Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
Alekseev, V., Dumont, H. J., Pensaert, J., Baribwegure, D. & Vaneteren J. R. (2006). A redescription of Eucyclops serrulatus (Fischer, 1851) (Crustacea: Copepoda: Cyclopoida) and some related
taxa, with a phylogeny of the E. serrulatus-group. Zoologica Scripta, 35, 123147.
Eucyclops serrulatus (Fischer, 1851), the type species of the genus Eucyclops, is redescribed from
specimens found in the St. Petersburg area, Russia (type locality) and compared with specimens
from Siberia, western Europe and North Africa. A neotype is selected. Cultures were set up,
and interpopulation hybridization as well as hybridization with related species was attempted.
The classical description of external body morphology was combined with pore signature
mapping and with DNA nuclear small subunit (18S) ribosomal gene sequence analysis.
Comparisons with E. dumonti Alekseev, 2000, E. hadjebensis Kiefer, E. speratus Lilljeborg,
E. turcomanus Lindberg, and E. pectinifer (Cragin, 1883) were carried out. A phylogenetic tree
based on molecular information shows that E. serrulatus and E. speratus should be regarded as
separate species. E. dumonti also deserves species status, but not E. hadjebensis. A cladistic tree
based on the pore pattern of the cephalosome agrees well with a tree based on the sequence
of the 18S rDNA gene. Cephalosome (and probably metasome) pore patterns seem useful to
elucidate relationships within genera, while urosomal pore patterns better reect the relationship
between genera. E. serrulatus occurs in three morphological forms over most of its range; one of
these (C) might be a rare (recessive) morphotype, while forms A and B differ in microhabitat
choice, but hybridize when living together. The same polymorphism also occurs in an American
species (E. prionophorus), and therefore two hypotheses regarding its origin are advanced: either
forms A and B evolved during the glacial episode (Pleistocene origin), separately on both sides of
the Atlantic, or the polymorphism was already present in the ancestor of the serrulatus-group,
and was later lost in some but not in all species (Pliocene origin.)
Victor Alekseev, Zoological Institute of the Russian Academy of Sciences, 199034, St. Petersburg, Russia.
E-mail: valekseev@yahoo.com
Henri J. Dumont, Deo Baribwegure & Jacques R. Vaneteren, Department of Biology, Ghent University,
Ledeganckstraat, B-9000, Ghent, Belgium
Jeannine Pensaert, Belgian Federal Public Service Health, Food Safety and Environment, Victor
Hortalaan, 1060 Brussels, Belgium
Introduction
The taxonomy of the large cyclopoid genus Eucyclops, burdened
with faulty species descriptions, is complex and chaotic. To
a certain extent this is because the original description of the
type species of the genus, Eucyclops serrulatus (Fischer, 1851),
was incomplete. This taxon was discovered in the mid-19th
century in a pond at Petershoff, the summer resort of the
Tsar close to St. Petersburg, Russia (Fischer 1851). An earlier
description, Cyclops agilis Koch, 1838, sometimes equated with
E. serrulatus, was published in the third volume of Kochs
study on the fauna from Germany. Cyclops agilis is only one of
a series of copepods described by Koch, very few of which are
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
123
genetics, we also collected animals in the Orlov Pond in Petershoff, in a roadside ditch on the way between Petershoff and
St. Petersburg, in the small River Sablinka, 50 km from St.
Petersburg in the direction of Moscow, and in the small River
Ravan, 100 km from St. Petersburg in the direction of
Moscow (Fig. 2, site 2). These samples yielded E. serrulatus,
but also E. speratus Lill. and E. macruroides, and were used in
DNA studies and for comparative morphology.
Western Europe (Ghent, Belgium: 5105N, 345E). For
hybridization, pore signature mapping and molecular genetics
we collected material in the pond of the botanical garden of
Ghent University in June 1999, as well as animals from
roadside ditches in the vicinity of Ghent in July 1999.
