MIRACIIDAE Dana, 1846 (Willen, 2002) (syn. DIOSACCIDAE Sars, 1906)

DIOSACCINAE Sars,1906


STENHELIINAE Brady, 1880

"The subfamily Stenheliinae Brady, 1880 is currently recognised as one of three well-defined suprageneric groups within the second largest harpacticoid family Miraciidae Dana, 1846, beside the nominotypical subfamily and Diosaccinae Sars, 1906 (see Willen 2000; Boxshall & Halsey 2004; Wells 2007; Huys & Mu 2008). Stenheliins are common inhabitants of the marine benthos" (from Karanovic & Kim. 2014)

"The subfamily Stenheliinae Brady currently accommodates 14 genera: Anisostenhelia Mu & Huys, 2002 (monotypic), Beatricella Scott, 1905 (monotypic), Cladorostrata Tai & Song, 1979 (two species), Delavalia Brady, 1869 (50 species), Itostenhelia Karanovic & Kim, 2014 (two species), Lonchoeidestenhelia Gómez, 2020 (monotypic), Melima Por, 1964 (five species), Muohuysia Özdikmen, 2009 (monotypic), Onychostenhelia Itô, 1979 (two species), Pseudostenhelia Wells, 1967 (three species), Stenhelia Boeck, 1865 (eight species), Wellstenhelia Karanovic & Kim, 2014 (nine species), Wellstenvalia Gómez & Cruz-Barraza (monotypic), and Willenstenhelia Karanovic & Kim, 2014 (six species) (Gómez 2021; Gómez & Cruz-Barraza 2021; Karanovic & Kim 2014; Wells 2007). Stenheliinae is a predominantly shallow-water taxon and only a few representatives (Wellstenvalia wellsi Gómez & Cruz-Barraza, 2021, and some species of Delavalia) are known from the deep sea (>200 m depth) (see Gómez & Cruz-Barraza 2021, Gómez 2021)" (from Gomez, 2021)


Stenhelia Boeck, 1864

Delavalia Brady, 1869

Anisostenhelia Mu & Huys, 2002

Archaeohuysia Gómez, 2021

Beatricella Mu & Huys, 2002

Diarthropodella Gómez, 2021

Lonchoeidestenhelia Gomez, 2020

Muohuysia Özdikmen, 2009

(syn. Hicksia Mu & Huys, 2002 )

Pseudostenhelia Wells, 1967

Cladorostrata Shen & Tai, 1963

Onychostenhelia Ito, 1979

Melima Coull, 1971

Wellstenhelia Karanovic & Kim, 2014

Willenstenhelia Karanovic & Kim, 2014

Wellstenvalia Gómez & Cruz-Barraza, 2021

Itostenhelia Karanovic & Kim, 2014

Archaeohuysia Gómez, 2021

Diarthropodella Gómez, 2021

ARCHAEOHUYSIA Gómez, 2021

  • Archaeohuysia huysi Gómez, 2021 [California]

    DIARTHROPODELLA Gómez, 2021

  • Diarthropodella prima Gómez, 2021 [California]
  • Diarthropodella secunda Gómez, 2021 [California]


    ITOSTENHELIA Karanovic & Kim, 2014

  • Itostenhelia polyhymnia Karanovic & Kim, 2014
  • Itostenhelia golikovi (Chislenko, 1978) Karanovic & Kim, 2014 [South Korea]


    LONCHOEIDESTENHELIA Gomez, 2020

  • Lonchoeidestenhelia prote Gomez, 2020 [Mexico]


    WELLSTENVALIA Gómez & Cruz-Barraza, 2021

  • Wellstenvalia wellsi Gómez & Cruz-Barraza, 2021

    WELLSTENHELIA Karanovic & Kim, 2014

  • Wellstenhelia bocqueti (Soyer, 1972)
  • Wellstenhelia calliope Karanovic & Kim, 2014 [South Korea]
  • Wellstenhelia clio Karanovic & Kim, 2014 [South Korea]
  • Wellstenhelia erato Karanovic & Kim, 2014 [South Korea]
  • Wellstenhelia euterpe Karanovic & Kim, 2014 [South Korea]
  • Wellstenhelia euterpoides Gomez & Barraza 2021 [California]
  • Wellstenhelia hanstroemi (Lang, 1948)
  • Wellstenhelia melpomene Karanovic & Kim, 2014 [South Korea]
  • Wellstenhelia qingdaoensis (Ma & Li, 2011) Karanovic & Kim, 2014 [South Korea]


