HARPACTICOIDA G.O. Sars, 1903

Karanovic, Tomislav, Cho, Joo-Lae & Lee, Wonchoel, 2012, Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea, Journal of Natural History 46 (25 - 26), pp. 1573-1613 : 1577-1579

Order HARPACTICOIDA G.O. Sars, 1903

Family PARASTENOCARIDIDAE Chappuis, 1940 Subfamily FONTINALICARIDINAE Schminke, 2010 Genus Proserpinicaris Jakobi, 1972

Synonyms

Nipponicaris Jakobi, 1972 , syn. nov.; Pannonicaris Jakobi, 1972 syn. nov.

Type species

Parastenocaris proserpina Chappuis, 1938 .

Other species

Parastenocaris admete Cottarelli, Fasano, Mura and Saporito, 1980 ; P. amalasuntae Bruno and Cottarelli, 1998 ; P. cantabrica Chappuis, 1937 ; P. cruzi Noodt and Galhano, 1969 ; P. fontinalis meridionalis Rouch, 1990 ; P. gorganensis Kovalchuk and Kovalchuk, 1990 ; P. hispanica Martínez Arbizu, 1997 ; P. ima Cottarelli, 1989 ; P. imjin sp. nov.; P. kalypso Pesce, Galassi and Cottarelli, 1988 ; P. mangini Rouch, 1992 ; P. moravica Šterba, 1965 ; P. nicolasi Rouch, 1996 ; P. nipponensis Chappuis, 1955 ; P. ondali Lee and Chang, 2009 ; P. pannonica Török, 1935 ; P. phyllura Kiefer, 1938 ; P. wangpi sp. nov.; P. young sp. nov.

Revised diagnosis

Small to medium-sized Fontinalicaridinae , with cylindrical habitus, smooth cuticule, somites ornamented with large sensilla, and with dorsal cuticular windows on cephalothorax and all urosomal somites, except first and last; spinules, if present at all, usually restricted to anal somite. Podoplean boundary between prosome and urosome inconspicuous. Genital complex in female occupying anterior ventral half of genital double somite; genital apertures and median copulatory pores covered by vestigial sixth legs, fused completely into relatively narrow flap. Caudal rami cylindrical or leaf-like, with lateral cuticular pore near posterior margin, armed with seven elements (three lateral, one dorsal, and three apical); lateral elements inserted close to each other and much more anteriorly than dorsal seta; one lateral element minute and hard to observe between two others. Male antennula eight-segmented, prehensile, with geniculation between third and fourth and sixth and seventh segments; last two segments in line; distal anterior corner of seventh segment produced into very small spiniform process, but larger proximal spiniform process present on fifth segment on anterior surface; aesthetasc on fifth segment usually massive and reaching tip of appendage. First swimming leg with no chitinous processes or inner basal armature. Second swimming leg with one-segmented endopod, armed with single slen- der seta apically, and with several apical spinules and sometimes with several lateral spinules. Third swimming leg endopod in female very small and linguiform segment unarmed, usually with few apical spinules. Third swimming legs in male transformed into strong grasping organs, with large intercoxal sclerite between them, each composed of praecoxa, coxa, basis, two-segmented exopod, and endopod reduced to single slender armature element; basis and proximal exopodal segment robust, latter usually with beaks or chitinous lobes on inner margin; outer spine on first exopodal segment smooth, large, and curved; ancestral distal exopodal segment (apophysis) small, usually cylindrical, oriented slightly inwards, unornamented, and armed with single short element on top, often leaf-like, or thumb-like. Fourth swimming leg in male with or without spinules on inner margin of coxa, but always with large and slen- der hyaline process on anterior surface of basis, between exopod and endopod, and no other chitinous structures on basis; first exopodal segment with longitudinal row of strong spinules on inner margin, and no other structures or depressions on this margin; endopod one-segmented, with or without apical armature, but usually curved and variously transformed, often knife-like (with serrated edges or tubercules), rarely pinnate, sometimes with scoop-like structure on tip. Fourth swimming leg in female without hyaline process on basis or inner spinules on first exopodal segment; endopod also one-segmented but cylindrical, straight, and not transformed, with strong element on tip, several apical spinules, and several spinules around midlength close to inner margin. Fifth legs very similar in shape in male and female, elongated and simple triangular plates, with inner distal corner produced into spiniform process, ornamented with spinules along inner margin (these often absent in female), usually another row of spinules on posterior surface proximally, and single large cuticular pore on anterior surface; armature consists of very long outermost seta (ancestral basal); two setae at outer base of inner distal process (probably ancestral endopodal setae), and sometimes small seta (or long spinule?) on posterior surface at base of basal seta; latter element most often reduced to minute spiniform process and hardly visible (smaller than most spinules). Sixth legs in male also fused (or right one reduced and left one enlarged?) smooth, unarmed, unornamented, forming simple operculum covering gonopore.

