Contributions to Zoology, 66 (4) 215-226 (1997)
SPB Academic Publishing bv, Amsterdam
Parastenocaris hispanica n. sp. (Copepoda: Harpacticoida:
Parastenocarididae) from hyporheic groundwaters in Spain and its
within the of phylogenetic position fontinalis- group species
Pedro Martínez Arbizu
Arbeitsgruppe Zoomorphologie, Fachbereich Biologie, Carl von Ossietzky Universität Oldenburg,
D-26111 Oldenburg, Germany
Keywords: Harpacticoida, Parastenocaris hispanica, new species, fontinalis-group, Fontinalicaris,
taxonomy, phylogeny, zoogeography
Abstract (Lang, 1948) and is described here in the
first of series of the a contributions to systematic s
Parastenocaris is described from hispanica n. sp. hyporheic of Iberian Parastenocarididae.
groundwaters in Spain. The phylogenetic position of the new The phylogenetic relationships within the Pa- within the is discussed. As a species fontinalis-group Lang still uncertain. rastenocarididae are The genus result P. fontinalis meridionalis Rouch is elevated to species Parastenocaris is without doubt rank. The fontinalis-group is characterized by six autapo- polyphyletic
morphic characters. (Schminke, 1993). Discussions about affinities of
the known species are based on the assignment of
these species to different "phyletic" groups (Lang, Resumen 1948). In order to elucidate the phylogenetic posi-
fon-tion of the newly discovered species, the Se describe Parastenocaris una nuevaespécie, hispanica sp. n., tinalis-groupis characterized and phylogenetically de subterráneas de se discute su aguas España y posición the methods of phylogenetic dentro del Como analysed applying filogenética grupo-fontinalis Lang. resultado,
as defined meridionalis Rouch debe ser elevado a systematics by Hennig (1982). P. fontinalis fontinalis-categoría de El está caracterizado seis espécie. grupo-fonlinalis por autapomorfías.
Methods
Introduction
Groundwater samples were taken by the Karaman-Chappuis
method (Chappuis, 1942) in Spanish rivers, filtered through a little is known about the Parastenocarididae Very of mesh and 5% nylon net 100 um preserved in formaldehyde. the of the Iberian Penin- inhabiting groundwaters Additional samples were kindly made available by Dr. Rouch
These taken in sula. Three papers have been published previ- (Saint-Girons, France). samples were Spanish
rivers Dr. R. Dr. F. Lescher-Moutoué and Dr. N. ously: Chappuis (1937) described two new species by Rouch,
Gourbault in the 1976 and 1978. The specimens were in Enckell years from caves Santander (northern Spain), to transferred a medium consisting of a mixture of lactic acid (1965) found two new species in the river Fuen- and glycerine gelatine and dissected for slide preparation. The southern and Noodt & girola (Andalucía, Spain) Leitz Dialux figures have been prepared with a phase contrast Galhano the fauna of (1969) studied groundwater microscope using a camera lucida. northern and described five Paraste- Portugal new nocaris species. During an intensive study of the freshwater harpacticoid fauna of the Iberian groupAbbreviations used in the text and figures are: AÍ =antennule, Peninsula, additional Parastenocarididae eight A2 = = Ae = Md = antenna, ap apomorphy, aesthetasc,
= four of science. One = = have been found, them new to mandible, Mxl maxillule, Mx2 maxilla, Mxp of these new species belongs to the maxillipede, P1-P6 = legs 1 to 6, pi =plesiomorphy. - Parastenocaris 216 P. Martinez Arbizu hispanica n. sp. from hyporheic groundwaters in Spain
1. of the Iberian Peninsula with stations where P. has been found indicated asterisk. Fig. Map hispanica n. sp. by an
The terms "pars incisiva", "pars molaris", and "lacina Sta. 91-7/4: River Magro, between Macastre and Casas de mobilis" are omitted from the description of the mandible Millas, Prov. Valencia, Spain; 4 m from shore, water temp.
> (Mielke, 1984); the terms "furca" and "telson" are used 22°C, 50 individuals, 10.7.'91. according to Schminke (1976a); terms of the phylogenetic Sta. 91-8/8: River Cabriel at Las Hondonadas, between El analysis are used sensu Hennig (1982). The cladogram was Cañigral and Salvacañete, Prov. Cuenca, Spain (type locality), made by hand using the so-called "Hennigian principle" (cf. spring at shore, water temp. 16°C, 15 o" or, 26 9 9 and 2
Meier, 1992). copepodids, 24.8.'91.