Siberia (Tumen, Western Asian Russia: 57N, 67E). A pond
in the park of the Tumen Agriculture Academy, May 1999,
sampled to collect material for hybridization experiments,
pore mapping and molecular genetics. For comparative
morphological research we also used a population collected
in temporary ponds, near roadside ditches and marshes close
to Tumen in AprilMay 1999.
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
125
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
Results
1. Eucyclops serrulatus (Fischer, 1851)
Synonymy: see Dussart & Defaye, 1985.
Eucyclops asymmetricus Dumont & Pensaert in Dumont
(1979) (nomen nudum).
Probable new synonyms: Eucyclops speratus Dumont & Decraemer, 1977; Eucyclops speratus ifniensis Dumont & Decraemer, 1977.
Material examined. Neotype: dissected female neotype, catalogue number 55031, dissected male (accession number 55032),
8 September 1983, from pond in Petershoff, possibly S. Fischers
type locality (Fischer 1851). The material was deposited in
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
127
Leg 1
Leg 2
Leg 3
Leg 4
Coxa
Basis
Exopodite
Endopodite
01
0-I
0-I
0-I
1-I
1-0
1-0
1-0
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
129
Male: body length 810 m, with caudal setae 1230 m. Cephalosome (Fig. 5: 2) 1.3 times as long as wide, with maximal
width close to its posterior end. Last segment of prosome
smooth, last urosomal segment with row of denticles on
caudal side. Caudal rami (Fig. 11: 7) 4.3 times longer than
wide, without lateral spinules. Slender innermost terminal seta
about twice the length of spine-like outermost seta. Lateral
seta shifted to dorsal side, with several spinules at base.
Dorsal seta near insertion of innermost seta, about 0.8 times
as long as outermost seta. Antennule (Fig. 9: 2) 14-segmented
with six setae and three aesthetascs on rst segment. Segments
2, 3, 4, 6 and 10 also with aesthetascs. Antennal basipodite
(not shown) basically as in female, with 4 6 long setules
posteriorly (N1, N2); anteriorly with three rows of strong
130
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
as long as spine. P6 (Fig. 11: 6) with inner spine and two setae
with length proportions, beginning from outer seta, 1/0.75/1.1.
Table 1 Variation of microcharacters on antennal basipodite in Eucyclops. Hair-setae in roman, denticles in arabic numerals; numbers of
characters correspond to Fig. 2.
Species
10
11
12
13
14
15
16
E. serrulatus form A
St. Petersburg, Russia
E. serrulatus form B
St. Petersburg
E.serrulatus form A
Norway
E. hadjebensis
Atlas Mts, Morocco
E. speratus
Lund, Sweden
E. dumonti
Central Mongolia
E. turcomanus
Herat Prov., Afganistan
E. pectinifer
Philadelphia, USA
E. macruroides
Lund, Sweden
VIVIII
III
89
78
1216
14
45
58
56
68
58
45
VIX
IIIIV
13
45
34
56
56
34
45
IV
III
610
89
1518
67
56
67
36
67
VI
12
12
58
35
45
25
46
15
47
68
89
45
58
VIII
III
18
VII
18
8+I
26
311
78
912
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
131
Table 2 Micro-characters on coxa and intercoxal sclerite of P4 in Eucyclops. Coding corresponds to Fig. 2.
Species
C+D
II
III
Hairsetae on
basipodite
E. serrulatus A StPetsb
E. serrulatus B StPetsb
E. serrulatus A Norway
E. serrulatus A Ghent
E. s. hadjebensis
E. speratus
E. dumonti
E. macruroides
E. turcomanus
E. pectinifer
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
45
5
5
4
5
5
23
12
1214
14
13
1011
27
1012
20
10
10
24
3
34
2
2
8
2
6
4
5
N
N
N
N
N
N
N
N
N
N
N
N
N
N
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
N
N
N
N
Y
Y
Y
Y
LH
SH
SH
H
LH
LH
SH
H
LH
LH
SH
SH
SH
H
LH
H
SH
H
SH
SH
LH
LH
LH
H
LH
LH
SH
H
LH
LH
Y
Y
Y
Y
Y
Y
N
Y
Y
Y
4
3
Y, present; N, absent; ?, character not seen; H, hair-setae; LH, long hair-setae; SH, short hair-setae.
