 AMEIRIDAE Monard, 1927, emend. Lang, 1936
NITOKRA Boeck, 1865
The genus Nitokra at present includes 81 described species and suspecies, widespread in different fresh, brackish and marine habitats; some species are inhabitants of subterranean waters, four species are known as commensals of marine invertebrates (Karanovic et al. 2015).
Bowman (1988) stressed that the "commonly used spelling Nitocra is replaced by Boeck's correct original spelling Nitokra" notwithstandig that the name Nitocra has been usually used since then. Suarez-Morales et al. (1996) followed this opinion, but Mielke proposed to maintain the spelling Nitocra which is recommended by the International Code of Zoological Nomenclature. In the present list we follow the Bowman's spelling.
Great interspecific variability caused in the past times several synonyms in the genus (Lang, 1948; Por, 1964; Apostolov & Marinov, 1988; Bodin, 1997), and to prove this it would be necessary to redescribe many species and subspecies since their descriptions do not fit within the standard of the modern alpha-taxonomy ( Marcus & Por, 1961; Kunz, 1975; Reid, 1988). Moreover, it will be necessary to rename the numerous homonyms in the genus (Karanovic & Pesce, in press).
In this genus, examples of strong reduction of the armature of the female fifth leg basoendopodite can be observed. Most species possess the basoendopodite of this leg with 5 spines/setae; in other species (N. balli, N. blocki, N. cari, N. laingensis, N. pseudospinipes, N. reunionensis, N. uenoi) the same basoendopodite bears 4 spines/setae; N. phreatica and N. pseudospinipes have 3 spines/setae; N. bisetosa only 2 appendages (Karanovic & Pesce, 2002).
"The taxonomy of the genus, similar to numerous other genera within the family, has posed challenges due to the lack of detailed species descriptions and taxonomic information (Karanovic and Pesce 2002). Initially designated as "Nitokra" by Boeck (1865), the nomenclature was later amended to "Nitocra" (Giesbrecht 1881), a spelling upheld until Bowman (1988) highlighted its erroneous usage. This viewpoint was endorsed by Wells (2007), referencing ICZN (1999: § 33.3.1), which advocates for prevailing name usage, thereby asserting the adoption of "Nitocra" (from: Yiedir & Karaytug, 2024).
Nitokra affinis affinis Gurney, 1927 [Europe, Brazil, Bermuda ?, Black Sea, Egypt; Australia; marine, littoral sand, wetlands ] 
Nitokra affinis colombiensis Fuentes-Reinés & Suárez-Morales, 2014 [Colombia; laguna waters]
Nitokra affinis californica Lang, 1965 [USA; marine, tidal pools]
Nitokra affinis rijekana Petkovski, 1954 [ Yugoslavia ]
Nitokra affinis stygia Por, 1968 [Dahlak Archipelago, Red Sea Israel;
ground waters ]
Nitokra alperi Yiedir & Karaytug, 2024 (Nitocra alperi in the original description)
Nitokra arctolongus Shen & Tai, 1973 [ China ]
Nitokra australis Soyer, 1974 [ Kerguelen ]
Nitokra balli Rouch,1972 [Long-Island, New Guinea] (6)
Nitokra balnearia Por, 1964 [ Israel; mineral spring ]
Nitokra bdellurae (Liddell, 1912) [ Europe, Mexico ]
Nitocrameira bdellurae Liddell, 1912 (synonym)
Nitokra bisetosa Mielke, 1993 [Costa Rica, Galapagos; sandy beaches ]
Nitokra blochi Soyer, 1974 [Kerguelen archipelago, Indian Ocean ]
Nitokra cari Petkovski, 1954 [Adriatic Sea, Yugoslavia ]
Nitokra delaruei Soyer, 1974 [ Kerguelen ]
Nitokra divaricata divaricata Chappuis, 1923 [Europe; commensal, Astacus, Austropotamobius, Potamobius ]
Nitokra divaricata caspica Behning, 1936 [ Caspian Sea ]
Nitokra dubia Sars, 1927 [ Africa, USA, Brazil; coastal lagoons ]
Nitokra elegans (T. Scott, 1904) [ Scotland ] (=Ameira?)
Nitokra elongata Marcus, 1968 [ Romania ]
Nitokra esbe Karanovic et al., 2015 [W- Australia; ground waters]
Nitokra evergladensis Bruno, Reid & Perry, 2002 [USA; ground waters]
Nitokra fallaciosa fallaciosa Klie, 1937 [ Spain, Bulgaria, Iran, north Africa ] (1)
Nitokra fallaciosa baltica Lang, 1965 [Europe, Africa, Asia ] (1)
Nitokra fluviatilis (Galhano,1968) [ Portugal ]
Nitokra fragilis fragilis Sars, 1905 [ New Zealand, India]
Nitokra fragilis paulistana Jakobi, 1956 [ Brazil; coastal lagoons ] (5)
Nitokra phreatica Bozic, 1964 (?)