    WILLENSTENHELIA Karanovic & Kim, 2014

  • Willenstenhelia mirabiilipes Ma & Li 2018 [China Sea]
  • Willenstenhelia reducta Gomez, 2020 [Mexico]
  • Willenstenhelia thalia Karanovic & Kim K., 2014
  • Willenstenhelia urania Karanovic & Kim K., 2014
  • Willenstenhelia terpsichore Karanovic & Kim K., 2014


    STENHELIA Boeck, 1865

    Willen (2003) hypothesized that deep sea colonization within the Stenheliinae and the genus Stenhelia occurred independently at least twice in two different lineages: S. normani group, characterized by the male P5 expod being fused to the basis and lacking two outer spines (S. giesbrechti T. and A. Scott, 1896; S. gundulae Willen, 2003; , S. tethysensis Monard, 1928; S. normani T. Scott, 1905 ; S. polluta Monard, 1928; S. acutirostris Willey, 1935) and S. longicaudata group, characterized by the apomorphic shape of the P1 (S. noodti Schriever, 1982; S. islandica Schriever, 1982; S. lima Becker and Schriever, 1979 ; S. diegensis Thistle and Coull, 1979)

    The traditional subgeneric division of Stenhelia is abandoned by Mu & Huys (2001)since both subgenera, Stenhelia (Stenhelia) and Stenhelia (Delavalia), are polyphyletic taxa. The latter is tentatively upgraded to generic rank pending a phylogenetic analysis of the Stenheliinae


  • Stenhelia (Stenhelia) asetosa Thistle & Coull, 1979 [Pacific] ( syn. Anisostenhelia asetosa (Thistle & Coull, 1979)
  • Stenhelia (Stenhelia) aemula aemula (T. Scott, 1893) [north Europe; psammon] (sin. Beatricella aemula (Scott T., 1893)
  • Stenhelia (Stenhelia) aemula bifida Lang, 1936 [Sweden, marine]
  • Stenhelia (Stenhelia) blanhardi Scott, 1895 [Scotland] (syn. Typhlamphiascus blanchardi (Scott T. & Scott A., 1895)
  • Stenhelia (Stenhelia) curviseta Lang, 1936 [Spitzbergen; marine]
  • Stenhelia (Stenhelia) diegensis Thistle & Coull, 1979 [Pacific]
  • Stenhelia (Stenhelia) divergens (Nicholls, 1940) [St. Lawrence; sea waters]
  • Stenhelia (Stenhelia) gibba Boeck, 1865 [palaearctic; brackish waters, psammon]
  • Sienhelia longirostris (?)
  • Stenhelia (Stenhelia) peniculata Lang, 1965 [USA, costal waters]
  • Stenhelia (Stenhelia) proxima Sars, 1906 [north Europe; sea waters]
  • Stenhelia (Stenhelia) pubescens Chislenko, 1978 (according to Karanovic et a., 2015) [Japan; coastal waters]
  • Stenhelia (Stenhelia) sheni Mu & Huys, 2002 [China; Bohai Sea]
  • Stenhelia taiae Mu & Huys, 2002 (according to Karanovic et a., 2015) [China; Bohai Sea]
  • Stenhelia (Stenhelia) xilophila Hicks, 1988 [New Zealand; coastal waters] (syn. Hicksia xylophila (Hicks, 1988); Muohuysia xylophila (Hicks, 1988)