132
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
133
Combinations
Offspring in
first generation?
Hybrids in
offspring
Offspring in
second generation?
Type A Type B
Type A Type C
Type B Type C
Type A Type A
Type B Type B
Type C Type C
+
+
+
+
+
+
AB
A
B
A
B
C
+
+
+
+
+
+
Combinations
Sex ratio
Offspring in
first generation
Offspring in
second generation
Tumen males x
StPetsb females
Tumen females x
StPetsb males
Tumen males x
Tumen females
StPetsb males x
StPetsb females
3:6
6:3
3:6
6:3
Combinations
Sex ratio
Offspring in
first generation
Offspring in
second generation
StPetsb males x
Ghent Females
StPetsb females x
Ghent males
StPetsb male x
StPetsb females
Ghent males x
Ghent females
Tumen females x
Ghent males
3:6
6:3
6:3
6:3
6:3
Hybridization experiments
Intrapopulation hybridization: for these experiments forms A, B
and C from Ghent were used (Table 3). A preliminary survey on
E. serrulatus from the type locality in 1983 had revealed much
intrapopulation variability. Based on differences in caudal
seta construction, caudal rami proportion and presence of
spine patterns on the body surface, morphotypes A, B,
and C were identied (Figs 5, 6, Tables 1, 2). In 1999, these
forms were found to co-occur in the same water-bodies, in
ponds in Tavricheski Garden, St. Petersburg, in the botanical
garden of the University of Ghent and in the park of the Agriculture Academy of Tumen (A and B only). No reproductive
isolation was found among any of them. Hybridization of
forms A and B produced animals with intermediate characters
Table 6 Hybridization of three forms of Eucyclops serrulatus with E. speratus and E. macruroides (all from St. Petersburg, MayJune 1999).
Combinations
Females E. serrulatus Type A
Females E. serrulatus Type B
Females E. serrulatus Type C
Males E. serrulatus Type A
Males E. serrulatus Type B
Males E. serrulatus Type C
Females E. macruroides
Females E. speratus
134
E. speratus
males
E. speratus
females
NO
NO
NO
E. macruroides
males
NO
NO
NO
NO
NO
NO
NO
YES
E. macruroides
females
NO
NO
NO
YES
NO
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
on distal half reduced to between half and one third the length
of a ramus. Outermost seta with spinules on both sides,
innermost seta with short stiff setules (Fig. 13: 9). Antennule
of 12 segments with hyaline, nely denticulated membrane
along three distalmost segments. Mouthparts (not shown) as
in E. serrulatus. Basipodite of antenna with six hair-setae in
position N1 and one setule in position N2 posteriorly, most
other characters on both sides as in E. serrulatus (Table 1).
P4: spine of coxopodite with a gap in the setules lining its
outer margin, intercoxal sclerite with long setules along
outer margin, other characters of P4 coxopodite as in
E. serrulatus (Table 2). A feature stressed as diagnostic by
Kiefer was that the setae on the distal segment of P4
exopodite are knife-shaped, with the constricted margin set
with spinules, while the inner rim has normal setules. This,
in fact applies to P3 and P2 as well; only P1 has more or
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
135
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
137
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
Fig. 17 Eucyclops pectinifer (Cragin, 1883) neotype from Gees Lake near Philadelphia, PA, USA. 18 female, 915 male: 1 body, dorsal; 2
urosomites and caudal rami, ventral; 3 P5; 4 maxillule; 5 maxilliped; 6 maxilla; 7, 8 A2 basipodite, anterior and posterior, respectively;
9 body, dorsal; 10 caudal rami, dorsal; 11 urosomites with P5P6; 12, 13 basipodite of A2, posterior and anterior, respectively;
14 P1; 15 distal segment of P4 endopodite. Scale bar: 1, 9 = 400 m; 2, 10 = 200 m, 14 = 150 m; 14 = 75 m; 36, 12, 13, 15 = 70 m.
