Acta%20Soc%20Zool%20Bohem%202002-03.pdf

Acta Soc. Zool. Bohem. 66: 161–168, 2002
ISSN 1211-376X
Morphological differentiation of some populations of the genus Cyclops
(Copepoda: Cyclopoida) from Bohemia (Czech Republic)
Zden!k BRANDL & Markéta LAVICKÁ* )
Faculty of Biological Sciences, University of South Bohemia, Branišovská 31,
CZ–370 05 "eské Bud!jovice, Czech Republic; e-mail: zdbrandl@bf.jcu.cz
Received June 3, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. Groups of spines forming the coxal ornamentation of the fourth pair of swimming legs and some
other morphological details are used to distinguish species of the genus Cyclops O. F. Müller, 1776, from
several habitats in the Czech Republic. The identity of populations based on these characters agrees with
that derived from allozyme analyses. Besides the easily distinguished species Cyclops vicinus Uljanin, 1857
and C. insignis Claus, 1857, three other species were found: C. furcifer Claus, 1857 (in temporary waters)
and probably C. strenuus Fischer, 1851 and C. abyssorum G. O. Sars, 1863 (in permanent waters).
Morphology, cryptic species, coxal ornamentation, species identification, habitats, Cyclops,
Czech Republic, Palaearctic region
INTRODUCTION
Although the detailed taxonomic differentiation of the common species of cladocerans in this
country has long tradition (e.g., Kurz 1874) and was greatly enhanced by Hrbá#ek in his early
publications (1959a, b), common cyclopoid copepods were believed to be a well-known group of
simply delimited species with good differentiating morphological characters (Šrámek-Hušek 1938,
1953). Even the most common limnetic genus Cyclops O. F. Müller, 1776, was supposed to be
represented by just two limnetic species, C. strenuus Fischer, 1851, and C. vicinus Uljanin, 1857,
and two others inhabiting rather small water bodies (C. insignis Claus, 1857, and C. furcifer Claus,
1857). The fifth species listed in the Bohemian faunistic records was originally described by
Šrámek-Hušek (1937) as C. bohemicus Šrámek-Hušek, 1937, from the lake "erné Jezero in the
Šumava Mountains and later assigned to the complex of C. abyssorum Sars, 1863. Now it is
supposed to be extinct at the original type locality.
On the other hand, the parts of Europe rich in numerous and deep lakes (both northern Europe
and the subalpine countries) are known to have a higher number of closely related and morphologically
very similar limnetic species of the genus Cyclops (e. g., C. scutifer Sars, 1863, C. abyssorum,
C. lacustris Sars, 1863, C. kolensis Lilljeborg, 1901, C. bohater Kozminski, 1933). Their species
identity and mutual relationships were quite complicated and unclear. Kozminski (1936) introduced
a detailed and elaborate morphometric analysis using about 30 statistically evaluated characters to
distinguish closely related species. Even using this approach the specific identity of some populations
remained uncertain (Einsle 1975). Their differentiation was enabled by two new techniques
*)
Present address: Department of Parasitology and Hydrobiology, Charles University, Vini#ná 7, CZ–128 44 Praha 2,
Czech Republic
This contribution is dedicated to Doc. RNDr. Jaroslav Hrbá!ek, DrSc., upon the occasion of the 80th anniversary
of his birthday ("2th May "92").
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introduced only a quarter of century ago. Beermann (1977) discovered the partial elimination of
chromatine from the chromosomes of early embryonic cells of a cleaving egg. The timing of this
process into the exact phase of the egg cleveage, specific for each of the very closely related
species, allowed Einsle (e.g., 1996a, b) to identify and describe some of the so far unrecognized
species. He also used another technique, i.e. the electrophoretic separation of allozyme (for copepods
used earlier by Boileau and Hebert 1988) to distinguish such very similar species. Combining
these two approaches with the analysis of previously neglected details of morphological structures
(Einsle 1985), he was better able to delimite the existing species and to describe some new
species (C. heberti Einsle, 1996, C. singularis Einsle, 1996, and C. stagnalis Einsle, 1996).
The aim of this work is to describe some detailed morphologic characters of several Bohemian
populations of the genus Cyclops which we have collected and to compare their species identity
with the recentmost descriptions.
MATERIAL AND METHODS
Plankton samples were collected with a coarse plankton net (0.38 mm mesh size) either by vertical hauls from
deep reservoirs and ponds or by pouring water gathered by 1-litre wide-mouth bottle from shallow pools through
the net. If possible, adult females were individually selected immediately from the fresh live material and then
preserved in 4% formalin; otherwise the whole sample was preserved in the same way. Preserved animals were
dissected in a drop of glycerin on a microscopic glass. The 4 th pair of swimming legs and its posterior face
ornamentation were examined. The ornamentation of the coxal segment of the basipodite of this pair of legs
consists of up to seven groups (Fig. 1, A to F) of little spines or setulae. They are arranged and placed in
characteristic positions (Einsle, 1996b) on the coxal segment. Moreover, the praecoxal plate connecting the
coxae (intercoxal coupler) may be either nake or covered with setulae and either simply even at the distal margin
or humped on each side near the inner lateral seta of the coxa (Fig. 1, H – the letter symbols follow Einsle, 1996b).
The humps may either be short and not exceeding the margin of the plate or they may project significantly
Fig. 1. Ornamentation pattern of the coxal segment of the 4 th pair of swimming legs in Cyclops O. F. Müller:
general scheme. The letters follow conventional labelling by Einsle (1985). A to F: groups of spines or setulae; H:
the humps of the intercoxal coupler.
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eyond the coupler margin. Other characters which we have recorded were the length of antennules as compared
with the length of the cephalothoracic body region, the shape of the genital segment of females, the position of
the lateral spine at the inner side of the distal segment of the 5 th legs, and the length of apical spines of the last
segment of endopodite of the 4 th pair of swimming legs.
