SUPERFAMILY THALESTROIDEA Sars, 1905

MIRACIIDAE Dana, 1846
(syn. Diosaccidae Sars, 1906)

Subfamily Diosaccinae Sars, 1906


AMPHIASCOIDES Nicholls, 1941 [syn. AMPHIASCELLA Lang, 1944]


The genus Amphiascoides was considered a member of the family Diosaccidae Sars,1906 since its creation in 1941; however, a recent revision by Willen (2002) synonymized Diosaccidae with the older (1846) family Miraciidae. Therefore, all the genera previously contained in this family, including Amphiascoides, now belong to the Miraciidae.

When first erected by Nicholls (1941) as the morphological link between Schizopera and Amphiascus, the genus Amphiascoides contained 20 species plus at least one more: A. brevifurca, of which he had not enough data at that time but that is currently known to be a valid species (Bodin 1997).

Later on, Lang (1948, 1965) described several other species assignable to this genus and recognized about 25 nominal species.

Lotufo & Fleeger (1995) considered that the number of species known was around 20. In the most recent and comprehensive revision of the marine harpacticoids by Bodin (1997), the number of species in Amphiascoides was reduced to only 16, but this catalog did not include four species of Amphiascella Lang, 1944, a genus synonymized to Amphiascoides by Lang (1948).

These species are: A. littoralis (T. Scott, 1903), A. neglectus (Norman & Scott, 1905), A. proximus (T. Scott, 1914), and A. sterilis (Monard, 1926). Amphiascoides arabicus, described by Noodt (1964), was considered by Lang (1965) as belonging to another genus; there are still other nominal species related to this group whose taxonomic status is still uncertain (see Bodin 1997).

Historically, the lack of information on basic morphology and of the key characters for many species has generated hesitation among taxonomists, particularly in classifying closely related forms (Suarez-Morales & Avilés Torres, 2003).

The most recent description of a species of this genus was published by Lotufo & Fleeger (1995) for A. atopus Lotufo & Fleeger, 1995, found in bioassay cultures, but with an uncertain geographical or ecological origin. These authors sorted out two species groups containing each about half the number of the 20 species they included in their analysis.

One group has 7 elements on the third exopodal segment of the fourth swimming leg, and the other only 6. They assigned their new species to the first one, sharing this character with at least 10 other species: Amphiascoides brevifurca (Czernivaski, 1886), A. breviarticulatus Kunz, 1983, A. bulbiseta Pallares 1975a, A. dimorphus Lang, 1965, A. koltuni Kunz, 1983, A. lancisetiger Lang, 1965, A. neglectus (Norman & Scott, 1905), A. nichollsi Lang, 1965, A. petkovskii Lang, 1965, and A. subdebilis (Willey, 1935) [from: Suarez-Morales & Aviles-Torres, 2003]


