehnemanni Pallares, 1970 (=Enhydrosomella kuehnemanni Pallares, 1968
  • Enhydrosoma lacunae Jakubisiak, 1933 [Cuba; lagoons]
  • Enhydrosoma latipes (A. Scott, 1909) [Africa; sea waters, plankton]
  • Enhydrosoma latipes (A. Scott, 1909) [Africa; sea waters, plankton]
  • Enhydrosoma littorale Wells, 1967 [Africa, Mozambico; sea waters]
  • Enhydrosoma longicauda Marinov & Apostolov, 1983 [Atlantic ocean]
  • Enhydrosoma longicaudata Boeck, 1872 accepted as Cletodes longicaudatus (Boeck, 1872)
  • Enhydrosoma longifurcatum Sars, 1909 syn. of Enhydrosoma serdarsaki Sonmez et al,. 2018 [Europe, Italy; psammon]
  • Enhydrosoma longum Shen & Tai, 1979 (=Kollerua longum (Shen & Tai, 1979)
  • Enhydrosoma mangroviae Jakobi, 1955 [Brazil; brackish coastal waters] *
  • Enhydrosoma micrurum Monard, 1928 [France; marine]
  • Enhydrosoma migoti Monard, 1926 [France; marine]
  • Enhydrosoma minimum Jakobi, 1955 [Brazil; brackish costal waters] *
  • Enhydrosoma minutum Scott T., 1903 (=Rhizothrix minuta (Scott T., 1903)
  • Enhydrosoma nicobarica Sewell, 1940 [USA?; sea waters] (=Triathrix nicobarica (Sewell, 1940)
  • Enhydrosoma parapropinquum Gómez, 2003
  • Enhydrosoma pectinatum Wells & Rao, 1987 [Andaman, Nicobar islands; coastal waters]
  • Enhydrosoma pericoense Mielke, 1990 [Panama; coastal waters]
  • Enhydrosoma perplexa (Scott T., 1899) (Acrenhydrosoma perplexa (Scott T., 1899) accepted as Acrenhydrosoma perplexum (Scott T., 1899)
  • Enhydrosoma pontica Lang, 1948
  • Enhydrosoma propinquum (Brady, 1880) [Europe, Italy, Algerie; coastal waters] *
  • Enhydrosoma radhakrishnai Reddy, 1979 [South India] (= Kollerua radhakrishnai (Ranga Reddy, 1979)
  • Enhydrosoma robustum Karanovic et al., 2015
  • Enhydrosoma rosae Fiers, 1996 [nom. nov. (Fiers, 1996) for E. propinquum]
  • Enhydrosoma sarsi (T. Scott, 1904) [north Europe; coastal waters]
  • Enhydrosoma sarsi (Scott T., 1904)
  • Enhydrosoma serdarsaki Sonmez et al,. 2018 [Turkey; Black Sea]
  • Enhydrosoma setiensis (Raibaut, 1965) accepted as Enhydrosomella setiensis Raibaut, 1965
  • Enhydrosoma solitarum Gomez, 2003
  • Enhydrosoma sordidum Monard, 1926 [European Atlantic coasts, Black and Mediterranean Seas, Brazil; coastal waters]
  • Enhydrosoma staufferi (Monard, 1935) (=Enhydrosomella staufferi Monard, 1935
  • Enhydrosoma stylicaudatum Willey, 1935 [Bermuda, ?] (=Stylicletodes stylicaudatus (Willey, 1935)
  • Enhydrosoma texana Park et al, 2020 [Northern Gulf of Mexico]
  • Enhydrosoma tunisense Monard, 1935 [Africa, Tunisia; marine] *
  • Enhydrosoma uniarticulatum Borutzky, 1928 [URSS; brackish waters]
  • Enhydrosoma variabile Wells, Hicks & Coull, 1982 [New Zealand; estuarine waters]
  • Enhydrosoma vervoorti Fiers, 1987 (=Schizacron vervoorti (Fiers, 1987)
  • Enhydrosoma vicinum Por, 1967
  • Enhydrosoma wellsi Bodin, 1968 [France; sea waters]
  • Enhydrosoma woodini Thistle, 1980