Sta. 92-6/2: River Argelita at Fuente de los Ignacios,
Argelita, Prov. Castellón,Spain; 1 m from shore; > 20 individu-
als, 15.6.'92. Descriptive part Sta. Rouch 1976, n° 18: Well between Velamazán and
Barca, road Soria-Cl 16, Prov. Soria, Spain; 1 o• and 1 9, leg. Family Parastenocarididae Chappuis, 1933 R. Rouch, 13.9.'76.
Sta. Rouch 1976, n° 51: River Guadalope at Mas de las Parastenocaris hispanica n. sp. Matas, Prov. Teruel, Spain; 2 Cf ce and 1 Q, leg. R. Rouch,
21.9.'76.
Material. - Sta. 91-3/16: River Maimona, near bridge at road Sta. Rouch 1978, n° 9: River Guarga, road CI36 at CS-V-2002, Mas del Molino at Villanueva de Viver Prov. (CS), Sabiñánigo, Pito de Momepos, Prov. Huesca, Spain; 2 er O" and
Teruel, Spain; 4 m from shore, water temp. 11°C, 8 erer and 11 3 9 9, leg. R. Rouch, 14.09.'78. 9 9, 28.3.'91. - 217 Contributions to Zoology, 66 (4) 1997
2. Parastenocaris male habitus: dorsal lateral view. Scale bar 100 Fig. hispanica n. sp., A, view; B, µm. 218
male female of left male Fig. 3. Parastenocaris hispanica n. sp.: A, antennule; B, antennule; C, proximal segments antennule; D,
Scale bar 10 antenna. µm.
Sta. Rouch 1978, n° 15: River Alcanadre, road M240 be- the localities where Parastenocaris hispanica n. tween Angiies and Ponzano, Prov. Huesca, Spain; 8 O" O" and 8 sp. has been found. The type locality is Sta. 91-8/8, 9 9, leg- R. Rouch, 14.9.'78. river Cabriel at Salvacañete, Prov. Cuenca, Spain. Sta. Rouch 1978, n° 8: River Alcanadre at Bierge, Prov.
1 2 and 2 R. Huesca, Spain; cr, 9 9 copepodids, leg. Rouch,
13.9.'78. Description of holotype and allotype. - Length 380
Sta. Rouch 1978 n° 8bis: River Alcanadre at Prov. , Bierge, with |im (from rostrum to operculum). Rostrum Huesca, Spain; 1 er, leg. R. Rouch, 13.9.'78. large base and two sensillae on tip. Céphalothorax The material is stored at the Copepod collection of the and abdominal segments 1-4 with dorsal "win- Arbeitsgruppe Zoomorphologie, University of Oldenburg, dows" For sensillae on see Germany. The type material examined for the description (Fig. 2A). tergites Fig. below (from Sta. 91-8/8) consists of 1 dissected o* (holotype, 2A-B. Operculum convex (Fig. 6C). Furca (Fig.
Coll. No. 1996.22/1-1996.22/4) and 1 dissected 9 (allotype, 7A) with 7 setae, all 3 anterolateral setae located in Coll. No. 1996.23/1-1996.23/4),each mounted on4 slides. first third oframus.
Distribution. - Northeastern Spain, from the Pyr- Male: AI (Fig. 3A, C) 8-segmented, prehensile, enees to Valencia, in the river sytems of Ebro, number of setae starting at proximal segment: 0/6/ Mijares and Júcar. Also at the river Duero on the 4/2/4+Ae/O/O/9+Ae. A2 with allobasis and 1-seg- Iberian Plateau (Meseta). This river flows into the mented exopod carrying 1 pinnate seta (Fig. 3D). Atlantic in north that the Portugal, so species may Distal with them endopod segment 7 setae (2 of have distribution. shows a wider westerly Fig. 1 Contributions to Zoology, 66 (4) - 1997 219
4. Parastenocaris male male female male male Fig. hispanican. sp.: A, pereiopod 1; B, pereiopod 2; C, endopod2; D, pereiopod3; E,
3 1978 male 3 female 3. Scale bar 10 pereiopod from sta. Rouch n° 8; F, pereiopod copepodid V; G, pereiopod µm.