We collected and examined material from various types of habitats ranging from deep reservoirs (Slapy
Reservoir on the Vltava River in Central Bohemia, $ímov Reservoir on the Malše River in South Bohemia) to
ephemeral puddles filled with rain water. Small permanent waters were represented by carp ponds and also by
permanent pools in the innundation areas of the Labe (Elbe) River nr. Velký Osek (north of Kolín, Central
Bohemia) and of the Upper Lužnice River (south of Suchdol nad Lužnicí). Besides these materials collected for the
purpose of this work, we also examined some of our older collections or samples kindly supplied by colleagues.
RESULTS
The specimens of cyclopoid copepods of the genus Cyclops we have examined can be distinguished
into five morphologically different types. The first of these types, which can be easily
recognized, is represented by C. insignis, the species differring from all the others within the
genus Cyclops by its 14-segmented antennules when adult. Adults of all the other species of this
genus have 17-segmented antennules, with a characteristic group of four short segments, which
are just as long as they are wide, in the middle of the antennule (segments 8 through 11).
One of the other four types we collected is undoubtedly a common species C. vicinus. The
typical characters of C. vicinus are the “wings”, i.e. widely extended lateral lobes of the 4 th thoracomer,
together with another constant character of this species, the spine formula 2.3.3.3. The last
character may occur in some of the other Cyclops species which otherwise have the spine formula
3.4.3.3. Therefore we describe some details of C. vicinus morphology in comparison to the details
of the remaining three types.
The ornamentation of the coxal segment of the basipodite of the 4 th pair of swimming legs and the
ornamentation of the intercoxal plate of this pair of legs (the coupler)
Cyclops vicinus. (Ornamentation pattern ABCDEF, Fig. 2). Group A consists of 20–35 mostly small
spines in a few rows. Some spines may be longer than the others. Group B: 4–7 short thick spines.
Group C: 7–12 long thick spines of equal length. Group D: 1 or 2 short thick spines. Group E:
numerous thin spines covering each other. Group F: short fine setae along the distal half of the
outer coxal edge. Intercoxal coupler: naked, with a pair of little humps.
Type I (Ornamentation pattern ACD, Fig. 2). Group A consists of 18–25 spines, the half of them
close to the coupler being finer, longer and set into slightly bent arch, the external half of them
being shorter, especially those in the middle of the row. Group C: 3–5 thicker and longer spines,
slightly bent outward from the intercoxal coupler. Group D: 1–3 short and thick spines. The humps
on the intercoxal coupler do not extend beyond the coupler margin, the coupler bears one row of
fine long setae in the middle of its length. Groups B, E, F: missing.
Type II (Ornamentation pattern ACDE, Fig. 2). Group A: 15–23 thick spines shorter than in type
I, often placed on the cuticular lobe. Group C: 6–8 thick spines, those in the middle being longer
than the external ones. Group D: 2–5 thick long spines. Group E: large number (up to more than 10)
of short or long thin spines partially covering each other. The intercoxal coupler with a pair of
humps extending well beyond its distal edge; the humps and the area between them are covered
with a row of very long fine setae. Groups B, F: missing.
Type III (Ornamentation pattern ABC, Fig. 2). Group A: a row of 25–30 short minute spines, with
the internal half of the row often in two parallel curved lines. Group B: from 3 up to 7 or 10 long weak
spines. Group C: long row of 14 to 22 short and strong spines of equal length. Intercoxal coupler:
without any extending hump, covered with numerous fine long setae in its distal part. Groups D, E,
F: missing.
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Position of the lateral spine of the 2 nd segment of the 5 th leg
Cyclops vicinus: inserted at the middle of the segment, not longer than the segment itself.
Type I : at the middle of the segment, as long as the segment.
Type II: at the middle of a very long and slender segment, shorter than the segment.
Type III: the spine shifted to the distal end of a shorter and wider segment.
Fig. 2. Ornamentation pattern of the coxal segment of the 4 th pair of swimming legs found in the examined
populations: CV = Cyclops vicinus Uljanin, 1857, 1 to 3 = Types I to III.
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The length of the outer apical spine of the 4 th endopodite
Cyclops vicinus: one third of the length of the inner apical spine.
Type I: 1/3 to 1/2 of the length of the inner apical spine.
Type II: 1/2 to 2/3 of the length of the inner apical spine.
Type III: 1/3 of the length of the inner apical spine.
Shape of the genital segment of female
Cyclops vicinus, Type I, Type II: Broadest at anterior part and gradually narrowing at posterior
part.
Type III: Anterior part wide, round-shaped, then abruptly narrowing into a cylindrical caudal part.
Habitats and localities
Cyclops insignis. Spring species in small permanent waters. Two permanent backwaters, Upper
Lužnice River nr. Dvory nad Lužnicí, many samples from 1992–1994, J. Hrbá#ek legit; the same
backwaters, 8.3. 1995, 21.3. 1995, Holý (1996); extensive flat shoal overgrown with willow and alder
shrub, Starý Vrbenský Pond nr. "eské Bud!jovice, 28.3. 1996, 11.4. 1996, 26.3. 1997, 9.4. 1997, M.
Lavická legit.
Cyclops vicinus. The most frequent species of the genus Cyclops in Bohemian ponds, reservoirs
and permanent backwaters. Almost permanent inhabitant of limnetic region with maximum density
in spring when its copepodid larvae emerge from the cocoons diapausing on the bottom. Examples
of localities: Slapy Reservoir, $ímov Reservoir (Brandl 1994); ponds nr. Blatná (Ko%ínek et al. 1987);
ponds Starý Vrbenský, Nový Vrbenský, Starý Haklovský, Nový Haklovský nr. "eské Bud!jovice,
year-round collections 1992, 1993, Z. Brandl legit; permanent backwater, Upper Lužnice River nr.
Lesní Chalupy, 27. 4. 1999, M. Lavická legit; permanent backwater, Labe (Elbe) River nr. Velký Osek,
22. 4. 1999, M. Lavická legit.
Populations of the Type I. Permanent backwater, Upper Lužnice River nr. Lesní Chalupy, 27. 4.
1999; permanent backwater, Labe (Elbe) River nr. Velký Osek, 22. 4. 1999; Starý Vrbenský Pond nr.
"eské Bud!jovice, 5. 5. 1999; $ímov Reservoir, 2. 6. 1999, all M. Lavická gblegit.
