SUPERFAMILY ECTINOSOMATOIDEA Sars, 1903

ECTINOSOMATIDAE Sars, 1903, Olofsson, 1917

Syn.: Ectinosomidae Sars, 1903 (1911); Olofsson, 1917; Monard, 1927; Rose, 1933; Lang, 1948; 1965; Gonzalez & Bowman, 1965;Zheng Zhong & al.,1984 (1989)


ECTINOSOMA Boeck, 1865 emend. Lang (1965)
syn. Helectinsoma Krichagin, 1877

Boeck (1865) established the genus Ectinosoma with the type-species E. melaniceps Boeck, 1865. Lang (1948) split the genus into the subgenera Halectinosoma, including species of Ectinosoma with one surface seta and 3 distal setae exopod P5, Ectinosoma embracing species with 4 distal setae exopod P5. In 1965 Lang "looked upon them as separate genera ... , since no transitional forms can exist between them ...". but sometimes it is difficult to decide whether there is a surface seta or not, as shown in the case of Halectinosoma porosum Wells, 1967. This species has a very peculiar P5, because the exopod is completely fused with the baseoendopod and there is no real surface seta but a seta on a lobule in distal position. This species shows pores on the cephalothorax that resemble the characteristic ushaped tubercles that can be found only in the Ectinosoma species. For this reason, H. porosum has to be transferred to the genus Ectinosoma.

Lang (1965) pointed out that tubercles can probably be found in all species of the genus Ectinosoma and that "pores of the very same appearance occur on all somites, the penultimate one excepted." Except E. soyeri Apostolov, 1975, all species of Ectinosoma have been shown to possess a tubercle on exopod P5. Most species are also reported to have tubercles on the abdomen or the whole body. The species of Halectinosoma, the sister group of Ectinosoma, have little round inconspicuous pores instead of tubercles. At the moment it is very difficult to determine Ectinosoma species. Very often there are no differences in the number of setae of the appendages and the distinctive features are very small. The drawings frequently are not good enough or lacking.

As Lang (1965) pointed out, the mandible carries very good diagnostic characters. The cutting edge is unique for a species in most cases. Unfortunately, the mandible is often not mentioned in the old descriptions! Mielke (1979) showed that characters of the antenna, female P5, male P5-P6, and the ornamentation of the abdominal somite reveal distinctive features. These become apparent when detailed descriptions are available. This also holds for the ornamentation of the body. The presence, number and distribution of rows of fine cuticular hairs, tubercles, sensillae and the shape of hyaline frills are constant in a species and provide many useful characters which can be seen in whole specimens without dissection. The form of the labrum also seems to be useful in distinguishing between the species, as in the case of E. papuarum and E. barbararum.

At present no revision of the genus is available, and "nothingis known about the phylogenetic relationships of this taxon (Gomez & Seifried, 2001).

The genus includes about 30 valid species and 13 species incertae or nomina nuda, some valid species probably being of doubtful position (Seifried, 1997; Seifried & Durbaum, 2000; Gomez & Seifried, 2001).


