[syn. PTEROPSYLLIDAE Nicholls, 1944 (Lang, 1948)]

The family Tetragonicipitidae Lang, 1944, contains 15 genera of which the genus Phyllopodopsyllus T. Scott, 1906 is the most speciose, consisting of more than 60 species and subspecies, most of them inhabiting coarse sand and shell gravel of the intertidal zone where strong water turbulences prevail.

ADOGINICEPS Gomez & Morales-Serna, 2015
DIAGONICEPS Willey, 1930
LAOPHONTELLA Thompson & A. Scott, 1903
MWANIA Fiers & De Troch, 2000
NEOGONICEPS Fiers & De Troch, 2000
ONISCOPSIS Chappuis, 1954
TARENGOTICEPS Gomez & Morales-Serna, 2015

Fiers (1995) described the genus Nidiagoceps from Great Britain, later synonymyzed with Paraschizopera Wells, 1981 (Bodin, 1997).



According to Bodin (1977) the tetragonicipitid genus Phyllopodopsyllus T. Scott, 1906, with 66 valid species and subspecies, is one of the most speciose marine harpacticoid genera, together with Halectinosoma Lang, 1944 and Stenhelia Boeck, 1865, and a marine harpacticoid genus with the largest number of new described species in the last fifty years.

Lang (1944) divided the original genus Phyllopodopsyllus in two genera (Phyllopodopsyllus and Paraphyllopodopsyllus Lang, 1944) on the basis of the shape of the second segment of antennula and the setation of the second and third swimming legs in the females. Many authors followed this revision, and Vervoort (1964) provided a useful key to the species of the genus Phyllopodopsyllus, although not for Paraphyllopodopsyllus. Lang (1965) reconsidered his previous vue and synonymized Paraphyllopodopsyllus with Phyllopodopsyllus after several new species were described possessing intermediate caharacteristics. He compiled an excellent key to 21 species recognized at that time, and subdivided the genus into three morphological groups on the basis of the morphology (i.e. presence or absence, and dimensions of an unguiform projection) of the second antennular segment.

Later on Coull (1973) presented a very detailed subjective survey of the genus Phyllopodopsyllus, and compiled a most instructive table listing the most salient morphological characteristics. His key, untill today the most recent one, comprised 33 species and subspecies.

The last critical review of the genus was provided by Kunz (1984), who pointed out relationships betwen the genera and species within the family Tetragonicipitidae, representing their phylogenetic affinities, as well as subdivided the genus Phyllopodopsyllus into nine different species-groups, viz. bradyi, furciger, borutzkyi, pauli, opistoceratus, aegypticus, mossmani, xenus and longipalpatus, according to the shape and oligomerization and setation of antenna, antennula and swimming legs 2 to 4. More recently, Bodin (1988) and Mielke (1989) followed the above subdivision; Fiers (1995) criticized the naturalness of the above groups definitions, even if he suggested that the group definition of the genus could be a most practical identification tool.

From an ecological point of view, the genus Phyllopodopsyllus is widespread in marine coarse sand and shell gravel of the intertidal zone, with a single citation (Phyllopodopsyllus sp.) for strictly interstitial fresh waters of Italy (Cottarelli et al., 1996).