geniculate) and hyaline frill. Md with 1-segmented seta and 6 spinules, outer seta of basis absent. P3
with both palp (Fig. 6D) carrying 2 setae. Mxl (Fig. 7D), (Fig. 4D) exopod segments fused, por-
precoxal arthrite with 5 setae (1 dorsal surface tion of proximal segment ending in a large seta
seta, 3 claw-like pinnate setae and 1 slender seta), ("thumb") reaching beyond end of whole exopod,
coxa with 1 seta, basis with 3 setae. Mx2 (Fig. 7C) inner margin with protuberances and tube pore
with 2 endite with inner of syncoxa endites, proximal 1, reaching beyond margin segment; portion
distal one with 3 setae, basis with strong apical of distal segment ("apophysis") terminally with
pinnate claw, endopod represented by 2 setae, strong spine (seta), on outer margin with protuber-
distal like bird exopod absent. Mxp 3-segmented (Fig. 6E), ance giving "apophysis" look a finger;
segment with 1 seta. endopod represented by 1 seta. P4 (Fig. 5A) with
PI (Fig. 4A) with 3-segmented exopod, segment 3-segmented exopod, segment 1 subterminally
2 without setae, segment 3 with 4 setae, endopod with row of 8 large and slender spinules on inner
of with inner 2 3 with 2 2 segments, segment 1 1 strong spine margin, segment without setae, segment
and inner 1 in middle of segment, segment 2 with 2 setae and apical setae hyaline frill, endopod -seg-
hyaline frill. P2 (Fig. 4B) with 3-segmented mented with terminal seta fused to it and 3
with 3-4 subordinate exopod, segments 1 and 3 with hyaline frill, seg- spinules, seta subterminally ment 1 without seta, segment 2 without setae, seg- spinules fused to its outer margin, basis with ment 3 with 3 setae, endopod 1-segmented with 1 hyaline spinule between exopod and endopod, 220 P. Martinez Arbizu - Parastenocaris hispanica n. sp. from hyporheic groundwaters in Spain
5. Parastenocarís male P4 and male 4 female Fig. hispanica n. sp.: A, opposite endopod; B, pereiopod copepodidV; C, pereiopod 4;
male 5 ventral male 5 lateral view. Scale bar 10 D, pereiopod view; E, pereiopod µm.
coxa with 6 slender spinules on inner margin. P5 without subordinate spinules fused to it, and 4
(Fig. 5 D-E) with convex outer margin bearing 4 spinules, basis without hyaline spinule between setae and inner thorn, and with row of spinules exopod and endopod, coxa without spinules on in-
3 along inner margin. ner margin. P5 (Fig. 6A) with only spinules at
inner margin. Genital field as illustrated in Fig.
6A, seminal receptacles dotted. Female: Sexual dimorphism in AI, PI, P2, P3, P4,
P5, and genital field. AI (Fig. 3B) 7-segmented, Derivado nominis. - The specific name hispanica not prehensile, number of setae starting at proxi- refers to Spain. mal segment: 0/4/5/2+Ae/l/l/9+Ae.
PI endopod segment 1 with 2 small spinules at Variability. - Material from Sta. Rouch 1978 n° 8 inner margin instead of 1 thick spinule. P2 and Sta. Rouch 1978 n° 8bis differs from the endopod (Fig. 4C) with 1 seta and only 4 spinules. holotype and allotype in the following characters:
P3 (Fig. 4G) exopod 2-segmented, segment 1 with Male: P3 exp (Fig. 4E) "apophysis" with outer 2 with 2 and inner tube orientated seta, segment apical setae stronger outer protuberance, pore frill, proximally with tube pore at inner margin, upwards. Telson (Fig. 7B) ventrally with 2 strong endopod 1-segmented with 4 spinules, without spinules at base of each furcal ramus. setae. P4 (Fig. 5C) exopod 1 apically with Female: Furca (Fig. 6B) modified as "bulbifera spinules, endopod 1-segmented, with terminal seta type" (Noodt, 1963; Schminke, 1991). Contributions to Zoology, 66 (4) -1997 221
female 5 and female furca from Sta. Rouch 1978 n° male Fig. 6. Parastenocaris hispanica n. sp.: A, pereiopod genital field; B, 8; C,
telson dorsal Scale bar 10 view; D, mandible; E, maxillipede. µm.