  • Amphiascoides arabicus Noodt, 1964 (syn.Robertgurneya arabica (Noodt, 1964)
  • Amphiascoides atopus Lotufo & Fleeger, 1995
  • Amphiascoides breviarticulatus Kunz, 1983 [Azores; marine
  • Amphiascoides brevifurca (Czerniavski, 1868) [Europe; marine]
  • Amphiascoides bulbiseta pallares, 1975 [Argentina; marine]
  • Amphiascoides commensalis (Seiwell, 1928) accepted as Paramphiascella commensalis (Seiwell, 1928)
  • Amphiascoides coreanus Lee, Soh & Suh 2007
  • Amphiascoides ctenophorus (Monard, 1928) (syn. Robertgurneya rostrata (Gurney, 1927)
  • Amphiascoides debilis (Giesbrecht, 1881) [Europe, north America, north Africa?; marine]
  • Amphiascoides dictydiophorus (Monard, 1924) (syn. Robertgurneya dictydiophora (Monard, 1924)
  • Amphiascoides dimorphus Lang, 1965 [USA; marine]
  • Amphiascoides dispar (T. & A. Scott, 1894) [north Europe; marine]
  • Amphiascoides ecaudatus (Monard, 1936) (syn. Robertgurneya ecaudata (Monard, 1936)
  • Amphiascoides golikovi Chislenko, 1977 [Franz Josef Land; marine]
  • Amphiascoides hyperboreus (Scott T., 1903) (syn. Paramphiascella hyperborea (Scott T., 1903)
  • Amphiascoides hispidus (Brady, 1880) (syn. Paramphiascella hispida (Brady, 1880)
  • Amphiascoides intermixtus (Willey, 1935)
  • Amphiascoides ilievecensis (Monard, 1935) (syn. Robertgurneya ilievecensis (Monard, 1935)
  • Amphiascoides invaginatus (Monard, 1926) (syn. Paramphiascella invaginata (Monard, 1926)
  • Amphiascoides koltuni Chislenko, 1977 [Franz Josef Land; marine]
  • Amphiascoides langi (Monard, 1936) (syn. Paramphiascella langi (Monard, 1936)
  • Amphiascoides lancisetiger Lang, 1965 [USA; marine]
  • Amphiascoides limicola (Brady, 1900) [north Europe; estuaries, muddy sediments] (syn Amphiascoides debilis (Giesbrecht, 1881)
  • Amphiascoides limicolus (Brady, 1899) (syn. Amphiascoides limicola (Brady, 1899), Amphiascoides debilis (Giesbrecht, 1881)
  • Amphiascoides linearis (Sars G.O., 1907) (syn. Amphiascoides neglectus (Norman & Scott T., 1905)
  • Amphiascoides littoralis (T. Scott, 1903) [Scotland; marine]
  • Amphiascoides nanoides (Sars, 1911) [Norway; marine sand]
  • Amphiascoides nanus (Sars, 1906) [Europe, north Africa; marine, fine sand]
  • Amphiascoides neglectus (Norman & T. Scott, 1905 [europe, Italy; marine, lagoons]
  • Amphiascoides nichollsi Lang, 1965 [USA; marine]
  • Amphiascoides paradebilis Chislenko, 1978 [Japan; marine]
  • Amphiascoides petkovskii Lang, 1965 [USA; marine]
  • Amphiascoides proximus (T. Scott, 1914) [Falklands; marine]
  • Amphiascoides pygmaea (Norman & Scott T., 1905)
  • Amphiascoides roberti (Monard, 1935) (syn. Paramphiascella roberti (Monard, 1935)
  • Amphiascoides robinsonii (Scott A., 1902) (syn. Paramphiascella robinsonii (Scott A., 1902)
  • Amphiascoides rostratus (Gurney, 1927) (syn. Robertgurneya rostrata (Gurney, 1927)
  • Amphiascoides speciosus (Brian, 1921) (syn. Robertgurneya speciosus (Brian, 1921), Amphiascella brevifurca (Czerniavsky, 1868), Amphiascoides brevifurca (Czerniavsky, 1868)
  • Amphiascoides spinulosus (Sars G.O., 1911) (syn. Robertgurneya spinulosa (Sars G.O., 1911), Robertgurneyella spinulosa (Sars G.O., 1911)
  • Amphiascoides sterilis (Monard, 1926)
  • Amphiascoides subdebilis (Willey, 1935) [Europe, Algerie, Bermuda, Australia; marine, wetlands]
  • Amphiascoides vararensis (Scott T., 1903) (syn. Paramphiascella vararensis (Scott T., 1903)
  • Amphiascoides walteri Suarez-Morales & Avilés-Torres, 2003 [Mexico; marine]



    World distribution of the known species of Amphiascoides based on Nicholls (1941), Noodt (1955a,b), Lang (1948, 1965), Por & Marcus (1972), Mielke (1974), Marcotte & Coull (1975), Pallares (1975a,b), Hicks (1977), Chislenko (1977, 1978), Ceccherelli & Rossin (1979), Ceccherelli & Ferrari (1982), Kask et al., (1983), Kunz (1983), Bodin (1997), and Lotufo & Fleeger (1995). Key for the abbreviations of names: Aa= Amphiascoides atopus; Abr= A. breviarticulatus; Abf= A. brevifurca; Abu= A. bulbiseta; Ade= A. debilis; Ad= A. dimorphus; Adi= A. dispar; Ag= A. golikovi; Ak= A. koltuni; Al= A. lancisetiger; Alt= A. littoralis; An= A. nanus; And= A. nanoides; Ang= A. neglectus; Ani= A. nichollsi; Apa= A. paradebilis; Ap= A. petkovski; Apx= A. proxima; As= A. sterilis; Asd= A. subdebilis; Aw= A. walteri n.sp.(after: Suarez-Morales & Aviles-Torres, 2003)


    go to: DIOSACCIDAE (www.luciopesce.net/copepods/diosacc.htm)


    LAOPHONTISOCHRA George, 2002 *

  • Laophontisochra maryamae K. H. George, 2002 [Chile; psammon]
  • Laophontisochra teruerae Björnberg, 2014 [Brazil]


    LEPTOPONTIIDAE Lang, 1948 sensu Martinez Arbizu & Moura, 1994


    Martinez Arbizu & Moura, 1994 dissolved the family Cylindropsyllidae (Sars, 1909), Lang, 1948 , allocating the subfamily Cylindropsillinae to the family Canthocamptidae and raising to family rank the subfamily Leptopontiinae.

