CYCLOPINAE Kiefer, 1927


MESOCYCLOPS G.O. Sars, 1914


The genus Mesocyclops G.O. Sars, 1914 today includes more than 73 valid, nominal species/subspecies, distributed worldwide in continental and coastal waters. The genus has diversified to a great extent in the Palaeotropics, appearing to be rare or absent from the Arctic, Antarctica and temperate regions (Dussart & Fernando, 1984; Samraoui et al., 1998; Holynska, 1997, 2000) (left: M. chaci).

From an ecological point of view, the genus includes species which live in large lakes, caves, and large groundwater bodies, cenotes, hypogean karstic crevices and wells; some species are known as predators on smaller freshwater animals, such as mosquito larvae, cladocerans, rotifers and other copepods.

Until the 1980's the taxonomy of the genus remained poorly known; later on, Kiefer (1981) reviewed the Old World species of Mesocyclops, pointing out that the type-species of the genus (Mesocyclops leuckarti), previously considered as cosmopolitan, is confined to the Palaearctic area. More recently Van de Velde (1984) provided a complete and updated review of the genus in the African continent; Mirabdullayev (1996) and Holynska (2000) reviewed the genus in Uzbekistan and the Australasian region, respectively.



Map with number of species/subspecies of Mesocyclops
from different geographical regions (after Holynska, 2000)


Within the genus Mesocyclops a "thermocyclopoides species-group" has been defined by Holynska & Fiers (1994, 1997) on the basis of the distinct spinule pattern on the frontal surface of the maxillary coxa, and other characters as well: most members of the this group show a common biogeographical feature in the easy dispersal and/or early colonization (Holynska & Fiers, 1994; Holynska, 1997; Fiers et al., 2000).

Recently Alekseev (2021) divided "the genus into four subgenera based on the morphological and habitat characteristics. The subgenus Neomesocyclops subgen. n. comprises species almost exclusively living in the New World. The single exception is M. (N.) tenuisaccus (Sars 1927), described from South Africa, which may prove to be a junior synonym of M. (N.) annulatus (Wierzejski 1892), from South America, and could have appeared in Africa as a result of anthropogenic invasion from the New World.

The subgenus Tethymesocyclops subgen. n. includes species the distributions of which coincide with the borders of the former subcontinent Gondwana, or rather, the coast of the Tethys Sea, during the separation of Gondwana from Laurasia. These two subgenera contain species with a seta on the inner outgrowth of the basal segment of the first swimming leg pair. This seta is absent in all representatives of the nominative subgenus Mesocyclops s. str. with bare caudal rami and last thoracic somite, as well as in the new subgenus Pilosomesocyclops subgen. n., which has hairs on these structures. Most species of the latter two subgenera live in water bodies of the Old World.