Populations of the Type II. Permanent backwater, Upper Lužnice River nr. Lesní Chalupy, 27. 4.
1999; $ímov Reservoir, 2. 6. 1999, all M. Lavická legit; Slapy Reservoir, material cultivated from the
female collected on 6. 3. 1999 by J. Hrbá#ek.
Populations of the Type III. Ephemeral pool filled only by rain water, with maximum depth of 0.07
m, grassland and shrubs at the abandoned millitary training area, NW edge of the town "eské
Bud!jovice, 27. 5. 1999, M. Lavická legit.
The identity of Types I to III
When the ornamentation patterns and the other details described above are compared with the
most recent descriptions of such characters given by Einsle (1996a, 1996b) we can state that:
Type I fits best to the description of these characters of either C. strenuus or C. stagnalis
Type II is closest to the description of the lowland populations (“C. divulsus” group) of C.
abyssorum or to the description of C. singularis
Type III is less ambiguous being closely similar to the description of C. furcifer with uniform
spine formula 3.4.3.3 (the rarer of the two possible formulas of this species).
DISCUSSION
All five Cyclops species mentioned in the previous paragraph may occur in Bohemian water
bodies. All these species are very similar in their morphology and differ in very small and unstable
details. Einsle (1996b) uses the point of insertion of the lateral spine on the 2 nd article of the 5 th legs
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(at the middle or nearer to the basis of the article). This character is very uneasy to quantify.
Another character separating C. strenuus from C. stagnalis is the length of the antennules. Acording
to Einsle (1996b), they reach just the end of the cephalothorax in C. stagnalis but up to the
middle of the 3 rd thoracomer in C. strenuus. This character was found to be very variable at least in
our populations by Lavická (2000). Similarly, the width of the 4 th thoracomer is used to differentiate
C. strenuus from C. abyssorum (e. g. Einsle 1975), the maximum width of this segment being in the
middle of its length in C. strenuus but in the slightly extended corners of the posterior part in C.
abyssorum. However, evaluation of this character is not always easy especially in preserved material.
Thus the morphological characters do not offer a reliable way to identify a given population up
to the species level. Another differentiation can be made using the stage of the egg cleavage in
which redundant heterochromatin is eliminated. This “chromatin diminution” takes place in a
definite cleavage stage specific for individual species which might differ also in the amount of
such eliminated material (from small particles up to huge amount of heterochromatin, Beermann,
1966, Einsle, 1996b). The timing of diminution into the 4 th or the 5 th cleavage stage also separates
the species in the two couples (C. strenuus + C. stagnalis and C. abyssorum + C. singularis) each
of which has the same coxal ornamentation in both species of a couple. The procedure to find out
this stage is relatively simple but hundreds to thousands individuals have to be examined until the
eggs are found in the relevant stage of their embryonic development. Therefore we lack this
information for any of the examined populations having the ornamentation of Types I, II or III.
However, the relevant five species differ from each other in their habitats. Three of them, C.
furcifer, C. stagnalis and C. singularis are known to dwell in the ephemeral habitats of temporary
ponds and pools which often completely dry off. On the other hand, both C. strenuus and C.
abyssorum (including its lowland group of populations known as the “C. divulsus-group”) are
inhabitants of permanent waters ranging from small permanent ponds to large and deep lakes,
although C. strenuus can be found in temporary pools, too. At least the populations of the Types
I and II we examined came from permanent waters which included even two large reservoirs. The
reservoir populations should therefore represent C. strenuus rather than C. stagnalis (Type I,
$ímov Reservoir) and C. abyssorum (Type II, Slapy Reservoir and $ímov Reservoir) rather than C.
singularis.
The question of species identity of the populations which belong to the type having the same
coxal ornamentation pattern was elucidated by parallel allozyme examination of the same populations
by Lavická (2000). She applied the electrophoretic separation of enzymes arginine phosphokinase
(APK) and glutamateoxaloacetate transferase (GOT) made on the cellulose-acetate
plates to all the populations of Type I, II and III we collected. With two loci of the GOT and three
of APK enzymes she was able to distinguish three genetic types within the examined material. (The
fourth one was represented by populations of Cyclops vicinus.)
The most important result is that each of these three genetic types corresponded to one of the
above described types of coxal ornamentation patterns. Thus, the populations having the same
type of coxal ornamentation represent also one genetically uniform type – and therefore they
belong to one species. When we take into account the properties of habitats of the Types I and II,
they are most probably identical with C. strenuus and C. abyssorum (lowland “C. divulsus-group”
of populations), respectively. The third Type III may represent an isolated local population of C.
furcifer with stabilized spine formula 3.4.3.3, which otherwise is the less common of the two
possible formulas of this species (Einsle 1996b).
The occurrence of other species of the genus Cyclops, e.g. C. stagnalis, C. singularis or even
C. heberti Einsle, 1996, and C. bohater Kozminski, 1933, is still possible in the Czech Republic. A
population of the mountain subalpine type of C. abyssorum (“C. praealpinus-group”) lived in the
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past in the Šumava Mountains’ lake “"erné jezero” (“C. bohemicus” of Šrámek-Hušek, 1937). In
more recent years it was found in another of the Šumava Mountains’ lakes, the “Prášilské jezero”
(e.g. Fott et al. 1994). Its relation to the lowland populations of C. abyssorum should be reexamined.
Numerous populations of C. abyssorum inhabit mountain lakes in neighbouring Austria
and Slovakia. Further reasearch of the genus Cyclops and its distribution in this country is still
needed and should be preferably made using all three possible approaches: detail examination of
morphology, enzyme electrophoresis and the analysis of chromatine diminution.
A c k n o w l e d g e m e n t s
We are honoured to be invited to participate in this special issue dedicated to Doc. Jaroslav Hrbá#ek on the
occassion of his 80 th birthday. The first author is especially happy to do so with a paper on the detail morphology
used to identify species – the approach which J. Hrbá#ek always promoted and taught his students. The study was
supported by the grant F0149/1999(G4) of the Ministry of Education, Youth, and Sports. J. Hrbá#ek kindly
supplied some samples of copepods. C. M. Steer greatly improved the English of this paper.