  • Phyllopodopsyllus aegypticus Nicholls, 1944 [Africa, Egypt]
    [syn. Paraphyllopodopsyllus aegypticus (Nicholls)]
  • Phyllopodopsyllus alatus Fiers, 1986 [New Guinea]
  • Phyllopodopsyllus ancylus Mielke, 1992 [Costa Rica]
  • Phyllopodopsyllus angolensis Kunz, 1984 [Africa, Angola]
  • Phyllopodopsyllus bahamensis Geddes, 1968 [Bahamas]
  • Phyllopodopsyllus bermudae Lang, 1948 [Bermuda]
  • Phyllopodopsyllus berrieri Monard, 1936 [Italy, Africa ]
    [syn. Paraphyllopodopsyllus berrieri (Monard)]
  • Phyllopodopsyllus biarticulatus (Wells, 1967) [ Africa ? ]
  • Phyllopodopsyllus borutzkyi Lang, 1965 [ USA, Californian coast ]
  • Phyllopodopsyllus bradyi (T. Scott, 1892) [ North Europe; marine sand ]
  • Phyllopodopsyllus briani briani Petkovski, 1955 [ Yugoslavia ]
  • Phyllopodopsyllus briani petkovskii Kunz, 1984 comb.nov. (Karanovic et al., in press) [Hawai]
  • Phyllopodopsyllus carinatus Mielke, 1992 [Costa Rica]
  • Phyllopodopsyllus chavei Coull, 1970 [ Bermuda ]
  • Phyllopodopsyllus crenulatus Wells & Rao, 1987 [Andaman] ****
  • Phyllopodopsyllus curtus Marcus, 1976 [ Mediterranean ]
  • Phyllopodopsyllus danielae Bodin, 1964 [France]
  • Phyllopodopsyllus furciger Sars, 1907 [Africa, Tanzania; Galapagos, Bermuda]
  • Phyllopodopsyllus galapagoensis Mielke, 1989 [Galapagos]
  • Phyllopodopsyllus geddesi Kunz, 1984 [Hawai]
  • Phyllopodopsyllus gertrudi Kunz, 1984 [Pacific Sea] *****
  • Phyllopodopsyllus gertrudi costaricensis Mielke, 1992 [Costa Rica] *****
  • Phyllopodopsyllus gracilipes Wells & Rao, 1987 [Andaman and Nicobar]
  • Phyllopodopsyllus hardingi (Roe, 1955) [Europe; marine]
    [syn. Paraphyllopodopsyllus hardingi Roe]
  • Phyllopodopsyllus hartmannorum Kunz, 1995 [Africa, Tanzania] [PDF]
  • Phyllopodopsyllus hermani Coull, 1969 [Bermuda]
  • Phyllopodopsyllus hibernicus (Roe, 1955) [ Europe; marine ]
    [syn. Paraphyllopodopsyllus hibernicus Roe]
  • Phyllopodopsyllus iuanamai Biornberg & Kihara, 2013 [Brazil] [PDF]
  • Phyllopodopsyllus kunzi Mielke, 1989 [Galapagos]
  • Phyllopodopsyllus langi Kunz, 1975 [south Africa]
  • Phyllopodopsyllus laspalmensis Marinov, 1973 [Atlantic Ocean ]
  • Phyllopodopsyllus laticauda Por, 1964 [Israel]
  • Phyllopodopsyllus longicaudatus A. Scott, 1909 [ Africa ]
    [syn. Paraphyllopodopsyllus longipalpatus Chappuis]
  • Phyllopodopsyllus longipalpatus longipalpatus (Chappuis, 1954) [Italy]
  • Phyllopodopsyllus longipalpatus madagascarensis Kunz, 1984 [Madagascar]
  • Phyllopodopsyllus longipalpatus hawaiiensis Kunz, 1984 [Hawai]
  • Phyllopodopsyllus medius Por, 1964 [Israel]
  • Phyllopodopsyllus mielkei mielkei Kunz, 1984 [Hawai] ****
  • Phyllopodopsyllus mielkei californicus Kunz, 1984 [San Francisco Bay] ****
  • Phyllopodopsyllus minor (Th. & A. Scott, 1903) [New Zealand] **
  • Phyllopodopsyllus minutus Lang, 1948 [Bermuda]
  • Phyllopodopsyllus mossmani T. Scott, 1912 [Argentine, Falkland Islands]
  • Phyllopodopsyllus mossmani chiloensis Mielke, 1992 [Chile]
    [syn. P. mossmani (T. Scott); P. trichophorus Kunz, 1951]
  • Phyllopodopsyllus opisthoceratus Geddes, 1968 [Bahamas]
  • Phyllopodopsyllus pallaresae Kunz, 1995 [Africa, Namibia] [PDF]
  • Phyllopodopsyllus paraborutzkyi Kunz, 1975 [South Africa]
  • Phyllopodopsyllus parabradyi Lang, 1965 [USA, Californian coast]
  • Phyllopodopsyllus parafurciger parafurciger Geddes, 1968 [Bahamas]
  • Phyllopodopsyllus parafurciger carolinensis Coull, 1971 [USA, North Carolina]
  • Phyllopodopsyllus paramossmani (Lang, 1934) [Stewart Island]
    [syn. Paraphyllopodopsyllus paramossmani (Lang)]
  • Phyllopodopsyllus parastigmosus Gomez & Morales-Serna, 2015
  • Phyllopodopsyllus paraxenus Coull, 1970 [Bermuda]
  • Phyllopodopsyllus pauli Crisafi, 1960 [Africa, Tunisia; Italy; Bulgaria; France]
    [syn. Phyllopodopsyllus ponticus Apostolov] [PDF]
  • Phyllopodopsyllus pirgos Apostolov, 1969 [Bulgaria] [PDF] ***
  • Phyllopodopsyllus ponticus Apostolov, 1968 [Bulgaria] [PDF]
    [syn. of P. pauli, according to Apostolov (1973)]
  • Phyllopodopsyllus pseudokunzi Biornberg & Kihara, 2013 [Brazil] [PDF]
  • Phyllopodopsyllus punctatus Kitazima, 1981 [Japan]
  • Phyllopodopsyllus setouchiensis Kitazima, 1981 [Japan; Costa Rica] [PDF]
  • Phyllopodopsyllus simplex Kitazima, 1981 [ Japan]
  • Phyllopodopsyllus sp. [New Zealand] (Wells, in litt.)
  • Phyllopodopsyllus sp. [Italy, Sardinia] (Cottarelli et al., 1996)
  • Phyllopodopsyllus stigmosus Wells & Rao, 1987 [Andaman and Nicobar]
  • Phyllopodopsyllus tenuis Wells & Rao, 1985 [ Andaman and Nicobar islands]
  • Phyllopodopsyllus thiebaudi thiebaudi Petkovski, 1955 [Yugoslavia]
    [syn. P. pirgos Apostolov: P. intermedius Noodt, 1955] *
  • Phyllopodopsyllus thiebaudi santacruzensis Mielke, 1989 [Galapagos]
  • Phyllopodopsyllus tristanensis (Wiborg, 1964) [ Tristan ] **
  • Phyllopodopsyllus wellsi Karanovic et al., 2001 [ W-Australia; anchialine] [PDF]
  • Phyllopodopsyllus xenus (Kunz, 1951) [Africa, Namibia]
  • Phyllopodopsyllus yucatanensis Fiers, 1995 [Yucatan]
    [synonym of P. parafurciger (Karanovic et al., in press)]