MESOCYCLOPS s. latu

  • Mesocyclops aequatorialis aequatorialis Kiefer, 1929 [Africa, Madagascar; fresh waters]
  • Mesocyclops aequatorialis pescei Petkovski, 1986 (=Mesocyclops pescei)
  • Mesocyclops aequatorialis similis Van de Velde, 1984 [Africa, Uzbekistan; fresh waters]
  • Mesocyclops aequatorialis sokotoensis Jeje & Fernando, 1992
  • Mesocyclops affinis Van de Velde, 1987 [from New Guinea to Indochina; holes with green algae]
  • Mesocyclops arcanus Defaye, 1995 [Israel; inland waters]
  • Mesocyclops aspericornis (Daday, 1906) [Asia, Africa, central and south America, Philippines, Malaysia, Oceania, Australia; fresh waters, marshes, karst holes, wells]
  • Mesocyclops australiensis (G.O. Sars, 1908) [Australia, Tasmania; fresh waters]
  • Mesocyclops brooksi Pesce et al., 1996 [Australia; hyporheic]
  • Mesocyclops dadayi Holynska, 1997 [India; fresh waters]
  • Mesocyclops dybowskii (Landé, 1890)
  • Mesocyclops dybowskii fortii Brehm, 1911
  • Mesopsyllus dimorphus Mu & Huys, 2017 [China]
  • Mesocyclops dissimilis Defaye & Kawabata, 1993 [Japan, China, north Vietnam; lakes]
  • Mesocyclops dussarti Van de Velde, 1984 [West-Africa; fresh waters]
  • Mesocyclops ferjemurami Holynska & Vu, 2000
  • Mesocyclops friendorum Holynska, 2000 [Indonesia; ricefields]
  • Mesocyclops geminus Holynska, 2000 [Borneo; stream pools]
  • Mesocyclops granulatus Dussart & Fernando, 1988 [India, Pakistan; lakes, pond]
  • Mesocyclops guangxiensis Reid & Kay, 1992 [China; muddy ponds] (8)
  • Mesocyclops jakartensis Alekseev, 2013 [Java; city ponds]
  • Mesocyclops kieferi Van de Velde, 1984 [Yemen, Africa, Brazil, Sri Lanka; fresh waters, wells]
  • Mesocyclops isabellae isabellae Dussart & Fernando, 1988 [India, Pakistan, Sri Lanka; water reservoirs]
  • Mesocyclops isabellae var. nepalensis Dussart & Fernando, 1988 [Nepal; fresh waters]
  • Mesocyclops medialis Defaye, 2001 [New Caledonia;fresh waters]
  • Mesocyclops microlasius Kiefer, 1981 [Philippines; fresh waters]
  • Mesocyclops notius Kiefer, 1981 [Australia; fresh waters]
  • Mesocyclops ogunnus Onabamiro, 1957 [Africa, Near East, south and southeast Asia, Malaysia, Brazil, Cayman Islands; fresh waters]
  • Mesocyclops parentium Holynska, 1997 [Sri Lanka, south India; fresh waters]
  • Mesocyclops roberti Holinska e Stoch, 2011[South Pacific islands]
  • Mesocyclops spiniferus Mu & Huys, 2017 [China]
  • Mesocyclops thermocyclopoides Harada, 1931 [from Flores through Burma, Malaysia, Java, Indochina, Taiwan to the Ryukyus; fresh waters]
  • Mesocyclops thermocyclopoides acutus Dussart & Fernando, 1988 [Laos; lakes, large ponds] (1)
  • Mesocyclops thermocyclopoides australiensis (Sars G.O., 1908)
  • Mesocyclops tobae Kiefer, 1933 [Indonesia, Sumatra; fresh waters]
  • Mesocyclops yesoensis Ishida, 1999 [Japan]
  • Mesocyclops woutersi Van de Velde, 1987 [Australia, Papua, Indochina, south China, Ryukyus, Honshu; coral gravel pit]
  • Mesocyclops wraniki Baribwegure & Dumont, 2000 [Island of Soqotra, Indian Ocean; interstitial]


    Within the above group, a "woutersi-superspecies group" has been recently defined (Holinska, 2000), including the following taxa: M. woutersi; M. parentium; M. friendorum; M. dissimilis [distinguishing characters of the species in : Holynska (2000)].

    Gutierrez-Aguirre & Suarez-Morales (2001) reviewed the distributional and taxonomic status of the genus, pointing out that "the main characters to recognize species (i.e. presence/absence of a spine on the basipodite ofthe first trunk limb, the presence/absence of teethlike projections on the intercoxal sclerite of the fourth trunk limb, and the shape of the seminal receptacle) have not been strong enough to make a clear, confident taxonomic border between morphologically similar species", and this is most evident in species of the following groups: "thermocyclopoides", "meridianus-brasilianus" and "reidae-ellipticus".

    Mirabdullayev (2007) synonymized Mesocyclops aequatorialism similis with Mesocyclops thermocyclopoides.