  • Ectinosoma acutorostratum Vervoort, 1962 [Melanesia; psammon]
  • Ectinosoma andamanica Rao, 1993 [Andaman; littoral sea waters]
  • Ectinosoma antarcticum Giesbrecht, 1902
  • Ectinosoma australe Brady, 1899
  • Ectinosoma barbararum Seifried, 1997[Papua, New Guinea; sublittoral sand]
  • Ectinosoma barbicauda Bozic, 1978 [Reunion island; brackish waters]
  • Ectinosoma breviarticulatum Lang, 1965 [USA; sea waters]
  • Ectinosoma californicum Lang, 1965 [USA; sea waters]
  • Ectinosoma canaliculatum Por (?)
  • Ectinosoma carnivora Seifried & Durbaum, 2000 [Bahamas; intertidal mudflat]
  • Ectinosoma compressum Sars, 1920 [north Europe; marine]
  • Ectinosoma couceiroi Jakobi & Nogueira, 1960
  • Ectinosoma christalii (?)
  • Ectinosoma veili Labbé, 1926
  • Ectinosoma curvifrons Sars G.O., 1927 (nomen nudum)
  • Ectinosoma dentatum Steuer, 1940 [Brazil, Bermuda, Ifaluk Atoll, Mediterranean and Marmara Seas; coastal lagoons] *
  • Ectinosoma diops Por (?)
  • Ectinosoma edwardsi (Richard, 1890) (syn. Ectinosoma edwardsi edwardsi (Richard, 1890)
  • Ectinosoma erithrops Brady, 1868
  • Ectinosoma finmarchicum Scot (?)
  • Ectinosoma herdmani T. & A . Scott, 1894 (?)
  • Ectinosoma inopinatum Por (?)
  • Ectinosoma internedium Marcus,1965 [Black Sea]
  • Ectinosoma ischnum Sars G.O., 1927 (nomen nudum)
  • Ectinosoma ghardaqesis (Noodt, 1964) [Roten Meeres; psammon]
  • Ectinosoma gracilicorne Brady, 1910
  • Ectinosoma henneguyi Labbé, 1927
  • Ectinosoma litorale (Noodt, 1958) [Canarie islands; psammon]
  • Ectinosoma mediterraneum Kunz, 1975 [France; coastal waters]
  • Ectinosoma melaniceps Boeck, 1864 [cosmopolitan;marine, fresh, brackish waters]
  • Ectinosoma mexicanum Gomez & Seifried, 2001 [Mexico; lagoon]
  • Ectinosoma nonpectinatum Mielke, 1979 [Galapagos; psammon]
  • Ectinosoma normani T. & A. Scott, 1894 [cosmopolitan, except Australia; marine sand]
  • Ectinosoma obtusum Sars, 1920 [Portugal, Israel; lake]
  • Ectinosoma papuarum Seifried, 1997 [Papua, New Guinea; sublittoral sand]
  • Ectinosoma paradentatum Bozic, 1965
  • Ectinosoma paranormani Lang, 1965 [USA; sea waters]
  • Ectinosoma pectinatum Mielke, 1979 [Galapagos; psammon]
  • Ectinosoma porosum (Wells, 1967) [Africa, Mozambico; psammon]
  • Ectinosoma porrectum Sars G.O., 1927 (nomen nudum)
  • Ectinosoma propinquum Scott T. & Scott A., 1896
  • Ectinosoma pruvoti Soyer, 1972 [west Mediterranean]
  • Ectinosoma pygmaeum Scott T. & Scott A., 1896
  • Ectinosoma reductum Bozic, 1955 [France; psammon]
  • Ectinosoma soyeri Apostolov, 1975 [Bulgarie; phreatic waters]
  • Ectinosoma scotti Brady, 1910
  • Ectinosoma spinipes Brady, 1868
  • Ectinosoma tegula Seifried, 1997 [Fiji islands; sublittoral sand]
  • Ectinosoma tenerum Sars G.O., 1920
  • Ectinosoma tenuipes T. & A. Scott, 1894 [north Europe; marine]
  • Ectinosoma tholomiges Jakobi, 1954
  • Ectinosoma tholophilos Jakobi, 1954
  • Ectinosoma veili Labbé, 1926
  • Ectinosoma vervoorti Soyer, 1972 [west Mediterranean]
  • Ectinosoma virginensis Coull, 1971 [Virgin Islands; psammon]

    * synonyms: Ectinosoma couceiroi Jakobi & Nogueira, 1960 (Lang, 1965); Ectinosoma paradentatum Bozic, 1965 (in Wells & Rao, 1987)

    Species incertae or nomina nuda: E. n.sp. aff. compressum Sars sensu Klie (1949); E. curvifrons Sars, 1927 (in Lang, 1948); E. porrectum 1927 (in Lang, 1948); E. ischnum (in Lang, 1948); E. henneguyi Labbé, 1926; E. spec.I-V Mielke, 1979; E. spec. Gruppe A Mielke, 1979 (not in Bodin, 1988); E. tholomiges Jakobi, 1954; E. tholophilos Jakobi, 1954.


