Table Of Content''
BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 66: 93-132, 1996
BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGIE, 66: 93-132, 1996
A taxonomic revision of European Herpetocypris BRADY and NORMAN, 1889
(Crustacea, Ostracoda)
by M.E. GONZALEZ MOZO, K. MARTENS & A. BALTANAS
Abstract Mediterranee. Des populations parthenogenetiques vi vent dans
toute !'Europe. Cette espece presente une plus grande variabilite
The European representatives of the genus Herpelocypris are morphologique. Herpetocypris mateusorum est limite a l'Iberie
revised, based on extant type material and various new collec occidentale. Herpelocypris chevreuxi-intermedia-helenae est un
tions. The revision is based on the hypothesis that species with groupe d'especes tres proches et que !'on rencontre dans toute
syngamic populations will show large intra-specific mor !'Europe, au moins H. helenae a apparemment une distribution
phological variability and that parthenogenetic taxa will have globale.
both less intra-specific variability and small morphological dif Certaines particularites morphologiques de ce genre sont
ferences between taxa. Of the nearly 20 extant nominal species, discutees, de meme que Ia variabilite de plusieurs "caracteres
diagnostiques". Quelques remarques sur l'evolution de ce genre
subspecies and varieties, 6 recent species are retained.
sont proposees.
Herpelocypris replans, with both the common nominal subspecies
Mots-clefs: Taxonomie, morphologie, distribution, modes de
and H. replans curvata (known from its type locality only), occurs
reproduction, syngamie, parthenogenese.
in most of Europe, north of the Mediterranean. Herpetocypris
brevicaudata is the only species in which syngamic populations
occur, these are thus far only found in the western part of the
Mediterranean. Parthenogenetic populations occur throughout
Introduction
Europe. This species shows larger morphological variability.
Herpetocypris maleusorum is confined to western Iberia.
Herpetocypris chevreuxi-intermedia-helenae form a closely related The genus Herpetocypris BRADY & NORMAN, 1889 is
species cluster, which occurs throughout Europe, at least H. common in small to medium-sized waterbodies and has
helenae appears to have a global distribution. thus far been reported from Eurasia, North and South
Some morphological peculiarities of this genus are discussed, America and northern and southern Africa. In Europe,
as is the variability in several of the so-called diagnostic about 20 species, subspecies and varieties have thus far
characters. Some remarks on the evolution in this genus are
been described, and most of these have at various occa
offered.
sions been lowered or raised in taxonomic rank or were
Key words: Taxonomy, morphology, distribution, reproductive
synonymized. Most taxa have been distinguished on the
modes, syngamy, parthenogenesis.
basis of carapace shape and size, length of nata tory setae
on A2, length of setae on Mx2-palp and size and number
Resume of spines on the ventral margin of the furcal rami. At pre
sent, the taxonomy of this genus in Europe is highly con
Les repn!sentants europeens du genre Herpetocypris sont revises, fused, especially for the Mediterranean populations, and
sur Ia base du materiel-type existant et de diverses nouvelles a revision is urgently required (GONZALEZ Mozo et al.
collections. La revision est basee sur !'hypothese que les especes 1996).
ayant des populations syngamiques ont une plus grande A European research network, consisting of six
variabilite morphologique intra-specifique et que les taxa par laboratories in five countries, has investigated the evolu
thenogenetiques presentent a Ia fois une plus petite variabilite tionary ecology of reproductive modes of the three extant
intra-specifique et des differences morphologiques peu pro
lineages of non-marine ostracods with a variety of techni
noncees entre les taxa. Parmi les quelque 20 especes, sous-especes
ques: morphological and morphometrical analyses, clonal
et varietes nominales existantes, 6 especes recentes sont retenues.