    More recently Recently Huys & Conroy-Dalton (2006) described 3 new species of the genus Evansula, including them in the re-established family Cylindropsillidae: Evansula cumbaensis Huys & Conroy-Dalton, 2006; Evansula polaris Huys & Conroy-Dalton, 2006; Evansula spinosa Huys & Conroy-Dalton, 2006.

    In this occasion the AA. retain the Cylindropsyllidae as a valid family comprising the following genera: Cylindropsyllus Brady, 1880; Evansula T. Scott, 1906b; Stenocaris Sars, 1909; Cylinula Coull, 1971; Boreopontia Willems, 1981; Stenocaropsis Apostolov, 1982; Willemsia Huys & Conroy-Dalton, 1993; Navalonia Huys & Conroy-Dalton, 1993; and Selenopsyllus Moura & Pottek, 1998. Of these, the genus Cylindropsyllus is possibly paraphyletic and Stenocaris undoubtedly polyphyletic.


    LEPTOPONTIA T. Scott, 1902

    "T. Scott (1902) claimed that Leptopontia T. Scott resembled Mesochra Boeck (now Canthocamptidae) and Tetragoniceps Brady (now Tetragonicipitidae) in certain aspects, a view also held by Monard (1927) who formally allocated it to the Canthocamptidae. The genus was not considered by Sars (1903–1911) but Gurney (1932) placed it, along with Evansula T. Scott and Leptastacus T. Scott, in the Evansula series, one of six generic groupings defined in Gurney’s classification of the Canthocamptidae.

    The two monotypic genera Leptopontia and Arenopontia Kunz were moved by Lang (1944) to the Cylindropsyllidae, i.e. in the Leptopontia-Reihe which subsequently acquired subfamilial status (Lang, 1948). This classification gained wide usage although some authors continued to include Leptopontia in the Canthocamptidae (Apostolov, 1971). Recently, Martinez Arbizu & Moura (1994) dramatically altered the concept of the Leptopontiinae and raised it to family rank.

    The genus Leptopontia, however, has remained monotypic since its establishment at the turn of the century (T. Scott, 1902). Leptopontia curvicauda T. Scott was one of the very few interstitial copepods discovered during the era marked by the numerous surveys of Thomas and Andrew Scott, long before Wilson (1935) and Nicholls (1935) drew attention on the existence of meiofaunal harpacticoids in sandy beaches on both sides of the Atlantic." (from: Huys & Conroy-Dalton, 1996).


  • Leptopontia americana Huys & Conroy-Dalton, 1996 [USA; subtidal sand]
  • Leptopontia breviarticulata Huys & Conroy-Dalton, 1996 [Galapagos; marine sand]
  • Leptopontia curvicauda T. Scott, 1902 [Europe; marine sand]
  • Leptopontia curvicauda sensu Apostolov Huys & Conroy-Dalton, 1996
  • Leptopontia dovpori Huys & Conroy-Dalton, 1996 [England; coarse gritty sand]
  • Leptopontia ferrolensis Candas et al., 2013 [Spain]
  • Leptopontia flandrica Huys & Conroy-Dalton, 1996 [North Sea; marine sand]
  • Leptopontia mediterranea Huys & Conroy-Dalton, 1996 [Corsica; marine]
  • Leptopontia punctata Huys & Conroy-Dalton, 1996 [Sweden; marine sand]



    PSAMMOPSYLLUS Nicholls, 1945 (syn. Paulianicaris Chappuis, 1952 )


    "The genus Psammopsyllus was first created by Nicholls (1945), who allocated it to the Stenocaridae Monard, 1927. Later, Krishnaswamy (1956) established a new genus, Sewellina, and placed it together with the closely related Psammopsyllus in a new subfamily Psammopsyllinae, within the Cylindropsyllidae.

    Martínez Arbizu and Moura (1994) upgraded the Leptopontiinae to family status to accommodate the subfamilies Leptopontiinae, Psammopsyllinae, and a new subfamily, Arenopontiinae. Huys et al. (1996) only partly accepted this classification and treated the Arenopontiinae as a distinct family.

    Bruno et al. (1998) also showed that Arenopontiinae are not closely related to the Leptopontiinae and recognized a relationship between the Psammopsyllinae and Parastenocarididae.

    The genus Psammopsyllus is characterized by the loss of the exopod on the first leg. (from: Karaytug & Sak, 2005).