    OTHER SPECIES AND SUBSPECIES IN THE GENUS MESOCYCLOPS


  • Mesocyclops acanthoramus Holinska & Brown, 2003 [Australia, Queensland; service manhole]
  • Mesocyclops americanus Dussart, 1985 [north America; fresh waters]
  • Mesocyclops annae Kiefer, 1930 [Madagascar, India, Sri Lanka; fresh waters]
  • Mesocyclops annulatus annulatus (Wierzejski, 1892) [south America; lakes]
  • Mesocyclops annulatus diversus Herbst, 1962 [Brazil; fresh waters, rivers]
  • Mesocyclops arakhlensis Alekseev, 1993 [Siberia; lakes, plancton] (2)
  • Mesocyclops augusti Papa & Holinska, 2013 [Philippines; fresh waters]
  • Mesocyclops bodanicola Kiefer, 1955 represented as Mesocyclops leuckarti bodanicola Kiefer, 1929
  • Mesocyclops borneoensis Dussart & Fernando, 1988 [Indonesie; water reservoirs] (4)
  • Mesocyclops bosumtwii Mirabdullayev et al., 2007 [Africa, Ghana; lakes]
  • Mesocyclops brasilianus Kiefer, 1933 [central and south America; fresh and brackish waters]
  • Mesocyclops brevisetosus Dussart & Sarnita, 1986 [Borneo; fresh waters]
  • Mesocyclops chaci Fiers, 1996 [Mexico, Yucatan; wells]
  • Mesocyclops cokeri Najam-un-Nisa, Mahoon & Irfan Khan, 1987 (nomen dubium)
  • Mesocyclops curvatus Kiefer, 1981 accepted as Mesocyclops salinus Onabamiro, 1957
  • Mesocyclops cuttacuttae Dumont & Maas 1985 [north Australia; caves]
  • Mesocyclops darwini Dussart & Fernando, 1988 [North-Australia, Indonesia, New Guinea; lagoons, ponds, ephemereal waterbodies]
  • Mesocyclops dayakorum Holynska, 2000 [Indonesia; swamps]
  • Mesocyclops edax (S.A. Forbes, 1891) [north and central America, Mexico, Cuba; fresh waters]
  • Mesocyclops evadomingoi Gutierrez-Aguirae & Suarez-Morales, 2001 [Mexico;ponds]
  • Mesocyclops ellipticus Kiefer 1936 [central and south America, Cuba; surface and ground waters, temporary water bodies]
  • Mesocyclops ferjemurami Holinska & Sinh Nam, 2000 [Vietnam, north India, Sri Lanka; wells]
  • Mesocyclops forbesi Najam-un-Nisa, Mahoon & Irfan Khan, 1987 (nomen dubium)
  • Mesocyclops francisci Hoynska, 2000 [Malaysia; lakes]
  • Mesocyclops frankfiersi Alekseev, 2021
  • Mesocyclops holynskae Karanovic, 2006 [Australia; fresh waters,springs]
  • Mesocyclops insulensis Dussart, 1982 [Madagascar; fresh waters]
  • Mesocyclops intermedius Pesce 1985 [West Indies; wells]
  • Mesocyclops iranicus Lindberg, 1936 [Iran; fresh waters] (3)
  • Mesocyclops kawamurai Kikuchi K., 1940
  • Mesocyclops kayi Holinska & Brown, 2004 [Northern Burma; fish pond]
  • Mesocyclops leuckarti leuckarti (Claus, 1857) [Palaearctic region; fresh waters] (9)
  • Mesocyclops leuckarti araklensis Alekseev, 1993 [Siberia; fresh waters]
  • Mesocyclops leuckarti f. bodanicola Kiefer, 1929 [Europe; fresh waters]
  • Mesocyclops leuckarti deccanensis Lindberg, 1935
  • Mesocyclops leuckarti f. fortii Mann, 1940 [Turkey; fresh waters]
  • Mesocyclops leuckarti vincentiana (Brian, 1925) accepted as Mesocyclops annulatus (Wierzejski, 1892)
  • Mesocyclops longisetus longisetus (Thiébaud, 1914) [north, central and south America; fresh and brackish waters, marshes, ground waters]