autecology, starch-gel electrophoresis of allozymes and
Herpetocypris reptans, avec aussi bien Ia sous-espece nominale
amplification and automated sequencing of both
commune, que H. replans curvata (connu uniquement de sa
mitochondrial and nuclear DNA. Although not one of
localite-type), apparalt dans Ia majeure partie de !'Europe, au
nord de Ia Mediterranee. Herpelocypris brevicaudata est Ia seule the initially selected network genera, the present tax
espece presentant des populations syngamiques, rencontrees onomic revision of Herpetocypris was conducted in its
jusqu'a present uniquement dans Ia partie occidentale de Ia framework and this for the following reason: (1) the
I I
94 M.E. GONZALEZ MOZO, K. MARTENS & A. BALTANAS
Cypridoidea constitute 70-80% of the specific diversity in 3. Material
non-marine habitats, and it was thus felt that this group
should be investigated on more than one representative; The present revision relies on type and non-type materials
(2) Herpetocypris is a classical example of a group with from museums, as well as on new collections from Europe
geographically segregated reproductive modes: par (Spain, Portugal, Belgium, Italy, England, Switzerland),
thenogenetic populations occur throughout Europe, but Israel, Africa (Algeria, St Helena, South Africa), and
bisexual populations are found only around the Mediter North America (Canada). A list of these collections is
ranean. (3) Several exclusively parthenogenetic species given in an appendix. Lists of synonymies in the text deal
have been reported. with literature on European localities only.
The present revision is thus functional at several levels:
(1) to assess the validity of geographical parthenogenesis
in this lineage; (2) to test the applicability of the concept Taxonomic descriptions
of clonal taxonomy to this lineage and (3) to solve the
present taxonomic muddle in this group and to present Class Ostracoda, LATREILLE, 1806
a workable classification of identifiable units. Subclass Podocopa G.W. MOLLER, 1894
The present paper presents the taxonomic and mor Order Podocopida SARS, 1866
phological results of the revision. Applicability for Surorder Podocopina SARS, 1866
geographical parthenogenesis and clonal taxonomy will Infraorder Cypridocopina JONES, 1901
be more extensively discussed elsewhere. Superfamily Cypridoidea BAIRD, 1845
Family Cyprididae BAIRD, 1845
Material & Methods Subfamily Herpetocypridinae KAUFMANN, 1900
= Stenocypridinae FERGUSON, 1964
I. List of Museums and collections
BM Natural History Museum, London, U.K. Diagnosis: Large (1-2 mm), mostly elongated and laterally
MHNG Musee d'Histoire naturelle de Geneve compressed ostracods; marginal valves structures mostly
KBIN Koninklijk Belgisch Instituut voor well developed; branched pore canals and marginal sep
Natuurwetenschappen, Brussels, Belgium tae present in some genera. Antennae in males with claw
(O.C. = Ostracod Collection) GM developed into a comb-like structure, with one row
MCSN Museo Civico di Storia Naturale, Firenze of strong teeth. Furcae well developed, symmetrical or
MNIP Museum fur Naturkunde, Institut fur PaHion asymmetrical; furcal attachment with a triangular basal
tologie, Berlin reinforcement (in some genera only weakly developed).
ZMB Zoologisches Museum, Berlin, Germany Hernipenis with large and sclerotized bladder-like part 'c'
ZMO Zoological Museum, Oslo, Norway of the labyrinth, post-labyrinthal internal sperrniduct with
a variable number of additional, circular whorls (1-4).
2. Abbreviations used in text and figures
AI = Antennula. A2 = Antenna. Cp = carapace. db = Extant genera: Herpetocypris BRADY & NORMAN, 1889;
dorsal branch of furcal attachment. dms = dorsal lobe of Acocypris VAVRA, 1895; Stenocypris, SARS, 1889;
lateral shield of hemipenis. f1 = flange. Fu = furca. H = Chrissia HARTMANN, 1957 (syn.: Gesa HARTMANN,
height of valves. il = inner list. im = inner margin. L = 1957); Parastenocypris HARTMANN, 1964; Candonocypris
length of valves. lc = line of concrescence. Is = lateral SARS, 1894; Ilyodromus SARS, 1894; Humphcypris
shield of hernipenis. LV = left valve. Md = Mandibula. MARTENS, 1996; Somalicypris MARTENS, 1996;
ms =medial shield ofhernipenis. M x 1 = Maxillula. Mx2 Ampullocypris DE DECKKER, 1981 ; Psychrodromus
= Maxilla. ol = outer list. pfz = primary fused zone. r = DANIELOPOL & MCKENZIE, 1977; Stenocypria G.W.