    * species inquirenda (Lang, 1965; Por et al., 1984)

    ** recently moved to the genus Spinapecruris Gee, 2001 (Gee, 2001)

    In a paper on the genus Enhydrosoma, Jakobi (1955) describcd several new species. However they are so superficially described that in Lang's opinion (1965) they should be simply ignored. Fiers (1996b) considered some of them as junior synonyms of E. lacunae

  • Enhydrosoma ivitteae Jakobi, 1955
  • Enhydrosoma minimum Jakobi, 1955
  • Enhydrosoma guaratubae Jakobi, 1955
  • Enhydrosoma cananeiae Jakobi, 1955
  • Enhydrosoma tunisense Monard, 1935


    KEY TO SPECIES OF ENHYDROSOMA


    GEEHYDROSOMA Kim et al., 2014

  • Geehydrosoma intermedia (Cihislenko, 1978) [Korea]
  • Geehydrosoma brvipodium Kim et al., 2014 [Korea]


    Cletocamptinae Gomes & Rivera, 2022

    CLETOCAMPTUS Schmankewitch, 1875

    Synonymised names Godetella Delachaux, 1918 (synonym) Marshia Herrick, 1895 (synonym) Regis Labbé, 1927 (synonym) Sinocamptus Shen & Sung, 1963 (synonym) Wolterstorffia Schmeil, 1894 (synonym)

    "The genus Cletocamptus is considered cosmopolitan (Suárez-Morales et al, 2013) and its species occur in a wide salinity range, being reported in estuaries, costal and beach lagoons but few species are found in full freshwater" (from Fuentes-Reines et al., 2015).

    The systematic status of the genus is still controversial. Several authors (Por, 1968; Bodin, 1997; Mielke, 2000, 2001; Gomez et al., 2007) include it in Canthocamptidae incertae sedis sensu Por, 1986. Pending a definitive review of the above family, the genus Cletocamptus is herein provisonally placed in the Cletodidae sensu Scott, 1904.

    At present the valid species within Cletocamptus are C. retrogressus Schmankewitsch, 1875, C. confluens (Schmeil, 1894), C. albuquerquensis (Herrick, 1894), C. trichotus Kiefer, 1929, C. feei (Shen, 1956), C. affinis Kiefer, 1957, C. gravihiatus (Shen and Sung, 1963), C. helobius Fleeger, 1980, C. merbokensis Gee, 1999, C. axi Mielke, 2000, C. schmidti Mielke, 2000, C. deborahdexterae Gomez, Fleeger, Rocha-Olivares and Foltz, 2004, C. stimpsoni Gomez, Fleeger, Rocha-Olivares and Foltz, 2004, C. gabrieli Loffler, 1961, C. sinaloensis Gomez, Fleeger, Rocha- Olivares and Foltz, 2004, C. bicolor (Wilson, 1932), C. xenuus Por, 1968 , C. kummieri (Delachaux,1917), C. stimpsoni Gomez, Fleeger, Rocha Oliovarez & Faltz, 2003, C. fourchensis Gomez, Fleeger, Rocha-Olivares and Foltz, 2004, C. cecsurirensis Gomez, Scheihing & La Barca, 2007. C. mongolicus (Sterba, 1967), C. levis Gomez, 2005, C. nudus Gomez, 2005 , C. samariensis Fuentes-Reines et al., 2015.

    Gomez et al. (2004) considered as species inquirendae some species within the genus and they also considered as doubtful some records of C. deitersi. Additional doubtful records of C. deitersi are those of Brehm (1936, 1965), Chappuis (1936), Ringuelet (1958a, 1958b, 1960, 1962), Ringuelet et al. (1965), Oliveira and Miranda (1971), Apostolov (1984), Ruber et al. (1994), and Loftus and Reid (2000).