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Acta Soc. Zool. Bohem. 66: 169–175, 2002
ISSN 1211-376X
Impact of predation by cyclopoid copepods (Copepoda: Cyclopoida)
on zooplankton in a carp pond in Czech Republic
Zden!k BRANDL 1) & Miroslava PRAŽÁKOVÁ 2)
1)
Hydrobiological Institute, Academy of Sciences, Czech Republic and Faculty of Biological Sciences, University
of South Bohemia, Branišovská 31, CZ–370 05 "eské Bud!jovice, Czech Republic; e-mail:
zdbrandl@bf.jcu.cz
2)
Department of Parasitology and Hydrobiology, Charles University, Vini#ná 7,
CZ–128 44 Praha 2, Czech Republic
Received June 3, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. Predation by cyclopoid copepods (Cyclops vicinus Uljanin, 1857, Acanthocyclops robustus (G.
O. Sars, 1863), C. strenuus Fischer, 1851) was measured in field experiments in the pond Velký Pálenec
(SW Bohemia, Czech Republic) during two consecutive years. The cyclopoid copepods fed on all common
species of rotifers and also on the young larvae of copepods and in some periods also on Daphnia spp. and
other cladocerans. The predatory impact of cyclopoid copepod feeding may influence considerably the
populations of other zooplankters.
Predatory feeding, impact of predation, Cyclops vicinus, Acanthocyclops robustus, rotifers, Czech
Republic, Palaearctic region
INTRODUCTION
Cyclopoid copepods represent a group with a dual position in trophic webs of plankton communities.
Their early larval stages are herbivorous and feed on small algal cells. Later copepodite larvae
and adults may feed on plankton algae but they gradually change their feeding mode to predation
of plankton animals. The adults of large species even need a sufficient supply of animal food to be
able to produce eggs (Smyly 1970) although Whitehouse & Lewis (1973) obtained repeated production
of egg clutches from Cyclops abyssorum G. O. Sars, 1863, reared just on algal diet.
Growing interest in the food relationships within the plankton community have led to a series of
studies on both herbivorous and carnivorous feeding of various limnetic cyclopoid copepods (for
review see Brandl 1998a). The published data show a very wide range of animal species preyed
upon by cyclopoid copepods and significant impact of their predatory feeding on the other zooplankton
populations. The aim of this work is to present data about the food composition of the
cyclopoid species and about the impact of their predation on the other zooplankton in a carp pond.
We believe that even relatively old data can contribute to the contemporary discussion on the
extent of herbivorous and carnivorous feeding of cyclopoid copepods.
MATERIAL AND METHODS
During the period of intensive studies on pond ecosystems in carp ponds near Blatná, SW Bohemia, made by the
team of the Department of Parasitology and Hydrobiology, Faculty of Science, Charles University, detailed data
were obtained on zooplankton and its seasonal changes in several carp ponds. In 1980 and 1981 we performed
This contribution is dedicated to Doc. RNDr. Jaroslav Hrbá!ek, DrSc., upon the occasion of the 80th anniversary
of his birthday ("2th May "92").
169

Tab. 1. Predation rates of the cyclopoid copepods on the rotifer, cladoceran and copepod prey in the pond Velký
Pálenec, 1980–1981
Prey species No. of exp. series prey density, predation rate,
prey×l –1 , range prey×predator –1 ×day –1 ,
median and (range)
Asplanchna sp. 6 1–1224 0.33 (0.05–0.80)
Brachionus angularis 2 6, 16 0.08 (0.02, 0.14)
Brachionus rubens 2 164, 409 2.60 (2.57, 2.63)
Conochilus unicornis 3 1–50 0.39 (0.10–0.50)
Keratella cochlearis 10 1–1485 0.86 (0.05–4.79)
Keratella quadrata 9 1–89 0.07 (0.02–0.64)
Polyarthra dolichoptera 15 2–261 0.40 (0.02–15.4)
Pompholyx sulcata 2 133, 390 0.08 (0.04, 0.13)
Synchaeta sp. 2 104, 245 2.76 (2.45, 3.06)
Daphnia galeata 6 3–283 0.35 (0.02–1.52)
Diaphanosoma brachyurum 1 24 0.03
Bosmina longirostris 2 1, 34 0.24 (0.03–0.46)
Copepoda, naupliar larvae 15 27–405 0.38 (0.06–2.32)
Cyclopidae, copepodite larvae 12 9–198 0.12 (0.05–0.70)
Eudiaptomus, copepodite larvae 3 1–40 0.02 (0.01–0.05)
paralell measurements of predatory feeding of carnivorous cyclopoid copepods in the pond Velký Pálenec (for
the description and detailed characteristics of this pond, see e.g., Hrbá#ek 1966, Fott et al. 1974 or Ko%ínek et al.
1987). This 31-ha eutrophic pond with an average depth of 1.44 m is used for carp cultivation, usually in a twoyear
cycle with complete draining at the end of each cycle.
The experimental procedure used to measure the predation of cyclopoid copepods was essentially the same as
described by Brandl & Fernando (1978). The natural pond zooplankton was collected from a known volume of
water and concentrated. The size fraction containing predatory stages of cyclopoid copepods was separated from
the prey zooplankton by screening through metal sieves of convenient mesh size. The “prey zooplankton” (both
smaller and larger than the predators) was then thoroughly mixed and evenly distributed into experimental and
control replicates (usually two of each). The size fraction containing the predators was added only to the
experimental series of replicates. The replicates placed in 5-litre plastic bottles were re-diluted with filtered pond
water to the original pond zooplankton concentration. The bottles were hung in a horizontal position in the pond
for 24 hours. Then their content was concentrated using 0.04 mm sieve, preserved in 4% formalin and counted
under the microscope. The difference between the experimental and control series after 24 hours was then
attributed to predation by the cyclopoid copepods present in the experimental replicates. When egg-bearing
females of either cyclopoid copepods or other plankton animals were present in the “predator” size fraction, a
correction was applied in the experimental series for the number of newborn naupliar larvae or cladoceran
neonates. The correction was calculated from the number of eggs of each species per replicate, counted from two
other replicates preserved in the beginning of the experiments, and from the known temperature-related rate of
embryonic development (e. g., Bottrell et al. 1976, Herzig 1983).