Rome organ. R V = right valve. sfz = secondary fused MOLLER, 190 I.
zone. sl = selvage. Tl = first thoracopod. T2 = second
thoracopod. vb = ventral branch of furcal attachment.
vms = ventral lobe of lateral shield of hemipenis. vm = Distribution: This group occurs on all continents, except
valve margin. Antarctica. None of the genera, however, is cosmopolitan.
Herpetocypris has the widest distribution and occurs on
Chaetotaxy of the limbs follows the model proposed by all continents, except for AustraliSt and Antarctica.
BROODBAKKER & DANIELOPOL (1982), revised for the
A2 by MARTENS (1987).
Habitat: Representatives of this group are common in all
types of habitats, both temporary and permanent,
although they are not typical of the former. Some genera
are adapted to crenobiontic and even hypogean habitats.
I I
Taxonomy of European Herpetocypris 95
Herpetocypris BRADY & NORMAN, 1889 Type locality: Yetho1m Loch, Berwickshire (S.E.
=
Erpetocypris BRADY & NORMAN, 1889 Scotland), approximate coordinates: 55°33'N, 02°19'W.
=
Siphlocandona BRADY, 1910
=
Exuocypris MANDELSTAM, 1956
=
Abbreviated diagnosis: Large species (L 2.2-2. 7 mm),
Type species: Cypris rep tans BAIRD, 1835 with almost evenly rounded anterior and posterior
margins and straight to weakly curved dorsal margin; LV
with relatively weak inner list, calcified inner lamella
Diagnosis
Valves elongated and laterally compressed, L = 1.5-2.5 evenly sloping between valve margin and selvage. A2 with
mm; LV with largely inwardly displaced selvage and wide 1st n.s. the longest, usually reaching the end of the
fused zone, pore canals straight, marginal septae absent. penultimate segment, n.s. 2-5 of constant length, all
LV overlapping R V on all sides; this valve with well equally short (reaching to the middle of that segment),
developed il along anterior, ventral and posterior margin. 6th n.s. shorter. Fu with delicate claws; proximal setae
In closed carapace, valve margin of RV fitting between longer than in the other species, reaching beyond tip of
il and valve margin of LV. Mandibular scars small. ramus; ventral margin of ramus set with variable groups
AI with small Rome organ. Natatory setae on A2 very of teeth (mostly 4 groups of relatively large teeth and 2
short to long (beyond tips of claws), but always present. groups with smaller teeth).
Mxl-palp spatulate, with length about equal to distal
width. Seta d2 on Tl about 1.5-2 the length of seta dl.
Furca symmetrical, proximal seta present, not claw-like, Additional description
spines on ramus short and grouped. Furcal attachment Largest species in the genus, with anterior and posterior
with typical basal triangle, and an additional, but weakly margins evenly rounded and straight to weakly curved
built small branch towards the distal part of the main dorsal margin; ventral margin straight; LV longer and
branch. Hemipenis without hook-like process on medial higher than RV. LV with calcified inner lamella evenly
shield. sloping between valve margin and inner list, not forming
an abrupt angle; anterior inner list small (relative to other
species), set with few and short ridges. RV with inner list
Remarks: Several species have erroneously been described
present, but weakly developed (not produced away from
or transferred to this genus. Herpetoycpris strigata KAUF
inner lamella); this valve with a weak indention at the
MANN, 1900 and H. peregrina KAUFMANN, 1900 (a
antero-dorsal corner. Anterior and posterior calcified
homonym of H. peregrina CRONEBERG, 1894) are
inner lamellae relatively narrow in both valves.
synonyms of Tonnacypris lutaria (KOCH, 1838).
Al (Fig. lB) with first podomere with 2 subapical ven
Herpetocypris tumefacta KAUFMANN, 1900 is a synonym
tral and one medio-dorsal setae, all hirsute. Second
of Eucypris pigra (FISCHER, 1851).