    * Fiers (1986) considers P. intermedius as a distinct species.

    ** incertae sedis

    *** synonym of P. briani (Coull, 1973; Apostolov & Marinov, 1988)
    juvenile male of P. thiebaudi, according to Kunz (1984)

    **** synonym of Phyllopodopsyllus setouchiensis (Mielke, 1992)

    ***** synonim of Phyllopodopsyllus aegypticus (Karanovic et al., in press)



    BJÖRNBERGT. & KIHARA T. 2013. On Tetragonicipitidae (Crustacea, Copepoda) from the Channel of São Sebastião, Brazil, with description of their nauplii and two new species of Phyllopodopsyllus. Zootaxa 3718 (6): 501–529

    BODIN P. 1997. Catalogue of the new marine Harpacticoid Copepods (1997 Edition). Documents de travail de l'Institut des Sciences naturelles de Belgigue, pp.304.

    COULL B.C. 1973. Harpacticoid copepods (Crustacea) of the family Tetragonicipitidae Lang: a review and revision, with keys to the genera and species. Proc. Biol. soc. Washington, 86 (2): 9-24.

    FIERS F. 1986. New and interesting copepods (Crustacea, Copepoda) from brackish waters of Laing Island (Northern Papua New Guinea) Leopold III Biological station, Laing Island - Contribution n° 96. Bull.Inst. Roy. sc.nat. Belg., 56.99-120. >

    FIERS F. 1995. New Tetragonicipitidae (Copepoda, Harpacticoida) from the Yucatean continental shelf (Mexico) including a revision of the genus Diagoniceps Willey. Bull. Inst. r. Sc. nat. belg., Biol., 65: 151-236.

    FIERS F. & M. DE TROCH. 2000. New tetragonicipitidae (Copepoda, Harpacticoida) from the Indo-Pacific. Hydrobiologia, 434: 97-144.

    KARANOVIC T., G.L. PESCE & W. F. HUMPHREYS. 2001 . Copepods from ground waters of Western Australia, V. Discovery of the genus Phyllopodopsyllus T. Scott, 1906 in Australia, and description of Phyllopodopsyllus wellsi n.sp. (Crustacea: Copepoda: Harpacticoida). Rec. Western Australian Mus., 20: 333-344 Mus.

    KUNZ H. 1984. Systematik der familie Tetragonicipitidae Lang (Crustacea, Harpacticoida). Mitt. Zool. Mus. Univ. Kiel, 2:33-48.

    KUNZ H. 1995. Contribution to the knowledge of the genus Phyllopodopsyllus T. Scott (Copepoda, Harpacticoida) from Africa with description of two new species. Hydrobiologia, 297: 83-98.

    MIELKE W. 1992. Six representatives of the Tetragonicipitidae (Copepoda) from Costa Rica. Microfauna marina, 7: 101-146.

    WELLS J.B. & G.C. RAO. 1987. Littoral Harpacticoida (Crustacea: Copepoda) from Andaman and Nicobar Islands. Mem. Zool. Survey India 16: 1-385


    COULL B.C.
    University of South Carolina
    Columbia SC 29208 USA
    e-mail: bccoull@sc.edu

    FIERS F.
    Royal Belgian Institute of Natural Sciences
    Vautierstraat 29, B-1000 Brussels - Belgium
    E-mail: frankfiers@kbinirsnb.be

    KARANOVIC T. University of Tasmania - School of Zoology
    Private Bag 5, Hobart - Tasmania 7001
    e-mail: tomislav.karanovic@utas.edu.au

    II Zoologisches Institut und Museum der Universitat Gottingen
    Berliner Strabe 28, D-37073 Gottingen - Germany

    PESCE G.L.
    Via Santa Maria al Colle 131/B - 67019 L'Aquila (Italy)
    tel. 39-3338769242
    e-mail: luciopesce@virgilio.it

    WELLS J.B.J.
    Victoria University of Wellington - Wellington - New Zealand
    e-mail: John.Wells@vuw.ac.nz

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