    | RELEVANT LITERATURE AND KEYS |




    HALECTINOSOMA Lang, 1944, 1965

    syn. Pararenosetella Lang, 1944


    The genus Halectinosoma Vervoort, 1962 is a large group composed of about 70 nominal species within the family Ectinosomatidae (Soyer 1972, Huys & Bodin 1997, Cl´ement & Moore 2000, Karanovic & Pesce 2001, Wells 2007, Huys 2009, Boxshall et al. 2010, Sua´ rez-Morales & Fuentes-Rein´es 2015). Initially it was described as a subgenus of the genus Ectinosoma, with the following short diagnosis (Lang, 1944): "Exp. P.5 mit 3 Rand- and 1 Flachenborste, welch letztere evtl. fehlen kann." Exactly the same was repeated later (Lang, 1948), but unfortunately Lang too omitted to designate the type species. This was done by Vervoort (1962), who designated Ectinosoma sarsi Boeck, 1872. Lang (1965) erected Halectinosoma at generic level, although he did not mention Vervoort's designation of the type species. Unfortunately Halectinosoma sarsi (Boeck, 1972) was placed as incertae sedis by Clement & Moore (1995), so the genus is now without the type species.

    Lang's (1944, 1948) division of Halectinosoma into the "sarsi"group (where the third exopodite segment of second, third and fourth swimming legs carried 3 outer spines) and the "curticorne"-group (where the third exopodite segment of at least fourth swimming leg bore 2 outer spines) was later (Lang, 1965) considered unworkable.

    Although the detailed morphology is quite conservative within the genus some of its species have obviously been misplaced and should be referred to other genera. One of them, H. porosum Wells, 1967, was formally transferred to the genus Ectinosoma by Huys & Bodin (1997); some other probably belong to yet undescribed genera. However, the systematic of the genus Halectinosoma is problematic and species identification is very difficult due to the inadequacy of most of the descriptions in the literature (Clement & Moore, 1995). This genus is indeed much closer to the genus Pseudobradya than to Ectinosoma, from which it was properly divided.

    One of the characters that is traditionally used to distinguish Halectinosoma from Pseudobradya is the condensed maxillary endopodite in the former (Huys & Bodin, 1997). Also, a position of the inner setae on the maxillary basis can be used as a very good distinguishing character between these two genera (they are always situated near the top of the basis in Pseudobradya). Probably the subgeneric status of Halectinosoma (of the genus Pseudobradya) could be more appropriate, but its generic status should be retained because it has already been the subgenus (of the genus Ectinosoma) and this could cause great taxonomic confusion.

    "Species of Halectinosoma Lang, 1944 are often dominant members of the harpacticoid copepod assemblage of marine sediments and yet their identification is notoriously difficult.

    Differences between species within this large genus of 58 species (Huys et al., 1996) are often subtle and a lack of appreciation of this subtlety has caused much taxonomic confusion and the production of descriptions which, in many cases, do not permit the accurate identification of species. Clement & Moore (1995) commented a revision of Halectinosoma with a reappraisal of H. sarsi (Boeck, 1872) and descriptions of 11 related species. In this paper was reexamined H. herdmani (Scott & Scott, 1894) and described a group of related species which share the distinctive, elongate body shape of H. herdmani.

    Several of these species can be found living sympatrically, generally in sandy sediments, and it is highly probable that considerable misidentification of these species has occurred in the past" (from: Clement & Moore, 2000).