P. aedes 1938 Affinities of Parastenocaris hispanica sp. n. similis Török, 1935, and P. Hertzog,
are not closely related to the nominal species P.
These three to what are with fontinalis. species belong The greatest similarities those species of Schminke (1993) calls "Parastenocaridinae" in the genus belonging to the fontinalis- group (Lang,
which all three anterolateral setae of the furca in- 1948). Unfortunately, both of Lang's discriminat- P. "endo- sert on its distal third. On the other hand, ing characters for the fontinalis- group, i.e., to what Schminke (1993) calls pod P4 male as female" and "endopod P2 1-seg- fontinalis belongs
The P4 "Fontinalicaridinae". The group con- mented", are useless. endopod of males is fontinalis- tains P. P. borea different from that of females (see Fig. 5 A, C; fontinalis fontinalis, fontinalis P. meridionalis Zinzenco, 1971; Songeur, 1961; Rouch, 1990) Kiefer, 1960, fontinalis Rouch, P. 1925, P. even in the species Parastenocaris fontinalis 1990, aquaeductus Chappuis, psammi-
ca and P. n. bohé- Schnitter & Chappuis, 1915 (cf. Herzog, 1938). Songeur, 1961, hispanica sp. (P. is of p. Furthermore, the endopod P2 is 1-segmented in all mica Sterba, 1968, a junior synonym
known Parastenocarididae, this being a symple- fontinalis).
All these share the same construction of the siomorphy within the family (cf. Martínez Arbizu taxa
male P4 as described above. However, P. & Moura, 1994). endopod
differs from the other taxa in some Lang grouped 5 species together in the fonti- aquaeductus characters: Male P4 fused nalis-group, of which P. entzi Török, 1935, important (1) endopod Martinez Arbizu - Parastenocaris 222 P. hispanica n. sp. from hyporheic groundwaters in Spain
7. Parastenocaris n. male telson lateral male telson from Sta. Rouch 1978 n° 8 lateral Fig. hispanica sp.: A, view; B, view; C, maxilla;
maxillule. Scale bar 10 D, µm.
seta with subordinate spinules on both margins, "apophysis" as long as wide. (6) Female P4 basis
the basis with between not only on outer margin. (2) Male P3 a row of spinules exopod and endo- with a row of spinules on inner margin. (3) Male pod.
P3 Characters 1-6 the exopodal portion of segment 1 ends distally on represent plesiomorphic inner with 2 little Male P4 within the This allows us to margin spinules. (4) state fontinalis-group. .
1 ends of define well-founded for the exopodal segment distally in a row a monophyletic group spinules on inner margin. (5) Male P3 exopodal remaining species. However, this does not exclude Contributions to Zoology, 66 (4) - 1997 223
Table I. Character matrix.
Characters 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
P. hispanica 111 1 1 100 11 00 0000
P. meridionalis 111 11 1110000 1000
0010 P. psammica 111 11 1110011
P.f. fontinalis 111 II 1110011 0100
P. f. borea 111 11 1110011 0101
the that another of of (5) Male P3 exopod "apophysis" twice as long as wide (ap); the possibility group species and in the "apophysis" is as long as wide (pi) in P. aquaeductus Parastenocarididae, characterized by an advanced fifth copepodid stage of P. hispanica (Fig. 4F). transformation of P4 the endopod male, e.g., group without between and (6) Female P4 basis any spinules exopod around P. 1969 kabyloides Enckell, (Martinez at endopod (ap); P. aquaeductus has a row of spinules this in be the sister of the Arbizu, prep.), may group position (pi).