Remarks

Fourth swimming leg in male with a large and slender hyaline process on the anterior surface of basis, between exopod and endopod, being the sole chitinous structure on this segment, is a synapomorphy that unites all species included in the currently redefined genus Proserpinicaris Jakobi, 1972 . Most other characters can be found in some other Fontinalicaridinae genera, or species of Parastenocaris Kessler, 1913 that are currently considered to be members of this subfamily by Schminke (2010) (see the Introduction section above). Their affinities are further discussed in the Discussion section below. Endopod of the fourth swimming leg in male is usually lanceolate or knife-like, with serrated margins, except in Proserpinicaris ima ( Cottarelli, 1989) comb. nov., where it is completely smooth, and in P. cruzi ( Noodt and Galhano, 1969) comb. nov. and P. cantabrica ( Chappuis, 1937) , where the endopod is pinnate (or plumose) along inner margin. We believe both conditions are secondary transformations, because the transformed endopod with serrulate or tuberculate margins can be found in some other unrelated taxa, and so is probably a plesiomorphic character state. All five Asian species have the endopod of the fourth leg in male with a scoop-like structure, formed by a bunch of basally fused apical spinules. They include the type species of the genus Nipponicaris Jakobi, 1972 , Parastenocaris nipponensis Chappuis, 1955 , and unquestionably form a monophyletic group of species. They all, however, have a very well developed hyaline process on the anterior surface of the fourth leg basis, which is a complex structure and highly unlikely to have arisen convergently a number of times. That is why we synonymize the genus Nipponicaris with Proserpinicaris .

The original description of Parastenocaris pannonica Török, 1935 , which was designated by Jakobi (1972) as the type species of his new genus Pannonicaris Jakobi, 1972 , is incomplete (see Török 1935, Lang 1948). Later redescriptions of this species by Damian (1958) and Kulhavý (1960) indicate that the large hyaline process on the fourth leg basis is between an exopod and endopod, which puts this species in the genus Proserpinicaris and renders the genus Pannonicaris its objective synonym. We speculate that the fourth leg illustrated by Török (1935) and repeated in Lang (1948) is actually a mix-up, because all other morphological details agree with those observed in Proserpinicaris . Török (1935) described two sympatric species from the Budapest water supply: P. pannonica (note: he wrongly spelled the species name as pannonicus, but being an adjective it has to agree with the feminine generic name), and P. budapestiensis Török, 1935 . The most probable explanation is that he swapped the male fourth legs of these two species (or just their drawings during the preparation of plates), as P. budapestiensis also has all other typical Parastenocaridinae characters.

The genus Proserpinicaris , as defined here, is Palaearctic in distribution, with its centre of diversity in southern Europe. Note that Parastenocaris fontinalis meridionalis Rouch, 1990 was correctly considered a separate species already by Martínez Arbizu (1997), although he considered it a member of the fontinalis -group. Karanovic (2005) pointed out that this species is a member of the proserpina -group, as well as P. hispanica Martínez Arbizu, 1997 . We discuss this further below.