podomere with small, but clear, medio-ventral Rome
organ and minute dorso-apical seta. Third podomere with
two apical setae, the ventral not reaching the edge of the
Herpetocypris reptans (BAIRD, 1835)
terminal segment; the dorsal one reaching well beyond
the edge of this segment. Fourth and fifth podomeres each
Herpetocypris reptans reptans (BAIRD, 1835)
with two dorsal and two ventral setae, the former almost
(Figs. 1(A-O,T), 2(A,B), 17(P), 18(A-G,J-M),)
twice as long as the latter. Sixth (penultimate) podomere
with four long and one shorter setae. Terminal segment
1835 Cypris replans n.sp. BAIRD: 99-100. Pl. 3 (I I) with one long aesthetasc Ya, a short, claw-like seta
I 845 Candona replans (BAIRD), I 53 (reaching beyond Ya) and two long setae. The longest
1889 Erpelocypris replans (BAIRD), BRADY & NORMAN 23,
nata tory setae 2-2.5 times the length of the last 5 segments
p. 84, Pl. 13(27)
combined.
1890 Herpetocypris replans (BAIRD), SARS 28, p. 17
A2 (Fig. 1D,N,T) with exopodite consisting of2 short and
1900 Herpetocypris replans (BAIRD), KAUFMANN: 282, Pl. 16
(1-3); Pl. 18 (21-26) I long seta, the latter reaching slightly beyond the tip of
1928 non Erpelocypris replans (BAIRD) sensu GAUTHIER, the first endopodal segment. Natatory setae short, mostly
fig. A-M. (= Herpetocypris brevicaudata) reaching halfway the penultimate segment, except for the
1938 Herpetocypris replans (BAIRD), KLIE: 125-126 (419-421) first n.s., always the longest and mostly reaching the tip
1947 Herpetocypris replans (BAIRD), ROME, fig. 7-15 of the penultimate segment. The latter segment with
1947 non Herpetocypris replans (BAIRD) sensu ROME, fig. 2 dorsal setae, 4 unequal ventral t-setae, 3 long subapical
16-20 (= male of Herpelocypris brevicaudata)
z-setae (not reaching beyond the tip of the endclaws),
?1955 Erpetocypris replans (BAIRD) ssp. aulicae n. ssp. LOTIIG:
2 long and one shorter claws (Gl-G3) and two
159, Pl. 18 (1)
aesthetascs: y I close to the t-setae and y2 near the inser
1969 Herpetocypris replans (BAIRD), NOCHTERLEIN: 263-265
1982 Herpetocypris replans (BAIRD), TETART: 162-163, fig. tion of the terminal segment. Terminal segment with one
89-103 long GM and one short (Gm) claw, aesthetasc y3 fused
1987 Herpetocypris replans (BAIRD), MEISCH: 106-107 with a seta (the seta longest) and one longer seta (g), the
1992 Herpetocypris replans (BAIRD), BALTANAS: 431, fig. 5 latter longer than Gm. All claws set with two rows of small
II
96 M.E. GONZALEZ MOZO, K. MARTENS & A. BALTANAS
Fig. 1. - Herpetocypris replans (female, Belgium).
A. Md-palp (OC.l895). B. Al (OC.l895). C. Tl, detail of apical chaetotaxy (OC.J894). D. A2 (OC.J895). E. Md-coxa
(OC.l895). F. AI, detail of apical chaetotaxy (OC.1895). G. Mxl, part of chaetotaxy (OC~J895). H. T2 (OC.l895).
I. Tl (OC.J894). J. T2, detail ofa pical chaetotaxy (Wl004). K. Fu, detail ofa pical chaetotaxy (OC.J894). L. Fu (OC.J894).
M. Fu (OC.J894). N. A2, detail of apical chaetotaxy (OC.l895). 0. Mx2 (OC.J895).
Scale: 100 fJ.mfor J; 102 fJ.mfor A, N; 200 fJ.mfor C, E, F, G, K, 0 ; 420 fJ.m for B, D, H, I, L, M.
II
Taxonomy of European Herpetocypris 97
)
Fig. l(bis). - Herpetocypris replans curvata (P-S, female, type, Switzerland), H. replans replans (T, female, Belgium).