Male P4 1 inner with 4-5 fontinalis- group excluding P. aquaeductus. This (7) exopod subterminally on margin strong spinules (ap); P. hispanica has 8 slender spinules (pi). species would then be the sister group of a taxon (8) Male P4 coxa with 2-3 strong spinules (ap); P. aquaeductus including the fontinalis- group and the group has 5, P. hispanica has 6 slender spinules (pi). around P. kabyloides. For this reason P. aquae- (9) Male P3 exopod, "apophysis" with a protuberance on outer ductus is excluded from the following phyloge- margin (ap); this protuberance is absent (pi) in P. aquaeductus,
it is considered P. P. P. and P. meridionalis. netic discussion, although an ap- psammica, f. fontinalis, f. borea, f.
(10) Male P3 curved at an angle propriate outgroup to polarize the characters with- exopod "apophysis" inwardly P.of60° (ap); P. aquaeductus, P. psammica, P. f. fontinalis, f. in the fontinalis-group. borea, and P. f. meridionalis have anangle of 45° (pi).
(11) Male P5 slender (ap); P. aquaeductus, P. f. meridionalis,
outer and P. hispanica have a broad P5 with convex margin
Phylogenetic relationships within the (pl)- (12) Male P3 "thumb" shorter than "apophysis" (ap); P. fontinalis-group aquaeductus, P. f. meridionalis, and P. hispanica have a
"thumb" longer than "apophysis" (pi). 8 shows the within Fig. phylogenetic relationships (13) Female P5 without spinules at inner margin (ap); P.
the The different and fontinalis-group. subspecies are aquaeductus, P. psammica, P. f. fontinalis, P. f. borea, P.
have spinules at inner margin (pi). treated for this analysis as separate taxa. A charac- hispanica (14) Operculum triangular (ap); P. aquaeductus. P. psammica, ter matrix is given in Table I. The characters are P. f. meridionalis, and P. hispanica have a round operculum listed and polarized as follows: (pl).
Male P5 inner distal with (15) corner two setae (ap); P. aquae-
(1) Male P4 endopod fused seta with subordinate spinules only ductus, P. f. fontinalis, P. f. borea, P. f. meridionalis, and P.
and other "Fontinali- have setae at this No setae on outer margin (ap); P. aquaeductus hispanica no position (pl). are pre-
sent caridinae" such as P. kabyloides have subordinate spinules on in the ground pattern of Parastenocarididae (Martinez
both margins (pi). Arbizu & Moura, 1994). The acquisition of these 2 setae in P.
is considered ofthis this (2) Male P3 basis without row ofspinules (ap);.P. aquaeductus, psammica an autapomorphy species,
P. kabyloides, and most Parastenocarididae have a row of being the most parsimonious assumption.
(16) Male with a and at inner spinules on inner margin of basis of P3 (pi), this being a P5 strong large spinule margin
symplesiomorphy of the family. (ap); P. aquaeductus, P. psammica, P. f. fontinalis, P. f. meri-
and have ofsmall at this (3) Male P3 exopod portion of segment 1 without spinules on dionalis, P. hispanica a row spinules
inner 2 in adult P. margin (ap); spinules are present aquae- position (pi).
ductus (pi) and also in the fifth copepodid stage ofP. hispanica
(Fig. 4F), while these spinules get lost with the moult to the
adult. Biogeography (4) Male P4 exopod 1 row ofspinules on inner margin inserting
subterminally (ap); these spinules insert terminally (pi) in P. has The new species its known distribution in east- aquaeductus and in the fifth copepodid stage of P. hispanica ern Spain, the Pyrenees being its northern bound- (Fig. 5B), being relocated to subterminal position with the adult the French side ofthe moult. ary (Fig. 1). On (north) Pyr- 224 P. Martinez Arbizu - Parastenocaris in hispanica n. sp. from hyporheic groundwaters Spain
Fig. 8. Phylogenetic position ofP. hispanica n. sp. Explanation of characters in text.
Parastenocaris P. enees f. meridionalis has been col- tion of the French species P.f. meridionalis and
lected at the River Nert (Rouch, 1990), this being psammica. The subspecies P. f. borea is distrib-
the only record of this taxon. Parastenocaris uted in groundwaters that where covered with ice
psammica is described from different locations in during the last glaciation. Assuming the extinction
Lorraine, northern France (Songeur, 1961). The of most groundwater organisms due to the forma-
widest distribution has been recorded for Paraste- tion of permafrost in the soil during glacial times,
of this nocaris fontinalis, which is present all over Ger- we may conclude a colonization areas after
many, The Netherlands, northern France, Switzer- glaciation by a sub-population of P. fontinalis that
and Czechoslovakia land, (Janetzky et al., 1996), was better adapted to cold conditions.
its subspecies P. fontinalis borea being distributed
in northern Germany and The Netherlands.