  • Mesocyclops longisetus araucanus Loffler, 1961 [Chile; fresh waters]
  • Mesocyclops longisetus curvatus Dussart, 1987 (Canada, Yucatan, Cuba, south America; fresh waters, ponds, reservoirs]
  • Mesocyclops major G.O. Sars, 1927 [Africa; fresh waters]
  • Mesocyclops mariae Guo, 2000 [China; fresh waters]
  • Mesocyclops meridianus (Kiefer, 1926) [central and south America; fresh and brackish waters]
  • Mesocyclops meridionalis Dussart & Frutos, 1985 [Brazil, Argentina; ricefields, fresh water ponds]
  • Mesocyclops microspinulosus Lindberg, 1942 represented as Thermocyclops microspinulosus Lindberg, 1942
  • Mesocyclops monardi (Perret, 1925)
  • Mesocyclops mongoliensis Kiefer, 1981 [Mongolia; fresh waters]
  • Mesocyclops neglectus G.O. Sars (?)
  • Mesocyclops nicaraguensis Herbst, 1960 [Nicaragua; fresh waters] (5)
  • Mesocyclops oblongatus G. O, Sars (?)
  • Mesocyclops ogunnus Onabamiro, 1957
  • Mesocyclops obsoletus (Koch, 1838)
  • Mesocyclops paludosus Lindberg, 1956 [Africa; swamps]
  • Mesocyclops papuensis Van de Velde, 1987 [Australia, New Guinea, Indonesia; swamps]
  • Mesocyclops paranensis Dussart & Frutos, 1985 [south America; water reservoirs]
  • Mesocyclops pehpeiensis Hu, 1943 [China, Birmania, Korea, Vietnam, India?, Japan, Sri Lanka, Mexico, Cuba; introduced in the USA; fresh waters]
  • Mesocyclops pescei (Petkovski 1986) [Crooked Island, Bahamas,Yucatan; ground waters]
  • Mesocyclops pilosus Kiefer, 1930 [Madagascar; fresh waters]
  • Mesocyclops pseudoannae Van de Velde,1987 [New Guinea, Australia?; marsh]
  • Mesocyclops pseudomeridianus Defaye & Dussart, 1988 [French Guyana; rivers]
  • Mesocyclops pseudospinosus Dussart & Fernando, 1988 [Indonesie; water reservoirs]
  • Mesocyclops pubiventris Holinska & Brown, 2003 [Australia, Queensland; service manhole]
  • Mesocyclops rarus Kiefer, 1981 [Africa; lakes]
  • Mesocyclops rectus Lindberg, 1937 [India; fresh waters] (6)
  • Mesocyclops reidae Petkovski 1986 [Colombia, Honduras, Yucatan; ricefields, ground waters, cenotes]
  • Mesocyclops restrictus Dussart & Fernando, 1985 [Burma; fresh waters]
  • Mesocyclops roberti Holynska & Stoch, 2011 [South Pacific; volcaniv lake]
  • Mesocyclops ruttneri Kiefer, 1981 [India, Sri Lanka, Uzbekistan,China, Japan, Thailand, Vietnam, USA; fresh waters] (7)
  • Mesocyclops salinus Onabamiro, 1957 [Africa, Madagascar?; brackish waters]
  • Mesocyclops salinus ssp. Samraoui et al., 1998 [Algerie; coastal wetlands]
  • Mesocyclops shenzhenensis Guo, 2000 [China; fresh waters]
  • Mesocyclops simillimus Brady, 1907
  • Mesocyclops sondoongensis Tran & Holynska, 2015
  • Mesocyclops spinosus Van de Velde, 1984 [Africa; fresh waters]
  • Mesocyclops splendidus Lindberg, 1943 [India, Sri Lanka, Bangladesh; fresh waters]
  • Mesocyclops tenuisaccus (G.O. Sars, 1927) [south Africa; lakes]
  • Mesocyclos tenuis Marsch, 1966 (?)
  • Mesocyclops varius Dussart, 1987 [south America; ponds]
  • Mesocyclops venezolanus Dussart, 1987 [south America; ponds]
  • Mesocyclops yenae Holinska, 1998 [Vietnam; fresh waters]
  • Mesocyclops yutsil Reid 1996 [Mexico, Yucatan; cenotes]