    Recently moved to Canthocamptidae Brady, 1880, subfamily Canthocamptinae Brady, 1880 (Walter, T.C.; Boxshall, G. (2021). World of Copepods Database. Cletocamptus Schmankevitsch, 1875. Accessed through: World Register of Marine Species at: http://www.marinespecies.org)


  • Cletocamptus affinis Kiefer, 1957 [China; brackish waters]
  • Cletocamptus albuquerquensis (Herrick, 1894) [Americas; fresh, brackish waters] (Cletocamptus chappuisi Gómez, Gerber & Fuentes-Reinés, 2017 (wrongly identified by Chappuis, 1933)
  • Cletocamptus assimilis Gomez & Gee, 2009 [Argentina; inland waters]
  • Cletocamptus axi Mielke, 2000 [Galapagos; lagoons] **
  • Cletocamptus blanchardi (Richard, 1889) accepted as Cletocamptus retrogressus Schmankevitsch, 1875
  • Cletocamptus bicolor (Wilson, 1932) [America; fresh, brackish waters] **
  • Cletocamptus brehmi Kiefer, 1934 accepted as Cletocamptus deitersi (Richard, 1897)
  • Cletocamptus brevicaudatus (Herrick, 1894)
  • Cletocamptus cecsurirensis Gomez, Scheihing & La Barca, 2007 [Chile; fresh waters]
  • Cletocamptus chappuisi Gomez et al., 2012 [Bonaire]
  • Cletocamptus confluens (Schmeil, 1894) [cosmopolitan, except America; fresh, brackish waters, wetlands] *
  • Cletocamtus cubaensis Apostolov, 2023 [Cuba ]
  • Cletocamptus deborahdexterae Gomez, Fleeger, Rocha Oliovarez & Faltz, 2003 [USA; fresh waters] (syn. C. deitersi partim)
  • Cletocamptus deitersi deitersi (Richard, 1897) [Cuba, Yucatan, south America, China, Australia, Israel; Ethiopie; brackish waters, wetlands, cenotes] **
  • Cletocamptus deitersi ecuadorianus Loffler, 1964 [Ecuador, brackish waters] **
  • Cletocamptus dominicanus Kiefer, 1934
  • Cletocamptus feei (Shen, 1956) [China; brackish waters]
  • Cletocamptus fourchensis Gomez, Fleeger, Rocha Oliovarez & Faltz, 2003 [USA; fresh waters]
  • Cletocamptus gabrieli Loffler, 1961 [Iran; brackish waters] **
  • Cletocamptus goenchim Gomez et al., 2013 [India; estuary waters]
  • Cletocamptus gomezi Suarez-Morales et al., 2013 [Mexico; saline lake]
  • Cletocamptus gravihiatus (Shen & Sung, 1963) [China; brackish waters]
  • Cletocamptus helobius Fleeger, 1980
  • Cletocamptus koreanus Chang, 2013 [Korea; estuaries]
  • Cletocamptus kummieri (Delachaux,1917) [south America; marine]
  • Cletocamptus levis Gomez, 2005 [Brazil; marine]
  • Cletocamptus mongolicus (Sterba, 1967) [Mongolia]
  • Cletocamptus nudus Gomez, 2005 [Brazil; marine]
  • Cletocamptus pilosus Gomez & Gee, 2009 [Argentina; inland waters]
  • Cletocamptus retrogressus Schmankevitch, 1875 [Europe, Africa, Asia; fresh, brackish waters]
  • Cletocamptus samariensis Fuentes-Reines et al., 2015
  • Cletocamptus servus (Labbé, 1926) accepted as Cletocamptus confluens confluens (Schmeil, 1894)
  • Cletocamptus schmidti Mielke, 2000 [Galapagos; lagoons]
  • Cletocamptus sinaloensis Gomes et al., 2004 [Mexico, Brazil; fresh waters]
  • Cletocamptus spinulosus Gomez & Gee, 2009 [Argentina; inland waters]
  • Cletocamptus stimpsoni Gomez, Fleeger, Rocha Olivarez & Faltz, 2003 [USA; fresh waters]
  • Cletocamptus tainoi Gomez et al., 2012 [Bonaire]
  • Cletocamptus tertius Gomez & Gee, 2009 [Argentina; inland waters]
  • Cletocamptus thailandensis Boonyanusith & Wongkamhaeng, 2023 [Thailand]
  • Cletocamptus trichotus Kiefer, 1929 [south Africa; fresh, brackish waters]
  • Cletocamptus xenuus Por, 1968 [Dahlak Archipelago, Red Sea; brackish waters] (=Dahlakocamptus xenuus (Por, 1968)