  • Stenhelia (Delavalia) acutirostris Willey, 1932 [?; ?]
  • Stenhelia (Delavalia) adriatica Marinov & Apostolov, 1981 [Adriatic sea]
  • Stenhelia (Delavalia) andamanica Rao, 1993 [Andaman; coastal waters]
  • Stenhelia (Delavalia) arctica T. Scott, 1898 [Arctic Ocean]
  • Stenhelia (Delavalia) arenicola Wilson, 1932 [north America, marine]
  • Stenhelia (Delavalia) asetosa Gomez, 2021
  • Stenhelia (Delavalia) bermudensis Coull, 1969 [Bermuda; coastal waters]
  • Stenhelia (Delavalia) bifidia Coull, 1976 [USA; coastal waters]
  • Stenhelia (Delavalia) bisetosa (Coull, 1971) [USA; meiobenthos]
  • Stenhelia (Delavalia) boqueti Soyer, 1972 [west Mediterranean] (syn. Wellstenhelia bocqueti (Soyer, 1972)
  • Stenhelia (Delavalia) breviseta Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia (Delavalia) californiensis Gomez, 2021
  • Stenhelia (Delavalia) caulerpae Por, 1964 [Israel; coastal waters] (syn. Melima caulerpae Por, 1964)
  • Stenhelia (Delavalia) clavus Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia (Delavalia) coineauae Soyer, 1972 [west Mediterranean]
  • Stenhelia (Delavalia) confluens lang, 1948 [north Europe; sea waters]
  • Stenhelia (Delavalia) cornuta Lang, 1936 [north Europe; sea waters]
  • Stenhelia (Delavalia) diegensis (Thistle & Coull, 1979)
  • Stenhelia (Delavalia) elizabethae Por, 1960 [Schwarzen; coastal waters]
  • Stenhelia (Delavalia) fustiger Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia (Delavalia) giesbrechti T. & A. Scott, 1896 [Europe; marine]
  • Stenhelia (Delavalia qingdaoensis Ma & Lee, 2011 [Yellow Sea]
  • Stenhelia (Delavalia) golikovi Chislenko, 1978 [Japan; sea waters]
  • Stenhelia (Delavalia) gundulae Wileln, 2003 [New Guinea; marine]
  • Stenhelia (Delavalia) hanstromi Lang, 1948 [north Europe; marine]
  • Stenhelia (Delavalia) hirtipes Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia (Delavalia) krisunensis Radhakriscia Ranga Reddy, 1878 [India]
  • Stenhelia (Delavalia) incerta Por, 1964 [Israel: sea waters]
  • Stenhelia (Delavalia) indica Krishnaswamy, 1957 [Madras; coastal waters]
  • Stenhelia (Delavalia) inopinata Scott, 1902 [India, Suez; coastal waters, lakes]
  • Stenhelia (Delavalia) intermedia Marinov & Apostolov, 1981 [Adriatic sea]
  • Stenhelia (Delavalia) islandica Schriever, 1982 [north Atlantic]
  • Stenhelia (Delavalia) latioperculata Ito, 1981 [Japan; sea waters]
  • Stenhelia (Delavalia) latipes Lang, 1965 [USA; coastal waters]
  • Stenhelia (Delavalia) latisetosa Sewell, 1940 [India?; sea waters]
  • Stenhelia (Delavalia) lima Becker & Schriever, 1979 [Helgoland?]
  • Stenhelia (Delavalia) longicaudata longicauda Boeck, 1872 [north Europe; sea waters]
  • Stenhelia (Delavalia) longicaudata finmarchica T. Scott, 1903 [north Europe; marine]
  • Stenhelia (Delavalia) longifurca Sewell, 1934 [India; brackish waters]
  • Stenhelia (Delavalia) longipilosa Lang, 1965 [USA; coastal waters]
  • Stenhelia (Delavalia) madrasensis Wells, 1971 [India; marine psammon]
  • Stenhelia (Delavalia) magnacaudata Monard, 1928 [France; marine]
  • Stenhelia (Delavalia) mastigochaeta Wells, 1965 [Scotland; marine muds]
  • Stenhelia (Delavalia) minuta Scott, 1902 [Suez, brackish waters]
  • Stenhelia (Delavalia) mixta Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia neglecta Norman ^ Scott (?)
  • Stenhelia (Delavalia) noodti Schriever, 1982 [north Atlantic]
  • Stenhelia (Delavalia) normani normani Scott, 1905 [Suez, France, brackish waters]
  • Stenhelia (Delavalia) normani polluta Monard, 1937 [Europe?; marine]
  • Stenhelia (Delavalia) nuwukensis Wilson, 1965 [Alaska; coastal waters]
  • Stenhelia (Delavalia) oblonga Lang, 1965 [USA, costal waters]
  • Stenhelia (Delavalia) ornementalia Shen & Tai, 1965 [China; brackish waters]
  • Stenhelia (Delavalia) ovalis Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia (Delavalia) palustris palustris (Brady, 1868) [Europe; brackish waters]
  • Stenhelia (Delavalia) palustris bispinosa Bodin, 1970
  • Stenhelia (Delavalia) paraclavus Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]
  • Stenhelia (Delavalia) pygmaea (Brady, 1905) (syn. Amphiascoides pygmaea (Norman & Scott T., 1905)
  • Stenhelia (Delavalia) polluta Monard, 1928 [Andaman, Nicobar, coastal waters]
  • Stenhelia (Delavalia) profunda Gomez, 2021
  • Stenhelia (Delavalia) qingdaoensis Ma & Lee, 2011 [Yellow Sea]
  • Stenhelia (Delavalia) reducta Gomez, 2021
  • Stenhelia (Delavalia) reflexa Brady & Robertson, 1875 [north Europe; sea waters]
  • Stenhelia (Delavalia) saharae Marinov & Apostolov, 1985 [Spanish Sahara; coastal waters]
  • Stenhelia (Delavalia) schminkei (Willen, 2002) [New Guinea; marine]
  • Stenhelia sheni Mu & Huys, 2002 [China]
  • Stenhelia simulans Norman & Scott (?)
  • Stenhelia (Delavalia) stephensoni Greenwood & Tucker, 1984 [Australia; brackish waters]
  • Stenhelia taiae Mu & Huys, 2002 [China]
  • Stenhelia (Delavalia) tethysensis Monard, 1928 [Europe, Tunisie; brackish waters]
  • Stenhelia (Delavalia) truncatipes Sewell, 1940 [India; sea waters]
  • Stenhelia (Delavalia) unisetosa Wells, 1967 [Africa; coastal waters]
  • Stenhelia (Delavalia) valens Wells & Rao, 1987 [Andaman, Nicobar; coastal waters]