    Wells (2007) included this genus, as well as the following Sewellina, Ichnusella,Parasewellina and Prosewellina in the family Psammopsyllidae Krishnaswamy, 1956.


  • Psammopsyllus arenarius Enckell, 1965 [Spain; marine interstitial]
  • Psammopsyllus arganoi Cottarelli, Puccetti & Saporito, 1984 [Mexico; brackish interstitial waters]
  • Psammopsyllus brevipes Cottarelli, Puccetti & Saporito, 1984 [Celebes; estuarine waters]
  • Psammopsyllus cornifer (Chappuis, 1952) [Madagascar; marine sand]
  • Psammopsyllus delamarei (Chappuis, 1954) [north Africa; psammon]
  • Psammopsyllus ertunci Karaytug & Sak, 2005 [Black Sea]
  • Psammopsyllus falciseta Mielke, 1983 [Panama; marine coastal interstitial]
  • Psammopsyllus imamurai Kikuchi, 1972 [Japan; lake interstitial]
  • Psammopsyllus limnicola Chappuis, 1954 [Madagascar; fresh water interstitial]
  • Psammopsyllus longipes Cottarelli, Puccetti & Saporito, 1984 [Lombok island; marine interstitial]
  • Psammopsyllus maricae Cottarelli, Saporito & Puccetti, 1983 [Italy, Scotland; estuarine waters]
  • Psammopsyllus operculatus Nicholls, 1945 [Africa, Australia, Sri Lanka, Brazil; marine sandy beaches]
  • Psammopsyllus pasquinii Cottarelli, 1969 [Italy; marine intersttial]
  • Psammopsyllus pori Kunz, 1993 [Namibia; marine sand]
  • Psammopsyllus stri Mielke, 1983 [Hawai, Panama; coastal interstitial]
  • Psammopsyllus tridentatus Soyer, 1975 [Kerguelen; mesopsammon]


    SEWELLINA Krishnaswamy, 1956

  • Sewellina reducta Krishnaswamy, 1956 [India; interstitial]
  • Sewellina subtilis Cottarelli, Saporito & Puccetti, 1986 [Sri Lanka; interstitial]


    PARASEWELLINA Cottarelli, Saporito & Puccetti, 1986

  • Parasewellina prima Cottarelli, Saporito & Puccetti, 1986 [Sri Lanka; interstitial]


    PROSEWELLINA Mielke, 1987

  • Prosewellina chilensis Mielke, 1987 [Chile; interstitial]


    BERERAIA Huys, 2009 nom. nov. for ICHNUSELLA Cottarelli, 1971

    Recently Hus (2009) pointed out that Cottarelli (1971) established the genus Ichnusella for his new species I. eione Cottarelli, 1971 (type species by original designation) and Psammopsyllus pasquinii Cottarelli, 1969, overlooking that the generic name Ichnusella had already been introduced by Dieni and Massari (1966) for a genus of fossil Foraminiferida.

  • Bereraia eione (Cottarelli, 1971) [Italy, Sardinia; hyporheic]
  • Bereraia ionica (Berera & Cottarelli, 2003) [Italy; freshwater interstitial]
  • Bereraia improvisa (Berera & Cottarelli, 2001) [Italy; interstitial fresh waters]
  • Bereraia longifurca (Cottarelli, Bruno & Berera, 1999) [Italy; fresh ground waters]
  • Bereraia pasquinii (Cottarelli, 1969)
  • Bereraia sibaritica (Berera & Cottarelli, 2003)
  • Bereraia tertia (Cottarelli, Bruno & Berera, 1999) [Italy; fresh ground waters]


    NOTOPONTIA Bodiou, 1977

  • Notopontia biarticulata (Wells, 1967) [Africa; marine interstitial]
  • Notopontia stephanieae Bodiou, 1977 [Kerguelen; marine interstitial]


    SYRTICOLA Willems & Claeys, 1982

  • Syrticola flandricus Willems & Claeys, 1982 [North Sea]
  • Syrticola galapagoensis (Mielke, 1982) [Galapagos; interstitial]
  • Syrticola intermedius Huys & Ohtsuka, 1993 [Japan; interstitial]
  • Syrticola mediterraneus Willems, Claeys & Fiers, 1987 [France, Corsica; marine interstitial]
  • Syrticola trispinosus Scott A., 1896


    LAOPHONTISOCHRA George, 2002 *

  • Laophontisochra maryamae K. H. George, 2002 [Chile; psammon]
  • Laophontisochra teruerae Björnberg, 2014 [Brazil]




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