P. Mx2-palp, distal part ( MG139). Q. Fu ( MGJ39 ). R. Fu ( MG139). S. A2, showing length ofnatatory setae (MG139).
T. A2, showing length of natat01y setae (MG139).
Scale: l02J1mfor P; 200 11m for Q-T.
spines, the 3 longest claws smooth in their distal fifth. segment with 4 subequal subapical smooth setae, one long
Md with coxa (Fig. IE) without special features. Md-palp and one minute ventra-apical setae and 3 smooth and one
(Fig. lA) of normal Cypridinid shape; alpha-seta narrow hirsute (gamma) apical setae. Terminal segment with three
and smooth, abruptly tapering towards the distal part, claws and three setae.
both beta and gamma-setae stout and hirsute. First seg Mxl (Fig. !G) with second palp segment spatulate, set
ment with respiratory plate (not drawn), 2 stout s-setae with three stout claws and three setae; first palp segment
(with alpha-seta in the middle) and an additional long with one medial and 5 subapical setae, two hirsute and
seta, normally smooth but here hirsute. Beta-bundle (bun three smooth. Third endite with 'Zahn-borsten' serrate.
dle of setae surrounding beta-seta on second palp seg First and second endites and respiratory plate without
ment) consisting of 3 smooth and 2 hirsute setae, all long. special features.
Second segment further with 3 dorsal setae. Penultimate Mx2 (Fig. 10) with two small a-setae, I smooth b- and
98 M.E. GONZALEZ MOZO, K. MARTENS & A. BALTANAS
d-setae. Endite (?)with a separate group of 3 smooth setae Differential diagnosis
between d-seta and 10 apical setae; palp with one long Herpetocypris rep tans is the largest species in the genus;
medial and two smaller (about half the length) lateral it can furthermore be distinguished from its congeners by
apical setae.Tl (Figs. IC,I) with seta d2 slightly longer the morphology of the anterior part of the LV, the short
than dl; penultimate segment divided, endclaw slightly n.s. on A2 of which the 1st is always the longest, and the
longer than length of three last segments combined. relatively long proximal furcal seta.
T2 (Figs. I H,J) a cleaning limb, as typical of the family.
Furcae (Figs. IK-M) symmetrical, rami set with variable
group of spines, proximal seta relatively long, reaching Herpetocypris repta11s curvata KAUFMANN, 1900
slightly beyond edge of ramus; distal seta longer than half (Figs. I (P-S), 18(H,I))
the length of the distal claw.
Males unknown. 1900 Erpetocypris replans var curvata, KAUFMANN, 282,
Pl. 16(4-5), Pl. 18(27).
Measurements
= = Type locality
New measurements: RV 2.30 mm (2.18-2.6) (n 6);
= = Mendriso, Switzerland.
LV 2.39 mm (2.23- 2.74) (n 7).
Measurements from literature: KAUFMANN (1900):
=
L 2.64 mm; H = 1.19 mm; W =I mm; KLIE (1938):
= = Diagnosis
L 2.1-2.6 mm; ROME (1947a): L 2.1-2.6 mm
All diagnostic features in valves and soft parts as in
H. replans replans, but smaller and with different valve
shape: dorsal margin more rounded.
Distribution
West and Central Europe.
Measurements (after KAUFMANN 1900):
L = 2.3 mm, H = 1.1 mm, W: 0.8 mm.
Habitat
The species occurs in ponds and small lakes; as well as
in slowly flowing waters. Distribution
Known from its type locality only.
Remarks
1. ROME (1947a) described a male of what he called H. Remarks
replans. Reinvestigation of the material used for this The population from Mendriso conforms in all details of
description, revealed that he was dealing with two species. soft part anatomy (typical length and configuration of the
At least some of the females he dealt with belonged to nata tory setae on the A2, length of proximal furcal seta,
H. replans s.s .. However, at least the male from Morocco etc) with H. replans replans. However, the shape of the
belonged to H. brevicaudata (see below), because of the valves is very different and analysis of several populations
largely inferior size, the different furca and especially the of H. replans replans showed that variability in valve
fact that in the male, the 5th n.s. of the A2 is the longest, shape in the nominal subspecies is minimal. We here con
not the 1st as in H. rep tans. It is here confirmed that all sider curvata a subspecies of H. replans.
males of Herpetocypris thus far described (GAUTHIER
1928 a, b, PETKOVSKI 1964, and various new populations
reported in the present paper) belong to H. brevicaudata; Herpetocypris brevicaudata KAUFMANN, 1900
all other species are exclusively known from par (Figs. 2(C,D), 3-9, 17(A-I), 19, 20)
thenogenetic populations.