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
139
Leg 1
Leg 2
Leg 3
Leg 4
Coxa
Basis
Exopodite
Endopodite
01
0-I
0-I
0-I
1-I
1-0
1-0
1-0
representing formula A B C + D E H (Fig. 18: 8). Intercoxal sclerite of P1 (Fig. 18: 5) with two protuberances, a row of
small denticles around midway and two groups of nest spinules
on body of protuberances, not extending beyond edge. Intercoxal sclerites of P2P3 also with high protuberances on free
edge and groups of setules as shown in Fig. 18: 6,7. Intercoxal
sclerite of P4 (Fig. 18: 8) without protuberances with row of
stiff incurved setae near outer edge and group of spinules at
mid-length; two groups of incurved setae along edge, two
groups of setules and spinules on body of sclerite. Coxa of P4
with strong spine bearing stiff setae on inner margin and only
12 setae on outer margin. Setae of P4Exp3 and apical seta of
P4Enp3 knife-shaped, as in type B of E. serrulatus. All other
limb setae cylindrical. P5 (Fig. 17: 3) 1-segmented with short
inner spine and two setae; outer seta subequal in length to
spine, middle seta about 2.5 times as long as spine. Egg sacs
(Fig. 17: 1) divergent, with 2025 eggs each.
Male: body length 716 m; with caudal setae, 1266 m.
Cephalosome (Fig. 17: 9) 1.2 times as long as wide, with
maximal width close to its posterior end. Last segment of
prosome without lateral setae. Caudal rami (Fig. 17: 10) 4.3
times longer than wide, seta III about half of seta VI; V seta
2.8 times longer than seta IV. Antennule 14-segmented;
antennal basipodite (Fig. 17: 1213) as in female, with four
long setules posterior (N1, N2) and two diagonal rows of
45 spines each; anterior with fewer shorter and slender spines
than in female. Morphology of mouth parts (not shown)
and P1 (Fig. 17: 14) as in female. Coxopodites and intercoxal
sclerites P1 as shown in Fig. 17: 14. Intercoxal sclerites P4
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
Female pore pattern (Figs 19, 20): rostral and postrostral positions as in E. serrulatus, and therefore likely to be diagnostic
at the genus level only. Cephalosome with reduced pore
complement. Positions 8, 10, 11, 15, 19 (?), 22, 23 (?), 31, 32 and
34 missing. The last couple of pores particularly signicant,
because they are part of the posterior U-shape, which is only
partly expressed in this species. Some positions are single,
against double in the serrulatus-group (4, 5 and 30), but here
as well as in other species, left-right asymmetries occur (e.g.
position 2, which could be either a singleton or a doubleton).