  • Halectinosoma abrau (Kritzagin, 1877) [Europe, Asia; surface and ground waters]
  • Halectinosoma abyssicola Bodin, 1968 [France; sea waters]
  • Halectinosoma angulifrons (Sars, 1919) [Europe, isle of Man; psammon]
  • Halectinosoma arangureni Suarez-Morales & Fuentes Reines, 2015 [Colombia; lagoons]
  • Halectinosoma arenicola (Rouch, 1962) [Brazil, Japan; marine interstitial]
  • Halectinosoma argyllensis Clement & Moore, 1995 [Scotland, England; psammon]
  • Halectinosoma armiferum (T. & A. Scott, 1894) [Norway, Scotland; psammon]
  • Halectinosoma barroisi (Richard, 1893 [Caspian sea; Palestina, marine] *
  • Halectinosoma bodotriaensis Clement & Moore, 2000 [British Isles; marine]
  • Halectinosoma brevirostre (Sars, 1904) [Norway; sea waters]
  • Halectinosoma britannicum Clement & Moore, 2000 [British Isles; marine]
  • Halectinosoma brunneum (Brady, 1905) [north Europe; salt-water ponds, marine]
  • Halectinosoma canaliculatum (Por, 1964) [north Europe,Mediterranean, Israel, Helgoland; sublittoral sandy muds] [syn. H. sarsi sensu T. & A. Scott (1894)]
  • Halectinosoma candelabrum Soyer, 1970
  • Halectinosoma chislenki Clement & Moore, 1995 [Arctic, Beaufort sea]
  • Halectinosoma chrystalli (T. Scott, 1894) [Canada, north Europe, Africa; psammon?]
    [syn. H. propinquum T. & A. Scott, 1894]
  • Halectinosoma clavatum (Sars, 1920) [north Europe; psammon]
  • Halectinosoma concinnum Akatova, 1935 [Caspian sea]
  • Halectinosoma cooperatum Bodin, Bodiou & Soyer, 1970 [west Mediterranean]
  • Halectinosoma crenulatum Clement & Moore, 1995 [Europe; sea waters]
  • Halectinosoma curticorne Boeck, 1872 [cosmopolitan, except Australia?; marine, mud, brackish waters]
  • Halectinosoma denticulatum Clement & Moore, 1995 [north Europe; sea waters]
  • Halectinosoma dimorphum Coull, 1970 [Barbados; Fore reef area]
  • Halectinosoma diops (Por, 1964) [Mediterranean; eulittoral sands]
  • Halectinosoma distinctum (Sars, 1920) [Norway; psammon]
  • Halectinosoma elongatum (Sars, 1904) [Europe; marine]
  • Halectinosoma erythops (Brady & Robertson, 1875) [Europe; marine interstitial]
  • Halectinosoma foveolata Jong Guk et al., 2017 [ sandy sediments; Korea]
  • Halectinosoma finmarchicum (T. Scott, 1903) [URSS, Alaska; sea waters] *
  • Halectinosoma fusiforme Wells, 1967 [Africa, Mozambico; psammon]
  • Halectinosoma fusum Wells, 1967 [Africa, Mozambico; psammon]
  • Halectinosoma gascognense Bodin, 1968 [France; sea waters]
  • Halectinosoma gothiceps (Giesbrecht, 1881) [Europe; marine]
  • Halectinosoma gracile (T. & A. Scott, 1894) [north Europe; marine, coarse sand]
  • Halectinosoma herdelongata Marcus, 1967 [Black sea] **
  • Halectinosoma herdmani (T. & A. Scott, 1894) [Europe, north Africa; sublittoral sandy muds, amongst algae]
  • Halectinosoma huysi Clement & Moore, 2000 [British Isles; marine]
  • Halectinosoma hydrofuge Wells, Hicks & Coull, 1982 [New Zealand; aestuarine waters]
  • Halectinosoma inhacae Wells, 1967 [Africa, Mozambico; psammon]
  • Halectinosoma inopinatum (Por, 1964) [Gulf of Haifa; marine interstitial]
  • Halectinosoma intermedium (Nicholls, 1940) [St. Lawrence; psammon]