The qualitative and quantitative data on species composition of zooplankton in the pond were derived from
representative samples (Hrbá#ek 1966) taken across the pond using a 30-litre Schindler sampler and resulting
usually in a sample from 450 l of water (data of the second author).
RESULTS
The cyclopoid copepods and their share in the plankton community
Three species of cyclopoid copepods represented predators in the pond community. The most
constant component was Cyclops vicinus Uljanin, 1875. The adults were present year-round in
densities usually between 1 and 10 per litre, rarely 3× more (June 1980). The later larval stages were
more numerous especially in spring and fall months when the density of copepodite larvae reached
170

almost 100. l –1 . The second important, and temporarily even more numerous species, was Acanthocyclops
robustus (G. O. Sars, 1863) with occurrence seasonally limited to the second half of the
vegetation season. In 1980, the adults appeared in late June and reached high densities in September
(28×l –1 ). In 1981, the period was the same: first adult males appeared in June, maximum density
of 28×l –1 was recorded in early September. The density of the later (IV+V) copepodite stages was
above 80×l –1 in September with more than 100×l –1 of younger copepodite larvae. This species
completely dissappeared from the community in late October.
The third species occurring in much lower densities (usually less than 1×l –1 and rarely between
1 and 5 per litre) was Cyclops strenuus Fischer, 1851. The adults of this species were present from
June to October in 1980 and from early spring till August in 1981. Other species of cyclopoid
copepods were recorded only occassionally in very low densities as Mesocyclops leuckarti (Claus,
1857) or Eucyclops serrulatus (Fischer, 1851) .
The most numerous representatives of cyclopoid copepods were always their naupliar larvae,
reaching densities of hundreds per litre (the maximum value 405×l –1 in May, 1980). However, due to
their herbivorous feeding mode they rather should be considered one of the possible prey species.
The herbivorous species available as a prey to cyclopoid copepods
The main component of the pond zooplankton community were cladocerans, represented mostly
by Daphnia galeata Sars, 1864. This is a relatively large species, namely in carp ponds with low
fish predation of large crustaceans, but its juveniles may become the prey of cyclopoid copepods.
Except for early spring and late autumn, the density of this species was always between 30 and 300
per litre, with more juveniles than adults. The other species of the same genus, the larger Daphnia
pulicaria Forbes, 1893, was rare in 1980 and more common in 1981 (up to 48×l –1 in May). Other
cladocerans were less important and occurred in low densities (Ceriodaphnia Dana, 1853) or with
seasonally limited period of higher densities (Bosmina longirostris (O. F. Müller, 1785): 16–30×l –1
Fig. l. Predatory impact of cyclopoid copepods on cladocerans, copepods and rotifers in the pond Velký Pálenec,
1980: density per litre and the part eaten daily by cyclopoid copepods.
171

Tab. 2. Predation by cyclopoid copepods on Daphnia galeata juveniles (females not yet reproducing) compared to
the reproduction of the Daphnia population, data for the 1980 cases of significant predation on Daphnia
Date June 23 September 3 October 1
temperature, °C 18.4 17.3 13.6
Cyclopidae, predatory stages, density per litre 25.8 52.5 116.9
Daphnia galeata, juveniles, density per litre 205.4 122.2 87.3
newborn Daphnia galeata, % per day 5.9 11.7 14.2
D. galeata eaten by copepods, % per day 2.7 9.4 3.5
in October 1981, Diaphanosoma brachyurum (Liévin, 1848): 24×l –1 in August, 1980 and 8×l –1 in
August, 1981). Another possible crustacean prey, the copepodite larvae of two herbivorous calanoid
copepods Eudiaptomus gracilis (G. O. Sars, 1863) and E. vulgaris (Schmeil, 1898) were
almost always present but they reached higher densities only in June (40×l –1 in 1980 and 13×l –1 in 1981).
The most numerous available prey were rotifers. We recorded 12 common species with up to 9
of them present at the same time. However, the occurrence of individual species was highly
seasonal and their maximum densities were usually limited to periods of few weeks. Then they
reached densities of up to hundreds per litre. In some species, these densities persisted for longer
periods – Keratella cochlearis (Gosse, 1851): 300–790×l –1 in August and September, 1980, and 110–
1485×l –1 in the same period of 1981. For the ranges of densities of some species, see Table 1.
Prey selection and predation rates of cyclopoid copepods
The list of species for which we recorded significant difference between the experimental and the
control treatments contained all possible prey species which were at least sometimes present in
higher densities. They included nine species of rotifers, three cladocerans, naupliar and the small
copepodite larvae of copepods (see Table 1). The most important component of the prey of
cyclopoid copepods were rotifers. The per capita predation rates by cyclopoid copepods of some
species of rotifers reached several specimens per day during the periods of high rotifer densities
(Polyarthra dolichoptera Idelson, 1925, Keratella cochlearis) but were low when a given rotifer
species was rare. Consistently high values were recorded for species with limited periods of
occurrence (Brachionus rubens Ehrenberg, 1838, Synchaeta sp.).
The rates of feeding on cladocerans were lower, especially when evaluated in relation to the
high possible gain of food in contrast to small rotifers. In most cases, the feeding rate on Daphnia
galeata was insignificant or not measurable. We recorded only six cases of significant values.
Those which we could compare with direct measurement of reproduction rate (1980) are given in
Table 2. They account for about a half of the production of Daphnia neonates per day.
The last, but not least, component of cyclopoid prey were younger copepodite and especially
naupliar larvae. The significant values of predation rates on one or the other of the larval stages
were recorded in all but one cases of our measurements.
The impact of predation on the whole zooplankton community
The summarized values of the daily eaten prey in terms of prey specimens compared to the overall
densities for the categories of cladocerans, copepods and rotifers are given in Figures 1 and 2. Since
the size and biomass are very different for individual prey species these figures give just a rough idea
of the impact of predation. Neverheless it is obvious that the predation is oriented mostly towards
the rotifer component of the zooplankton community especially during periods of the high rotifer
density. The relatively high impact of copepod predation on their own category is the result of
feeding on naupliar larvae rather than on the larger copepodite larvae. A high impact of the copepod
predation on newborn Daphnia galeata was found as well, but only occasionally (Tab. 2).