The metasomal pattern is puzzling, because many small, asymmetric pores were found. It appears that metasome segments
1 and 2 have at least two extra pairs of lateral pores, while
metasomal segment 3 has extra pores in the basal zone of the
mid-dorsum. Apparently, this species shows a tendency for a
E. pectinifer
Range
10001080
1.001.02
4.855.35
1.001.14
0.560.59
2.002.63
3.123.21
1.681.85
0.911.21
1.231.54
Range
12
12
24
24
24
19
24
24
24
22
9701130
0.8421.09
3.866.00
0.951.53
0.5130.956
1.2671.568
2.133.00
1.2141.583
0.8821.125
1.2671.54
Mean SD
1043 0.018
0.942 0.034
4.79 0.061
1.486 0.16
0.690 0.126
1.482 0.085
2.54 0.247
1.317 0.105
1.04 0.069
1.427 0.082
n
12
12
28
27
28
22
28
28
28
12
SD, standard deviation; n, number; StPetsb, St. Petersburg; L/W, length to width ratio; gen. som., genital somite; c.r., caudal rami; Si/Se, ratio of innermost setae to outermost seta;
Si/L c.r., ratio of innermost setae to caudal ramus. Smi/Sme, ratio of inner medial setae to outer medial seta; endP4, terminal endopodite segment of P4; SPi/SPe, internal to outer
spine ratio; SPi/L, ratio of internal spine to length of distal endopodite segment of P4; AS1/AS3, ratio of first to third segment of antennula.
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
141
Phylogenetic relationships
Cephalosome pore pattern-based phylogeny: the coding system
dening the state of the pore-characters is shown in Table 8.
A consensus tree including eight taxa, with E. macruroides and
A. gibsoni as outgroups, is shown in Fig. 21: it reveals all taxa
in the serrulatus-group to be closely related. E. serrulatus and
E. hadjebensis cluster together, and form a clade distinct from
that containing E. speratus, E. dumonti and E. pectinifer. Eucylops
dumonti and E. speratus come out as close relatives, a position
also supported by morphology. E. turcomanus is the most
distant member of this group although with low bootstrap
support. Eucyclops macruroides and Afrocyclops gibsoni have high
bootstrap support and belong to a clade that could be called
a non-serrulatus group.
142
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
Table 8 Coding system for pore positions on dorsum of cephalosome (0 = position vacant, 1 = 1 pore, 2 = doublet; 3 = triplet).
123456789012345678901234567890123456789012
222222122222122222222222222222222222200000
222222122022220222222212222222212222200000
202222122111220221212202222222212222200000
212222122211220221122212222122222222221300
212212122002120113220232110100212222200000
222112101002120222022002222122102022000000
211222122012210212222012222122222220100021
212022102220020002311102200010112022200000
Discussion
By any standard and method, considerable intra- and interpopulation variability was encountered in E. serrulatus. The type
population contained three forms (A, B and C) that differ in
setae morphology of the caudal rami and the swimming legs
(Table 1), but all easily mated and had fertile offspring across
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
143
Table 9 Gene length, GC content and accession number of the ribosomal 18S gene and geographical origin of the species used in this study.
Species
18S (bp)
GC percentage
EMBL acc.
Geographical origin
Collector
Eucyclops serrulatus FA
Eucyclops serrulatus FA FB
Eucyclops serrulatus FA
Eucyclops serrulatus FA
Eucyclops serrulatus
Eucyclops speratus
Eucyclops dumonti
Eucyclops macruroides
Ectocyclops polyspinosus
Macrocyclops albidus
1809
1809
1809
1809
1809
1809
1808
1809
1808
1808
49.25
49.25
49.25
49.25
49.25
49.14
49.28
49.25
49.56
50.61
AJ746330
AJ746331
AJ746332
AJ746328
L81940
AJ746333
AJ746335
AJ746329
AJ746336
AJ746334
V. Alekseev
V. Alekseev
V. Alekseev
V. Alekseev
T. Spears
V. Alekseev
V. Alekseev
V. Alekseev
V. Alekseev
V. Alekseev
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
145
Acknowledgements
This study was supported by NATO Linkage Grant no. 974986
to H. J. Dumont, a Biodiversity grant from the Presidium of
the Russian Academy of Sciences, and grant N 04-04-49053
from RFBR to V. Alekseev. Dr P. Weekers (Ghent) assisted
146
with the technical molecular work and calculated the phylogenetic trees. Dr Hans Mittmann (Karlsruhe Natural History
Museum, Germany) helped one of us (V. A.) during his examination of Kiefers collection. Dr Frank Ferrari (Smithsonian
Institution, Washington DC, USA) kindly assisted us with
inquiries into the fate of Cragins collections. Dr Trisha Spears
(Florida, USA) provided information about the origin of an
Eucyclops serrulatus from the US, used in gene sequencing.