Nitokra galapagoensis Mielke, 1993 [Galapagos; sandy beaches ]
Nitokra hamata Bodin, 1970 [ France ] ]
Nitokra hibernica hibernica (Brady, 1880) [ Europe, Israel, Asia, URSS, Iran; fresh waters ]
Nitokra hibernica bulgarica Apostolov, 1976 [ Bulgaria ] (?)
Nitokra humphreysi Karanovic & Pesce, 2002 [West-Australia; anchialine waters]
Nitokra husmanni (Kunz, 1976) [ Germany ]
Nitokra hyperidis Jacobi, 1956 [ Brazil; coastal lagoons]
Nitokra incerta (Richard, 1893) [Israel, Turkey, Iran ]
Nitokra intermedia Pesce, 1983 [ Italy ]
Nitokra inuber (Schmankevich, 1875) [ Italy, ? ] (= N. hibernica ?)
Nitokra karanovici Chullaton,2014 [Thailand]
Nitokra kastjanensis Kornev & Chertoprud, 2008
Nitokra koreanus Chang, 2007 [Korea; brackish waters]
Nitokra lacustris lacustris (Schmankevitch, 1875) [cosmopolitan ] (2)
Nitokra lacustris azorica Kunz, 1983 [ Azorres ]
Nitokra lacustris colombianus Reid, 1988 [ Colombia ]
Nitokra lacustis richardi (Chappuis, 1933) [ Curacao ]
Nitokra lacustris pacifica Yeatman, 1983 [ Fiji islands; New Guinea; W-Australia ]
Nitokra lacustris sinoi Marcus & Por, 1961 [ Romania ]
Nitokra lacustris cf. sinoi Marcus & Por, 1961(in: Suarez-Morales et al., 1996) [Yucatan]
Nitokra cf. lacustris (Schmankevitch, 1875) (in: Suarez-Morales et al., 1996) [Yucatan]
Nitokra laingensis Fiers, 1986 [New Guinea, Pacific ]
Nitokra langi (Arlt, 1983) [ Baltic sea ]
[junior syn. of N. fallaciosa baltica Lang, 1965]
Nitokra loweae Özlem Vildiz & Karaytug 2024 (Nitocra loweae in the original description)
Nitokra malaica Kiefer, 1929 [ Indonesie ]
Nitokra marinae Jaschnov, 1935
Nitokra mediterranea mediterranea (Brian, 1928) [ Greece ]
Nitokra mediterranea jakubisiaki (Apostolov, 1980) [ Bulgaria ]
[junior syn. of N. pontica Jakubisiak, 1938]
Nitokra minor minor Willey, 1930 [Bermuda]
Nitokra minor mozambicoe Wells, 1967 [ Mozambique ]
Nitokra phreatica Bozic, 1964 (?)
Nitokra pietschmanni (Chappuis, 1934) [ Hawai, Chine?, Japan ] (3)
Nitokra platypus platypus Daday, 1906 [ Asia ]
Nitokra platypus bakeri Chappuis, 1930 [ Philippines, Japan ]
Nitokra platypus pietschmanni Chappuis, 1934
Nitokra pontica (Jakubisiak, 1938) [ Bulgaria ]
Nitokra pori (Apostolov,1969) [ Bulgaria ]
[junior syn. of N. affinis stygia Por, 1968]
Nitokra psammophila Noodt, 1952 [ Germany ] (1)
Nitokra pseudospinipes Yeatman, 1983 [Tonga, Fiji islands; crab holes ]
Nitokra puebloviejensis Fuentes Reines et al., 2022 [Colombia]
Nitokra pusilla Sars, 1911 [ Europe, Mexico; marine ]
Nitokra quadriseta Wells & Rao, 1987 [ Andaman and Nicobar islands ]
Nitokra reducta reducta (Schafer, 1936) [ Europe, Near East ] (7)
Nitokra reunionensis Bozic, 1969 [Reunion Island ]
Nitokra serdarsaki Özlem Vildiz & Karaytug 2024 (Nitocra loweae in the original description)
Nitokra sewelli sewelli Gurney, 1927 [ India ]
Nitokra sonmezi Özlem Vildiz & Karaytug 2024 (Nitocra sonmezi in the original description)
Nitokra sphaeromata Bowman, 1988 [ Costa Rica; associated with wood-boring isopods ]
Nitokra spinipes spinipes Boeck, 1865 [ cosmopolitan; brackish waters ] (4)
Nitokra spinipes armata Lang, 1965 [ Usa, marine sand]
Nitokra spinipes orientalis Sewell, 1924 [Red Sea; lakes]
Nitokra cf. spinipes Boeck, 1865 (in: Suarez-Morales et al., 1996) [Yucatan]
Nitocra sewelli husmanni Kunz 1976
Nitokra taylori Gomez et al., 2010 (described as Nitocra taylori) [South Africa; aetuarine lake]
Nitokra typica typica Boeck, 1865 [ Europe, Baltic sea, Africa, America; marine sand, brackish waters ]
Nitokra cf. typica Boeck, 1865 (in: Suarez-Morales et al., 1996) [Yucatan]
Nitokra typica adriatica Petkovski, 1954 [ Yugoslavia ]
Nitokra uenoi Miura,1962 [ Ryukyu Islands, Pacific]
Nitokra vietnamensis Tran & Chang, 2011 [Vietnam] (described as Nitocra vietnamensis)
Nitokra yeelirrie Karanovic et al., 2015 [W- Australia; ground waters]
INCERTAE SEDIS
Nitokra chelifer Wilson, 1932 [ USA ]
Nitokra gracilimana Giesbrecht, 1902 [ Antarctic ]
Nitokra medusaea Humes (?)