172

DISCUSSION
Cyclopoid copepods inhabiting the pond Velký Pálenec (i.e. mainly Cyclops vicinus and Acanthocyclops
robustus) in both years caused measurable differences in prey densities between the
control and the experimental series of containers for at least part of the possible prey species
present in the pond. Whenever some species of rotifers were present in high densities they
became the preferred type of prey and per capita daily predation rates grew up to several specimens
of species like Polyarthra dolichoptera and Keratella cochlearis. Also the species which
occurred for just short periods, but in high numbers, were intensively preyed upon. The preference
of cyclopoid copepods for rotifer prey agrees with the data of numerous authors (from more
recent e.g. Diéguez & Gilbert 2002 or Plassman et al. 1997, for review of earlier data see Brandl
1998a). Plassman et al. (1997) found that Cyclops vicinus preferred Synchaeta spp. to species of
Keratella Bory de St. Vincent, 1822, and Polyarthra Ehrenberg, 1834, in Lake Constance. This
agrees with our data on high predation of Synchaeta Ehrenberg, 1832, in early spring when Cyclops
vicinus and C. strenuus were present in the pond whereas Acanthocyclops was not. Brandl
(1998b) found per capita daily predation rates by Cyclops vicinus in two Czech reservoirs up to 4.2
for preying on Polyarthra dolichoptera and up to 3.4 for Keratella cochlearis. However, the
rotifer population densities were also lower in reservoirs than in the pond Velký Pálenec. Moreover,
high copepod predation of rotifers of the other two most common genera, Polyarthra and
Keratella, can be at least partially attributed to the third cyclopoid predator, Acanthocyclops
robustus. This species is probably also responsible for predation of juvenile specimens of Daphnia
galeata since we found the significant values of Daphnia consumption right in the periods of
high densities of adult and later larval stages of Acanthocyclops occurring at the same time as high
density of Daphnia. The increase of the Acanthocyclops robustus predation of Daphnia sp. was
also reported by Roche (1990). Similarly Caramujo & Boavida (2000) who studied the tail spine
Fig. 2. Predatory impact of cyclopoid copepods on cladocerans, copepods and rotifers in the pond Velký Pálenec,
1981: density per litre and the part eaten daily by cyclopoid copepods.
173

elongation in Daphnia as a defence mechanism against Acanthocyclops predation found significantly
high values of predation rates by Acanthocyclops robustus on the youngest instars of
Daphnia hyalina x galeata.
An even more important component of cyclopoid food than Daphnia are the naupliar and
younger copepodite larvae of both cyclopoid and calanoid copepods. Cannibalistic predation is a
common feeding mode in cyclopoid copepods (see e.g. Gabriel 1985). Brandl & Fernando (1978,
1981) reported the predation of naupliar larvae by Cyclops vicinus in pond and reservoir populations.
The extent of the predatory impact of cyclopoid copepods on the other zooplankton in the pond
Velký Pálenec varied from very low or zero values for some of the three groups of plankton animals
(usually cladocerans or rarely copepods) to a relatively high percentage, between 10 and 20
percent for the rotifer populations. Occasional high values for copepods occurred in the late
spring when the cyclopoid copepods were slowly receding from the plankton community and
entering their diapause phase on the bottom.
During the past decade, algae have been more closely examined as an important food source for
adult cyclopoid copepods and their later copepodite stages, and the share of the herbivorous and
the carnivorous feeding mode have been discussed (e.g. Adrian 1991, Hansen & Jeppesen 1992,
Santer 1993, Santer & Bosch 1994, Hansen 1996). There is no doubt that especially Cyclops
vicinus feeds opportunistically on large algal cells, coenobia or colonies since the algal material
can often be seen in the guts of living animals. However, the predation by cyclopoid copepods on
the other zooplankton species is an equally important mode of feeding and its impact on the other
species may often be essential for the further population development of the prey species.
A c k n o w l e d g e m e n t s
We are especially happy to contribute to this special issue dedicated to Jaroslav Hrbá#ek on the occassion of his
80 th birthday. The first author thanks J. Hrbá#ek for many hours of invaluable discussions on various productivity
problems of water ecosystems and for the orientation to prospective topics in this research. The evaluation of
data was supported by the grant No. K 6005114 GA AV "R. C. M. Steer greatly improved the English of this paper.
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175

Acta Soc. Zool. Bohem. 66: 176, 2002
ISSN 1211-376X
176
BOOK REVIEW
MEHLHORN H. (ed.): Encyclopedic Reference of Parasitology. Second edition. Springer-Verlag: Berlin-
Heidelberg-New York, etc., 2001. Format 190×268 mm. First volume: Biology, Structure, Function. XXI+670
pages. Hardcover. Price sFr 326.00. ISBN 3-540-66819-5. Second volume: Diseases, Treatment, Therapy.
Hardcover. Price sFr 257.00. XXI+678 pages. ISBN 3-540-66829-2
The editor is professor at the Institute of Zoomorphology, Cell Biology and Parasitology of the Heinrich Heine
University in Düsseldorf (Germany). The project of this publication involved 38 acknowledged authorities from
Europe, USA and Brasil. As declared in the preface by the editor, this second edition intends to give a comprehensive
overview of the facts and trends in veterinarian and human parasitology. It is to point up that this is an
encyclopaedia, not a specialized textbook. The lay-out of the book presents an encyclopaedic arrangement of
2157 entries (key-words) in alphabetical order along the subject index being included in the flow of the text. For
a book review, entries may be assembled into several thematic blocks. Particular entries embrace the extent from
a simple reference to another page or a space-limited paragraph – up to an essay length of >10 textual pages.