    (1) synonym of M. affinis Van de Velde, 1987 (Holynska, in litt.)

    (2) Holinska (in litt.) suggest that this species could be an eastern subspecies of M. leuckarti rather than a genetically isolated species.

    (3) younger synonym of M. aspericornis (Ghenne & Fiers, 2000)

    (4) younger synonym of M. papuensis Van de Velde, 1987 (Holynska, 2000)

    (5) synonym of M. edax (Reid & Moreno, 1999)

    (6) it is unambiguously a species belonging to the genus Thermocyclops (Holinska, in litt.)

    (7) Guo (2000) synonymized M. ruttneri with M. pehpeiensis.

    (8) = M. woutersi (Holynska, 1997)

    (9) The record of this species by Knott & Jasinska (1998) from Australia is questionable.


    The species M. cokeri and M. forbesi, by Nisa & Mahoon (1987) from Pakistan, are inadequately and incompletely described; the species M. albicans and the subspecies M. leuckarti deccanensis Lindberg, 1935 are questionable. All the above taxa should be considered as "nomina dubia" (Holynska, 1997; in litt.).


    Recently, evolutionary relationships were investigated in the genus Mesocyclops by Holynska (2006), and the following clades were pointed out: reidae-clade (M. reidae, M. chaci, M. yutsil); rarus-clade (Mesocyclops annae, M. pseudoannae, M. splendidus, M. rarus, M. paludosus, M. darwini, M. dayakorum); annulatus-clade (Mesocyclops intermedius, M. ellipticus, M. paranaensis, M. annulatus, M. tenuisaccus); meridianus-clade (Mesocyclops meridionalis, M. varius, M. venezolanus, M. brasilianus, M. pseudomeridianus, M. meridianus); major-clade (Mesocyclops major , M. pilosus, M. insulensis); dussarti-clade (M. dussarti, M. dadayi, M. isabellae, M. thermocyclopoides); pubiventris-clade (M. pubiventris, M. medialis, M. brooksi, M. notius).

    In the above analysys, however, "there were some components for which the procedure, regardless of the outgroup choice and/or character set, suggested different relationships. Basal relationships of Mesocyclops[between M. edax (North and Central America), the Neotropical species (M. longisetus, M. araucanus, M. evadomingoi, meridianus- and annulatus-clade), Old World group (P1 basipodite without medial spine) and the rarus-clade (Old World; P1 basipodite with medial spine) remained unresolved" (from Holynska, 2006).

    More recently Alekseev (2021) "divided the genus into four subgenera based on the morphological and habitat characteristics. The subgenus Neomesocyclops subgen. n. comprises species almost exclusively living in the New World. The single exception is M. (N.) tenuisaccus (Sars 1927), described from South Africa, which may prove to be a junior synonym of M. (N.) annulatus (Wierzejski 1892), from South America, and could have appeared in Africa as a result of anthropogenic invasion from the New World. The subgenus Tethymesocyclops subgen. n. includes species the distributions of which coincide with the borders of the former subcontinent Gondwana, or rather, the coast of the Tethys Sea, during the separation of Gondwana from Laurasia. These two subgenera contain species with a seta on the inner outgrowth of the basal segment of the first swimming leg pair. This seta is absent in all representatives of the nominative subgenus Mesocyclops s. str. with bare caudal rami and last thoracic somite, as well as in the new subgenus Pilosomesocyclops subgen. n., which has hairs on these structures. Most species of the latter two subgenera live in water bodies of the Old World. Along with a detailed characterization of the identified subgenera, a new species is described: Mesocyclops (Neomesocyclops) frankfiersi sp. n. As part of the revision, species in the two largest subgenera, Neomesocyclops subgen. n. and Pilosomesocyclops subgen. n., are further divided into three sections by the presence of hair-like structures on the last thoracic somite and/or caudal rami" (from Alekseev, 2021)



    KEY FOR AFRICAN SPECIES OF MESOCYCLOPS

    KEY FOR AUSTRALIAN SPECIES OF MESOCYCLOPS


    RELEVANT LITERATURE




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