  • Halectinosoma islandicum Apostolov 2007 [Iceland; brackish waters]
  • Halectinosoma itoi Clement & Moore, 2000 [British Isles; marine]
  • Halectinosoma japonicum (Miura, 1964) [Japan; fresh ground waters, wells]
  • Halectinosoma kliei Clement & Moore, 2007 [North Atlantic]
  • Halectinosoma kunzi Lang, 1965 [USA; marine tidal pools]
  • Halectinosoma langi Wells, 1967 [Africa, Mozambico; psammon]
  • Halectinosoma latisetifera Clement & Moore, 2007 [North Atlantic]
  • Halectinosoma limnophilum (Sterba, 1968) (= Arenosetella limnophila Sterba, 1968)
  • Halectinosoma littorale (Nicholls, 1940) [USA, Europe; marine psammon]
  • Halectinosoma longicorne (T. & A. Scott, 1894) [Scotland, France; marine interstitial]
  • Halectinosoma longisetosum Lang, 1965 [USA; marine tidal pools]
  • Halectinosoma mandibularis Clement & Moore, 2007 [North Atlantic]
  • Halectinosoma monardi Soyer, 1973 [west Mediterranean; brackish waters]
  • Halectinosoma mixtum (Sars, 1904 [north Europe; marine] *
  • Halectinosoma neglectum (Sars, 1904) [Europe, Iceland, north America, psammon]
  • Halectinosoma munmui Bang, 2020 [Korea]
  • Halectinosoma oblongum (Kunz, 1949) [Helgoland; marine interstitial]
  • Halectinosoma ornatum Lang, 1965 [USA; marine tidal pools]
  • Halectinosoma otakoua Wells, Hicks & Coull, 1982 [New Zealand; aestuarine waters]
  • Halectinosoma paradistinctum Soyer, 1972
  • Halectinosoma parejae Sciberras et al., 2017 [Argentina]
  • Halectinosoma paragothiceps Clement & Moore, 2007 [North Atlantic]
  • Halectinosoma paraspinicauda Bodin, 1979 [France; brackish waters]
  • Halectinosoma perforatum Ito, 1981 [Japan; sea waters]
  • Halectinosoma pilosum Clement & Moore, 2000 [British Isles; marine]
  • Halectinosoma porosum Wells, 1967
  • Halectinosoma propinquum (T. & A. Scott, 1896) (syn. Halectinosoma chrystali (Scott T., 1894)
  • Halectinosoma proximum (Sars, 1919) [north Europe; psammon]
  • Halectinosoma pseudosarsi Clement & Moore, 1995 [north Europe, Canada; sea waters]
  • Halectinosoma pterinum Moore, 1974 [isle of Man; psammon]
  • Halectinosoma rouchi Soyer, 1973 [west Mediterranean; brackish waters]
  • Halectinosoma sarsii (Boeck, 1872)
  • Halectinosoma similidistinctum Lang, 1965 [USA; marine interstitial]
  • Halectinosoma smirnovi (Chislenko, 1967)
  • Halectinosoma spinicauda Wells, 1961 [Scilly island; marine interstitial]
  • Halectinosoma spinipes (Brady, 1880)
  • Halectinosoma tenerum (Sars, 1920) [north Europe; marine] (syn. Ectinosoma tenerum Sars G.O., 1920)
  • Halectinosoma tenuireme (T. & A. Scott, 1894) [Europe; marine interstitial]
  • Halectinosoma travei Soyer, 1973 [west Mediterranean; brackish waters]
  • Halectinosoma uniarticulatum Borutzky, 1972 [Asia, Kyzylkum; fresh waters]
  • Halectinosoma unicum Lang, 1965 [USA; marine interstitial]
  • Halectinosoma valeriae Soyer, 1973 [west Mediterranean; brackish waters]
  • Halectinosoma winonae Coull, 1975 [USA; brackish waters]
  • Halectinosoma wiesei (Smirnov, 1932)
  • Halectinosoma ssp. Werner & Martinez-Arbizu, 1999 [Arctic ocean]
  • Halectinosoma sp. Schizas & Shirley, 1996 [Alaska; psammon]