First volume comprises 1592 entries concerning biological/zoological aspects of parasitology. Entries that
describe systematic assemblages of parasites consist of sections illuminating classification, distribution, morphology,
genetics, reproduction, life cycles, feeding behaviour and transmission. Other thematic block directs attention
on morphology and structural parts of parasitic organisms placing emphasis upon the fine structure. Unicellular
protozoans are represented by entries describing external and internal organellae. Entries related to metazoan
parasites, helminths and arthropods, encompass terms relevant to external and internal morphological
structures and organ systems. Various physiological and metabolic functions and general outlines of evolution,
ecology and other sciences related to parasitism are introduced by terms such as behaviour, coevolution, ecosystems,
energy metabolism, encystation, environmental conditions, favorisation, host-parasite interface, local
adaptation, nutrition of parasites, etc. Entries dedicated to biochemistry, molecular biology and genetics are
concerned with terms such as aminoacids, deoxynucleotides, hybridization, extended phenotype, genome projects,
proteases, glycosylphosphatidylinositols, nucleic acids, lipids, RNA editing, etc. Entries on other fields of interest
and importance ensure coverage of arboviruses, host finding, immune diagnostic methods, phylogenetic relationships,
serology, transmitted pathogens, and vector biology. Second volume embraces 565 entries covering
topics related to pathogenical effects of parasitism in humans and animals, clinical aspects, diagnostic procedures,
therapeutical regimens and schedules. Thematic block based on parasitic diseases is focused on general information,
pathology, immune responses, evasion mechanisms, main clinical symptoms, chemotherapy, diagnosis,
prophylaxis and vaccination, and control measurements. A special credit deserves the entry that gives insights
into miscellaneous aspects of immunity. Moreover, particular sections surveying immune responses are situated
in each entry examining a particular parasitic disease. These sections delineate in detail innate defense mechanisms,
B cells and antibodies, T cells and cytokines, control of parasites by activated macrophages, immunopathology,
autoimmunity, evasion mechanisms and vaccination. In other thematic block explored are general
pathological terms such as respiratory, genital and nervous system diseases in animals, eye parasites, pathology,
parasitaemia, parasite load, disease control, etc. Entries devoted to chemotherapy are introduced partly by terms
related to parasitic agents, partly by terms characterizing their biological effect. Entries discussing other fields of
interest are concerned with epidemiology, host reactions, opportunistic agents, serology, vector transmitted
diseases and zoonoses.
Both volumes are extensively illustrated by 497 (371+126) figures composed of schematic line drawings and
photographs. In volume 1 featured are individual parasitic organisms – protozoans, helminths and arthropods in
adult forms and developmental stages, enternal and internal structural parts, organs and tissues, structures of
biomolecules, diverse biosynthetic and metabolic pathways, schematic representations of cell division, modes of
reproduction and miscellaneous biological phenomena, genealogy of different chromosomes, phylogenetic trees
and evolutionary schemes. Numerous original schematic life cycle drawings are of particular didactic value.
Stereoscans depict most instructive aspects of parasitic arthropods. In volume 2 featured are pathological
abnormalities and lesions, clinical conditions, fundoscopic findings, distribution maps, transmission cycles, structural
formulae and models of action of drugs. 183 (76+107) detailed tabular summaries give excelent overwiews
of data presented in the textual part.
Compiled by a broad spectrum of internationally respected experts, this remarkable opus provides an authoritative
and most exhaustive encyclopaedia embracing all aspects of parasitology. Without doupt, this publication
will become an indispensable necessity for libraries of professionals, universities and research institutes.
Jind%ich Jíra

Acta Soc. Zool. Bohem. 66: 177–188, 2002
ISSN 1211-376X
Taxonomy and conservation problems of the native salmonids
(Pisces: Salmonidae) in the Danube river system: a review
Juraj HOL"ÍK
Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, SK–842 06 Bratislava, Slovakia;
e-mail: uzaeholc@savba.savba.sk
Received May "5, 2002; accepted Septeber 3, 2002
Published November 4, 2002
Abstract. Five species of salmonids (Salmonidae sensu Reshetnikov, 1980) are recognized in the Danube
River basin. The anadromous Salmo labrax Pallas, 1811, known from the lower course of the Danube proper
is now very rare. Its lake derivative Salmo labrax (morph lacustris) still lives in some Austrian lakes. Its
present status was described as disastrous, as it disappeared in some lakes and in other ones it was replaced by
intentionally introduced lake trout of different origin. Another species of lake trout, the deepwater Salmo
schiefermuelleri Bloch, 1784, endemic to some Austrian lakes seems to be completely extinct. Its proper
taxonomic status is not clear yet. It is suggested that this taxon represented those specimens of the adult
Salmo labrax lake form, which left out spawning for one or two years. The upper and middle Danube and its
mountain and foothill tributaries are inhabited by the brook trout Salmo labrax (morph fario). Genus
Salvelinus is represented by Salvelinus umbla (Linnaeus, 1758), and the deepwater species Salvelinus profundus
Schillinger, 1901. From three forms of the former, the fast growing and predatory Wildfangsaibling and the
zooplankton feeder Normalsaibling became probably extinct and only the deepwater Tiefseesaibling still
survives in one lake. Also Salvelinus profundus, known from two Austrian lakes, is now probably extinct.
Conservation status of the Danubian huchen Hucho hucho (Linnaeus, 1758) may be evaluated as critically
endangered. Its range reduction recorded since 1945 seems to be halted in some countries, but in other it is
decreasing and its populations are maintained mostly by extensive stocking.
Taxonomy, conservation, Pisces, Salmonidae, Salmo labrax, Salmo schiefermuelleri, Salvelinus
salvelinus, Salvelinus profundus, Hucho hucho, Danube river basin, Palaearctic region
INTRODUCTION
The purpose of this paper is to initiate and/or promote research on the status of native salmonids
inhabiting the Danube River basin, including their ecology and conservation and in most of them
even their taxonomy and nomenclature. Danube basin is noted for both the highest total number
and the highest ratio of endemic fish species in Europe (Berg 1932, Lindberg 1972). The Danube
basin is also known as the Quaternary refuge of its freshwater fauna which, after the retreat of the
ice sheat, expanded into other European river basins (Lindberg 1972). It is therefore astonishing,
that relatively little is known about most Danubian native salmonids. This criticism deals both with
their taxonomy and ecology, including their population density and in some respect even the fish
catch statistics. It seems that most European ichthyologists were, and still are, interested in fishes
from remote, exotic countries and not in those inhabiting rivers within reach of their hands. This is
reason why the following data, although they were excerpted from all available information, give
only an indistinct and in some respect confused picture on the present status of given species. In
addition it seems, that most now belong to the endangered, and some species even among the
extinct freshwater fishes of our Old continent.