    * Cletocamptus confluens, according to its variability and widespread occurrence, could be considered an "assemblage of barely discernible species" (Mielke, 2000). For the same reason, the subspecies Cletocamptus confluens cyrenaica Brehm, 1908 and Cletocamptus confluens meridionalis Kiefer, 1929 are questionable.

    **taxa inquirenda, according to Wells (2007)




    Map of the distribution of Cletocamptus confluens (after Mielke, 2000 modif.)



    The species Cletocamptus deitersi and Cletocamptus retrogressus are geographically widspread, and morphologically and ecologically variable thus implying the existence of morphologically indistinguishable sibling species (Mielke, 2001).


    | Key to the species of Cletocamptus |


    PONTOCLETODES Apotolov, 1980 (genus incertae sedis)

  • Pontocletodes ponticus Apostolov, 1980 [Black sea; psammon]


    NANNOPODELLA Monard, 1928

  • Nannopodella denisi Monard, 1928


    KOLLERUA Gee, 1994

  • Kollerua birsteini (Borutzky, 1971) [Aral Sea]
  • Kollerua breviarticulatum (Shen & Tai, 1964) [China; fresh waters]
  • Kollerua longum Shen & Tai, 1979 [China; fresh waters]
  • Kollerua radhakrishnai (Ranga Reddy, 1979) [India; lakes]
  • Kollerua uniarticulatum (Borutzky, 1928) [URSS; brackish waters]


    STRONGYLACRON Gee & Huys, 1996

    "The harpacticoid copepod genus Strongylacron Gee & Huys, 1996 belonging to the family Cletodidae T. Scott, 1904 was established by Gee and Huys (1996) as a part of an effort to resolve the relationship among members of Enhydrosoma Boeck, 1873. The latter genus was recognized as a heterogeneous group by Fiers (1987) and Mielke (1990). After the studies of Lang (1936, 1948), this genus was distinguished from Cletodes Brady, 1972 by subtle morphological characteristics such as the form and setation of the distal segment on legs 3 and 4 (Gee 1994, 2001; Gee and Huys 1996). In the revision of Enhydrosoma, Gee (1994) first proposed the buchholtzi-species group representing differences from others in the rostrum, antenna, maxillule and male P3 endopod. In addition, he suggested that this group should be removed from Enhydrosoma. Thereafter, the buchholtzi-species group was revised and the following three genera were established: Schizacron Gee & Huys, 1996, Spinapecruris Gee, 2001, and Strongylacron Gee & Huys, 1996 (Gee 1994, 2001; Gee and Huys 1996). Strongylacron is a monospecific genus containing Strongylacron buchholtzi (Boeck, 1873) (Walter and Boxshall 2016)" (Kim et al., 2016),

  • Strongylacron buchholzi (Boeck, 1872) [north Europe, sea waters]
  • Strongylacron glabrum Kim et al., 2016 [Korea]