    The species H. herdmani, H. bodotriaensis, H. pilosum, H. britannicum, H. itoi, H. huysi, H. tenerum and H. kunzi are referred to the «herdmani species group», on the basis of close similarity (Clement & Moore, 2000).

    ** syn. of H. herdmani (Clement & Moore, 2000)

    Incertae sedis: H. sarsi (Boeck, 1872) (Clement & Moore, 1995).


    A revision of the genus Halectinosoma


    | RELEVANT LITERATURE AND KEYS |




    PSEUDECTINOSOMA Kunz, 1935

    Kunz (1935) created the genus Pseudectinosoma to accomodate harpacticoids from benthic biotopes of Kiel Bay (Germany) close to Sigmatidium Giesbrech, 1881. Pseudectinosoma is a poorly diversified genus among the Ectinosomatidae, its geographical distribution suggesting an ancient origin (Galassi, 1997; Galassi & De Laurentiis, 1997).


  • Pseudectinosoma galassiae Karanovic, 2006 [Australia; rivers]
  • Pseudectinosoma janineae Galassi, Dole-Olivier & De Laurentiis, 1999 [France; phreatic, hyporheic]
  • Pseudectinosoma kunzi Galassi 1997 [Italy; phreatic lake]
  • Pseudectinosoma minor Kunz, 1935 [Baltic Sea, Germany, USA; brackish marsh]
  • Pseudectinosoma reductum Galassi & De Laurentiis, 1997 [Italy; karstic spring]
  • Pseudectinosoma vandeli (Rouch, 1969) [Europe; karstic springs]
  • Pseudectinosoma sp. (unpublished) [central Italy, krenal]



    Distribution of the genus Pseudectinosoma (after Galassi, 1997 mod. and Karanovic, 2006):
    1. P.minor; 2. P. vandeli; 3. P. kunzi; 4. P. reductum; 5. P. janineae; 6. P. galassiae



    | PHYLOGENY AND BIOGEOGRAPHY OF THE GENUS PSEUDECTINOSOMA |

    | RELEVANT LITERATURE |




    NOODTIELLA Wells, 1965 (= Lineosoma Wells, 1965, by Kihara & Huys, 2009)


  • Noodtiella arenosetelloides (Noodt, 1958) [Canary islands, India; marine interstitial]
  • Noodtiella chilensis (Mielke, 1987)
  • Noodtiella coquinbensis Mielke, 1987 Mielke, 1987 [Chile; marine interstitial]
  • Noodtiella enertha Lindgren, 1975
  • Noodtiella frequentior Mielke, 1979 [Panama, Galapagos; marine interstitial]
  • Noodtiella gracile Mielke, 1975 [Germany; marine interstitial]
  • Noodtiella enertha Lindgren. 1975 [USA; sandy beaches]
  • Noodtiella hoodensis Mielke, 1979 [Panama, Galapagos; sandy beaches, interstitial]
  • Noodtiella intermedia Wells, 1967
  • Noodtiella iscensis (Wells, 1965)
  • Noodtiella larinconadensis Mielke, 1987 Mielke, 1987 [Chile; marine interstitial]
  • Noodtiella lusitanica Wells, 1965 [Portugal; marine interstitial]
  • Noodtiella mielkei Wells & Rao, 1987 [Andaman, Nicobar; marine interstitial]
  • Noodtiella ornamentalis Wells & Rao, 1987 [Andaman, Nicobar; marine interstitial]
  • Noodtiella pacifica Mielke, 1987 [Chile; marine interstitial]
  • Noodtiella pectinata (Chappuis, 1954)
  • Noodtiella problematica (Rouch, 1962) [Brazil; marine interstitial]
  • Noodtiella tabogensis Mielke, 1981 [Panama; marine interstitial]
  • Noodtiella toukae Mitwally & Montagna, 2001 [Egypt; sandy beaches] (= Noodtiella pectinata (Chappuis, 1954), synonomized by Kihara & Huys, 2009)
    [syn. of N. pectinata, according to Kihara & Huys (2009)]
  • Noodtiella wellsi Apostolov, 1974 [Bulgaria; marine interstitial]


    | KEY TO SPECIE OF NOODTIELLA |




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