References
Alekseev, V. R. (1985). Rol photoperioda v formirovanii diapauzu y
presnovodnuh tsiklopov (Role of photoperiod responses in
diapause in freshwater cyclopids). Trudy Gosniorh, 223, 95104 (in
Russian, English summary).
Alekseev, V. R. (1989). Eucyclops arcanus sp. n. (Copepoda, Cyclopoida)
from waterbodies of Bolshezemelskaya Tundra and Prebaikalia.
Zoologicheskii Zhurnal, 69, 135139 (in Russian, English summary).
Alekseev, V. (1990). Diapauza u rakoobraznuh (Diapause in Crustacea).
Moscow: Nauka Academic Publishers, 169 pp. (in Russian, English
summary).
Alekseev, V. (2000). Eucyclops dumonti sp. nov. from Central Mongolia.
Hydrobiologia, 441, 6371.
Alekseev, V. & Naumova, E. (2005). Express methods for integumental pore study. Zoologicheskii Zhurnal, 5, 234247 (in Russian).
Baribwegure, D. & Dumont, H. J. (1999). The use of the integumental
pore signature in the characterisation of the species of the genus
Thermocyclops Kiefer, 1927: the case of Thermocyclops emini (Mrazek,
1895) (Crustacea: Copepoda: Cyclopoida). Belgian Journal of Zoology,
129, 187200.
Baribwegure, D. & Dumont, H. J. (2001). The integumental pore
signature of Thermocyclops oblongatus (Sars, 1927) and Thermocyclops
neglectus (Sars, 1909), with the description of Thermocyclops africae
new species, and a comparison with Thermocyclops emini (Mrazek,
1895). Hydrobiologia, 458, 201220.
Cragin, F. W. (1883). A contribution to the history of the freshwater
Copepoda. Transactions of the Kansas Academy of Sciences, 8, 115.
De Lattin, G. (1967). Grundriss der Zoogeographie. Jena: Fischer, 602 pp.
Dumont, H. J. (1979). Limnologie van Sahara en Sahel. Unpublished
D. Sci. Thesis, Ghent University, 557 pp.
Dumont, H. J. & Decraemer, W. (1977). On the continental copepod
fauna of Morocco. Hydrobiologia, 52, 257278.
Dumont, H. J., Pensaert, J. & Van de Velde, I. (1981). The crustacean
zooplankton of Mali (West Africa). Hydrobiologia, 80, 161187.
Dussart, B. H. (1986). Some Crustacea Copepoda from Venezuela.
Hydrobiologia, 113, 2567.
Dussart, B. & Defaye, D. (1985). Rpertoire Mondial des Coppodes
Cyclopoides. Paris: CNRS, 236 pp.
Einsle, U. (1985). A further criterion for the identication of species
in the genus Cyclops s. str. (Copepoda, Cyclopoida). Crustaceana,
49, 299309.
Farris, J. S. (1989). The retention index and the rescaled consistency
index. Cladistics, 5, 417 419.
Felsenstein, J. (1985). Condence limits on phylogenies: an approach
using the bootstrap. Evolution, 39, 783791.
Fiers, F., Reid, J. W., Iliffe, T. M. & Suarez-Morales, E. (1996). New
hypogean cyclopoid copepods (Crustacea) from the Yucatan
Peninsula, Mexico. Contributions to Zoology, 66, 65102. Amsterdam:
SPB Academic Publishing.
Zoologica Scripta, 35, 2, March 2006, pp123 147 The Norwegian Academy of Science and Letters 2006
The Norwegian Academy of Science and Letters 2006 Zoologica Scripta, 35, 2, March 2006, pp123 147
147