    SCHIZACRON Gee & Huys, 1996

  • Schizacron barnishi (Wells, 1967) [Africa, Mozambico; coastal waters]
  • Schizacron bifurcarostratum (Shen & Tai, 1965) [China; rivers]
  • Schizacron intermedius Gee & Huys, 1996 [Thailand; mangrove]
  • Schizacron vervoorti (Fiers, 1987) [India; brackish waters]


    ACRENHYDROSOMA Lang, 1944


    The genera Acrenhydrosoma Lang, 1944, Dyacrenhydrosoma Gee, 1999, Paracrenhydrosoma Gee, 1999 and Neoacrenhydrosoma Gee and Mu, 2000 constitute a well defined lineage (Acrenhydrosoma-lineage) within the Cletodidae. (see Bodin 1997 and Gee. 1999).

    The strongly modified endopodal lobe of the fifth legs in both sexes, the position of the exopods in the fith legs when present, the armature of the female sixth legs, separate these four genera from all other known cletodids (from: Kotwicki & Fiers, 2005).


  • Acrenhydrosoma karlingi Lang, 1965 [USA; sea waters]
  • Acrenhydrosoma maccalli Schizas & Shirley, 1994 [Alaska; sea waters]
  • Acrenhydrosoma perplexa (T. Scott, 1899) [Europe; sand sediments]


    KEY TO SPECIES OF ACRENHYDROSOMA


    DYACRENHYDROSOMA Gee, 1999

  • Dyacrenhydrosoma breviseta Gee, 1999 [S.E.Asia; mangrove forest]


    PARACRENHYDROSOMA Gee, 1999

  • Paracrenhydrosoma karlingi (Lang, 1965)
  • Paracrenhydrosoma kiai Song et al., 2014 [Korea; subtidal muddy bottom]
  • Paracrenhydrosoma maccalli (Schizas & Shirley, 1994) [Alaska; mudflat intertidal]
  • Paracrenhydrosoma oceaniae Kotwicki & Fiers, 2005 [Kongsfjorden (Svalbard)]
  • Paracrenhydrosoma normani Gee, 1999 [Norway; sea waters]

    A key to aid in the identification of the Acrenhydrosoma-complex


    NEOACRENHYDROSOMA Gee & Mu, 2000

  • Neoacrenhydrosoma zhangi Gee & Mu, 2000 [China; muddy sand]


    CHALAROSTHRIX Gomez & Nazari, 2022

  • Chalarosthrix bisetosa Gómez & Nazari, 2022
  • Chalarosthrix mayae (Fiers, 1997)
  • Chalarosthrix nicobarica (Sewell, 1940)


    SPINAPECRURIS Gee, 2001

  • Spinapecruris curvirostris (T. Scott, 1894) [North-West Europe; marine muddy sediment]


    | RELEVANT LITERATURE |



    CRISTACOXIDAE Huys, 1990

    The family Crystacoxidae Huys, 1990 presently includes four genera living in marine psammon (Noodtorthopsyllus Lang, 1965; Cristacoxa, Huys, 1990; Laophontisochra George, 2002) and ground (Cubanocleta Petkovski, 1977) waters.

    NOODTORTHOPSYLLUS Lang, 1965

  • Noodtorthopsyllus psammophilus (Noodt, 1955) [Canarie islands; psammon]
  • Noodtorthopsyllus tageae Huys & Kihara, 2010 [Brazil; interstitial]


    CUBANOCLETA Petkovski, 1977

  • Cubanocleta noodti Petkovski, 1977 [Cuba, cave waters]


    CRISTACOXA Huys, 1990 (syn. of Noodtorthopsyllus according to Huys & Kihara, 2010)

  • Cristacoxa petkovskii Huys, 1990 [West Indies; psammon]


    ACUTICOXA Huys & Kihara, 2010 *

  • Acuticoxa ubatubaensis Huys & Kihara, 2010
  • Acuticoxa biarticulata Huys & Kihara, 2010


    *tentatively assigned to Nannopodidae (ex Huntemanniidae) by Huys & Kihara (2010)


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