Nitokra phlegraea Brehm, 1909 [ Italy ]
Nitokra wolterecki Brehm, 1909 [ Italy ]
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(1) N. fallaciosa reported by Petkovski (1964) and Noodt, (1953) and N. fallaciosa baltica described by Lang (1965) are probably the same taxon; N. fallaciosa presented by Dussart (1981) most probably do not belong to any of the above mentioned taxa. Notwithstanding the scepticism of Noodt (1953) and Lang (1965), and the interpretation of Dussart & Defaye (1990), N. psammophila described by Nood (1952) could be a valid species since several characters, such as the shape of furcal ram, width and armature of the anal operculum, female fifth leg, quite separate this species from N. fallaciosa (Klie, 1937; Noodt, 1953). (2) Synonyms: N. muelleri van Douwe, 1904; N. simplex Schmeil, 1894; N. treforti Chappuis, 1929; Canthocamptus yahiai Blanchard, 1891 (3)
Synonym of N. platypus (?); Ishida & Kikuchi (2000) report this species from Japan as valid taxon. (4)
For this variable species, Well & Rao (1987) rejected all the described subspecies. Synonyms: N. medusae Humes, 1953; N. palustris Breemen, 1907; N. parafragilis Roe, 1959 (5) species inquirenda, probably a subspecies of Nitokra spinipes (Lang, 1965).
(6) could be synonym of Nitokra uenoi Miura,1962 (Karanovic, pers. com.) (7) According to Dussart (1990), intermediate taxon between Nitokra and Nitocrella |
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RELEVANT REFERENCES
Bodin, P. 1997. Catalogue of the new marine Harpacticoid Copepods. Studiedocumenten van Het K.B.I.N., 89, pp.1-304.
Bowman, T.E. 1988. Nitokra sphaeromata, a new harpacticoid copepod crustacean associated with the wood-boring isopod, Sphaeroma peruvianum, in Costa Rica. Proceedings of the Biological Society of Washington, 101 (1):171-175.
Defaye, D. 1996. Redescription of Nitokra divaricata Chappuis, 1923 (Copepoda, Harpacticoida) with first records from Austropotamobius torrentium Schrank. Acta Zoologica Academiae Scientiarum Hungaricae, Prague, 42 (2):145-155.
Dussart, B. (H.) & D. Defaye. 1990. Répertoire mondial des crustacés copépodes des eaux intérieures. III. Harpacticoďdes. Crustaceana, suppl. 16:1-364.
Karanovic T. & G.L. Pesce. 2002. Copepods from ground waters of Western Australia, VII. Nitokra humphreysi sp. nov. (Crustacea: Copepoda: Harpacticoida). Hydrobiologia, 470: 5-12
Karanovic T., Eberhard T.,
Cooper S.J.B., Guzik M.T. 2015. Morphological and molecular study of the genus Nitokra
(Crustacea, Copepoda, Harpacticoida) in a small palaeochannel
in Western Australia. Org Divers Evol (2015) 15:65–99
Mielke, W. 1993. Species of the taxa Orthopsyllus and Nitocra (Copepoda) from Costa Rica. J. Microfauna Marina, 8:247-266.
Mielke, W. 1997. Interstitial Fauna of Galapagos. XXXIX. Copepoda, part 7. Microfauna marina, 11: 115-152.
Por, F.D. 1964. The genus Nitocra Boeck (Copepoda, Harpacticoida) in the Jordan Rift Valley. Israel J. Zool. ,13: 78-88.
Rouch, R. 1984. Copepoda: les harpacticoides souterraines des eaux douces continentales. Stygofauna Mundi: 321-355.
Vervoort W. 1986. Bibliography of copepoda, up to and including 1980., 1-3. Crustaceana: 1-1316.
Walter, T.C.; Boxshall, G. (2021). World of Copepods Database. Nitokra Boeck, 1865. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=115198 on 2021-06-07
Wells, J.B.J. 1976. Keys to aid in the identification of marine harpacticoid copepods. Aberdeen Univ. press, pp.1-215.
Wells J.B. & G.C. Rao. 1987. Littoral Harpacticoida (Crustacea: Copepoda) from Andaman and Nicobar Islands. Mem. Zool. Survey India 16: 1-385
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