A projection of the cladogram on a map showing Discussion
the approximate distributions of the species of the
shows a amount of concor- fontinalis-group high The polarization of the characters was done fol- dance between and distribution. The phylogeny lowing Hennig's criteria (Hennig, 1982: 98-116)
higher the latitude, the more derived the relatively and by outgroup comparison, not only with Para- This condition species (Fig. 9). suggests an origin stenocaris aquaeductus, but with all known Para-
ofthe fontinalis-group in southwestern Europe and from stenocarididae. Only some relevant examples
a slow colonization northwards. The first vicariant other taxa were considered in the text to explain
event is the of the the origin Pyrenees isolating the polarization of the characters chosen. In addi- basins south and north of it. Other groundwater tion, the Hennigian "criterion of ontogenetic char-
vicariant cannot be events explained satisfactorily acter precedence" has been successfully applied to
at the of of the distribu- present state knowledge polarize characters 3, 4, and 5. Some structures Contributions to Zoology, 66 (4) - 1997 225
in present in P. aquaeductus appear P. hispanica n.
sp. during ontogenetic development and are sec-
ondarily lost with the final moult to the adult. The
polarization ofthe characters as presented above is
also in concordance with the Hennigian "criterion
of the correlation of series of transformations"and
the "criterion of biogeographic progression". As a
result of this analysis, Parastenocaris meridionalis
Rouch (syn. P. fontinalis meridionalis Rouch),
be of P. cannot considered a subspecies fontinalis
is the sister of anymore. It group a taxon consisting
of P. and P. that psammica fontinalis, are separate
biological species as they are sympatric, having
been found together in the same sample (Songeur,
1961). P. meridionalis must therefore be con-
sidered a separate species, of which the Pyrenees
are the southern distribution limit. P. fontinalis
borea is further considered a subspecies of P. fon-
tinalis.
The phylogenetic relationships of the different
species-groups of Parastenocaris are unknown.
for the fam- Jakobi (1972) proposed a new system
ily Parastenocarididae by splitting it into 26 new
genera. This artificial system has been strongly
criticized by Schminke (1976b) and has not been
adopted by Dussart & Defaye ( 1990).
One of Jakobi's new genera was Fontinalicaris,
having as type species P. fontinalis. In this genus,
Jakobi included second P. a species, phyllophora Fig. 9. Projection of the cladogram (Fig. 8) on a map of the Noodt, 1954, that is quite unrelatedto P.fontinalis. western of middle part of Europe showing the distribution of
A of Parastenocaris will result in the the the revision rec- species belonging to fontinalis- group. 1, Parastenocaris
hispanica n. sp.; 2, P. meridionalis; 3, P. psammica; 4, P. ognition of a number of monophyletic groups, one fontinalis; 5, P. f. borea. of which will be the genus Fontinalicaris in-
cluding the species P. fontinalis, P. psammica, P-
meridionalis, and P. hispanica as is discussed unconditional support and continuous discussions especially here. Fontinalicaris with its type species Fontina- on parastenocaridid phylogeny. This work has been supported licaris will be the of the fontinalis type genus partially by a grant ofthe Deutscher Akademischer Austausch- Fontinalicaridinae advanced subfamily as by dienst (D.A.A.D.).
Schminke this I (1993). Waiting upon revision,
prefer to use the concept of species-group (in this
case rather than the Fonti- References fontinalis-group) genus
nalicaris.
1937. Subterrane Chappuis, P.A., Harpacticoiden aus Nord-
Spanien. Bull. Soc. Sei. Cluj., 8: 556-571.
Chappuis, P.A., 1942. Eine neue Methode zur Untersuchung
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Saint-Girons, France), who kindly loaned me abundant material tacés Copépodes des eaux intérieures. III. Harpacticoïdes. for this work. I wish to thank Prof. Dr. H.K. Schminke for his Crustaceana, Suppl. 16: 1-384. 226 P. Martinez Arbizu - Parastenocaris hispanica n. sp. from hyporheic groundwaters in Spain
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