2. LOITIG (1955) described a new subspecies of H. rep 1900 Herpetocypris brevicaudata n.s. KAUFMANN: 132, Pl. 16
tans, as Erpetocypris replans aulicae, from Elze and (6-7); Pl. 18 (28-31)
distinguished it from the nominal subspecies on a number 1928 partim male Erpetocypris rep tans (BAIRD) sensu
of valve characteristics (almost equally rounded anterior GAUTHIER, fig. A-M.
and posterior margin straightly convex RV, greatest 1932 Herpetocypris ghigii n.sp. MAS!, pp. 213-218, fig. a-e.
height in the middle, valves slightly higher) but provided 1938 Herpetocypris brevicaudata KAUFMANN, KLIE: 126-127,
fig. 422-425.
no figures. Because of this, the validity of this subspecies
1947 Herpetocypris lenta n.sp. ROME: 4-8, fig. 2 (a-g)
is difficult to interpret, while the size of this subspecies
1964 Herpetocypris brevicaudata KAUFMANN, PETKOVSKI:
also falls well into the range of H. replans replans. Until
24-27, fig. 56-60
illustrations can show that H. replans aulicae is a valid
1967 He1petocypris jlumendosa n.sp. ANIC HIN!: 17-22
and morphologically distinct subspecies, we here consider 1968 Herpetocypris brevicaudata KAUFMANN, SYWULA: 37-39,
it a synonym of H. replans replans. fig. 14 (a-b), 15
II
Taxonomy of European Herpetocypris 99
Fig. 2. - Herpetocypris replans replans (A-B, female, Belgium), Herpetocypris brevicaudata (C-D, female, Belgium).
A. LV, internal view (W1027). B. RV, internal view (WJ027). C. LV, internal view (OC.l954). D. RV, internal view
(OC.1954).
Scale: 410 pmfor C, D; 960 pmfor A,B.
100 M.E. GONZALEZ MOZO, K. MARTENS & A. BALTANAS
Fig. 3. - Herpetocypris brevicaudata (A-B, female, Algeria), Herpetocypris brevicaudata (C-D, male, Algeria).
A. RV, internal view (OC.J960). B. LV, internal view (OC.J960). C.RV, internal view (OC.J991). D. LV, internal view
(OC.J99J).E. RV, detail of internal view (OC.J960).
Scale: 102 pm for E; 410 pm for A, B, C, D.
I'
Taxonomy of European Herpetocypris 101
Fig. 4. - Herpetocypris brevicaudata (female, Belgium) A. Al (OC.J953). B. A2 (OC.J955 ). C. Md-palp ( OC.l954 ). D. A2, detail
of apical chaetotaxy ( OC.J955). E. Mxl, part ofc haetotaxy ( OC.J955 ). F. Md ( OC.J955 ). G. T2, detail ofa pical chaetotaxy
(OC.l953). H. Fu, detail ofa pical chaetotaxy (OC.I955). I. T2 (OC.J953). J. Mx2 (OC.J953). K. Fu attachment (OC.J954).
L. Tl (OC.J955). M. Fu (OC.J955). N. Fu (OC.J955).
Scale: 110 pmfor C, D, E, G, H, J; 156 pmfor A, B, F, I, K, L, M, N.
I I
102 M.E. GONZALEZ MOZO, K. MARTENS & A. BALTANAS
G
~
M
Fig. 5. - He1petocypris brevicaudata (female, Algeria)
A. AI (OC.I990). B. AI, detail of apical chaetotaxy (OC.I990). C. T2 (OC.I990). D. T2, detail of apical chaetotaxy
(OC.I990). E. Md-pa/p (OC.1990). F. A2 (OC.1990).G. A2 detail of apical chaetotaxy (OC.1990). H. Fu (OC.1990).I.
Fu (OC./990).1. Tl (OC.1990). K. Fu attachment (OC.1990). L. Md (OC./990). M. Mx1 (OC.I990). N. Fu, detail
of apical chaetotaxy (OC.I990). 0. Mx2 (OC./990). P. Md, detail of apical chaetotaxy (OC.I990).
Scale: 100 ~Jmfor G; 110 11m forB, D, E, M, N, 0, P; 156~Jmfor A, C, F, H, I, J, K, L.