    SPECIES INCERTAE SEDIS

  • Delavalia pygmaea Brady, 1905
  • Stenhelia glacialis Brady, 1910

    SPECIES INQUIRENDAE

  • Stenhelia brevicornis Herdman 1902 (?)
  • Stenhelia knoxi Herdman 1901 (?)
  • Stenhelia dentipes Herdman 1901 (?)
  • Styenhelia gracilicaudata Herdman 1921 (?)
  • Stenhelia longicornis Herdman 1901 (?)
  • Stehelia perplexa Herdman 1902 (?)
  • Stenhelia reflexa (Brady) sensu Bodin, 1964
  • Stenhelia minuta (A. Scott) sensu Por, 1964
  • Stenhelia normani (T. Scott) sensu Gurney, 1927
  • Stenhelia aemula (T. Scott) sensu Marinov, 1977
  • Stenhelia varians (?)


    Recently, Mu & Huys (2002) upgraded the subgenera Stenhelia and Delavalia to generic rank


    | REVIEW OF THE GENUS AND KEY TO SPECIES |



    MELIMA Coull, 1971


  • Melima bisetosa Coull, 1971
  • Melima caulerpae Por, 1964
  • Melima indica (Krishnaswamy, 1957)
  • Melima ovalis (Wells & Rao. 1987)
  • Melima papuaensis Willen, 2002



    MUOHUYSIA Özdikmen, 2009

    (syn. HICKSIA. Mu & Huys, 2002)


    "The genus Hicksia was described by Delgado (1904) with the type species Hicksia elvensis Delgado, 1904 in Trilobita from Haut Alemtejo, Portugal. Subsequently, the monotypic copepod genus Hicksia was erected by Mu & Huys (2002) with the type species Hicksia xylophila (Hicks, 1988) in Crustacea. Thus the genus name Hicksia Mu & Huys, 2002 is a junior homonym of the valid genus name Hicksia Delgado, 1904" (Özdikmen, 2009). According to this Author Hicksia Mu & Huys, 2002 should be replaced with the new name Muohuysia, as a replacement name.

  • Muohuysia xylophila (Hicks, 1988)


    BEATRICELLA Sars, 1906

  • Beatricella calidafornax Gómez, 2021
  • Beatricella aemula (Sars, 1906)
  • Beatricella mimica (Scott T., 1897) (syn. Stenhelia gibba Boeck, 1865)


    ANISOSTENHELIA Mu & Huys, 2002


  • Anisostenhelia asetosa (Thistle & Coull ,1979) Mu & Huys, 2002





    MIRACIINAE Dana, 1846


    Distioculus Boxshall, 1979

    Miracia Boxshall, 1979

    Oculosetella Boxshall, 1979

    Macrosetella Sewell, 1940




    DISTIOCOLUS Boxshall, 1979

  • Distiocolus minor Huys & Böttger-Schnack, 1994


    MACROSETELLA Sewell, 1940 (syn, Dwightia Wilson C.B., 1924;Setella Dana, 1846)

  • Macrosetella acicula (Dana, 1847)
  • Macrosetella crassicornis (Dana, 1847)
  • Macrosetella longicauda (Dana, 1847)
  • Macrosetella norvegica (Boeck, 1865) (syn. Microsetella norvegica (Boeck, 1865)
  • Macrosetella oculata (Sars G.O., 1916)
  • Macrosetella gracilis Huys & Böttger-Schnack, 1994
  • Macrosetella messinensis (Claus, 1863)
  • Macrosetella tenuicornis (Dana, 1847)
  • Macrosetella tenuis (Lubbock, 1860)


    MIRACIA Boxshall, 1979

  • Miracia efferata Huys & öttger-Schnack, 1994
  • Miracia gracilis Dana, 1849 (=Oculosetella gracilis (Dana, 1849)
  • Miracia minor Scott T., 1894 (= Distioculus minor (Scott T., 1894)


    OCULOSETELLA Boxshall, 1979

  • Oculosetella gracilis Boxshall, 1979



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