To Dr. Jaroslav Hrbá!ek, Scholar, Teacher and Personage at the occassion of his 80th birthday.
177

Genus Salmo Linnaeus, 1758
Until now most authors suppose that European rivers are inhabited, besides forms of unknown
systematic status, by three species of the genus Salmo, i.e. the Atlantic salmon Salmo salar
Linnaeus, 1758 and the brown trout Salmo trutta Linnaeus, 1758, and by the marble trout Salmo
marmoratus Cuvier, 1817. The brown trout forms three ecological forms, the sea trout –S.trutta
(morph trutta), the brook trout – S. trutta (morph fario), and the lake trout – Salmo trutta (morph
lacustris). This nomenclature implies close relationships of the migratory, stream resident and the
lake resident forms respectively. However, it has been documented, that all three ecological forms
inhabiting one river basin represent one gene pool and/or they are much more closely related each
other, than to any of those from another river basin (Ferguson et al. 1995 and references therein).
Moreover, recent data coming from population genetic analyses of various trout populations
using both nuclear-gene (biochemical), nested clade and mtDNA markers throughout all over the
Europe (e.g. Bernatchez et al. 1992, Riffel et al. 1995, Apostolidis et al. 1996 a, b, Bernatchez &
Osinov 1995, Giuffra et al. 1995, Osinov & Bernatchez 1996, Bernatchez 2001) suggest that actual
biological diversity of the brown trout might be far greater than it is recognised by the current
taxonomy with its nomenclatorial limits (Kottelat 1997). This situation resembles that found recently
in the another salmonid genus Oncorhynchus Suckley, 1861(Behnke 1992).
In other words, recent data coming from using the karyotype pattern (Phillips & Ráb 1997) and
the mtDNA and biochemical analysis of various stocks (Kottelat 1996, and literature herein) indicate
the more complex species composition of European trouts. This postulate is now verified by
the thorough analysis of the allozymes and mtDNA of the “brown trout” stocks from many localities
covering various European and also Central Asian river basins, made by Osinov & Bernatchez
(1996). Within five phylogenetic groups distinguished earlier (Bernatchez et al.1992, Giuffra et
al.1994, Bernatchez & Osinov 1995, Bernatchez 1995, 2001) there are two large ones: the “Atlantic”
group and the “Danubian” group, the last one involving “brown trout” stocks inhabiting streams
and lakes belonging to the basins of the Black, Caspian and Aral seas. The migratory trouts of the
Black Sea and the lower Danube have been described as a distinct species Salmo labrax Pallas,
1814, later on considered to be subspecies of Salmo trutta by Berg (1916). Afterwards Balon (1968,
1969) refused even the subspecific status of the Black Sea trout and the migratory trout from its
watershed he considered to be a morph – Salmo trutta m. labrax – only. However, observations by
Dorofeeva (1967, 1977, 1985), Osinov (1984, 1989), Bernatchez & Osinov (1995), Largiader & Scholl
(1995), Togan et al.(1995) and Osinov & Bernatchez (1996) revealed some osteological, karyological,
biochemical and molecular differences among particular subspecies of Salmo trutta, suggesting
to attribute them specific status. This is particularly true for the Black Sea trout populations
and Kottelat (1997) tentatively recognizes it as a distinct species, in spite that the recent list of the
salmonids inhabiting the territory of Russia, only Salmo salar and Salmo trutta are still recognised,
the latter with four subspecies – S. t. caspius Kessler, 1870, S. t. ciscaucasicus Dorofeeva,
1967, S. t. labrax Pallas, 1814, and S. t. ezenami Berg, 1948 (Dorofeeva & Savvaitova 1998) In this
paper I follow the opinion of Kottelat (l. c.) and consider the Black Sea trout to be a valid species.
Salmo labrax Pallas, 1814
The main anatomical characters distinguishing this species from its closest relative Atlantic trout
Salmo trutta Linnaeus, 1758, occurring in the watershed of the Atlantic Ocean is the higher
number of gill rakers, the high caudal peduncle depth (Berg 1948, Pavlov et al. 1994) and the
structure and dentition of some skull bones (Dorofeeva 1967). Also, as revealed just recently, there
is a set of distinct genetic markers suggesting that it represents one gene pool (Ráb pers. comm.).
178

The Black Sea trout is a migratory species, living as adult in the littoral of the Black Sea and
ascending rivers for spawning. The following details on its ecology and life history are taken from
the papers by Berg (1948), B&n&rescu (1964), Svetovidov (1964), Elanidze et al. (1970), Marinov
(1978), Pavlov (1980), Elanidze (1983), and Pavlov et al. (1994). Spawning migration begins in
February, it peaks in April and May and terminates in July. The spawning proper is in October,
peaks in November and lasted till the end of January. Juveniles stay 2–4 years in the brooks where
they hatched and only then move downstream to their feeding grounds in the sea. Characteristic
feature of this species is strong predominance of females: in some rivers the migratory and resident
trouts form one stock, where migrants are represented by females which mate with males belonging
to the resident brook form (m. fario). In this case part of hatched juveniles does not transform
to smolts and stay in a stream. Smoltification is determined by sex, as only females leave the river
while males remain there. Sexual maturation sets in at the age of 3–5 years and 350–900 mm of the
total length (Tl). The diet of juveniles is composed of invertebrates only, but smolts start to eat
also fish. The major food of adults in the sea is fish, mostly anchovy [Thryssa (syn. Engraulis)
encrasicholus (Linnaeus, 1758)]. The main distribution of this trout are eastern shores of the Black
Sea and the Azov Sea but it is known also from the shores of Crimea, Ukraine, Romania, Bulgaria
and Turkey. It ascended the rivers of Crimea and Caucasus and also the rivers Dnieper (formerly up
to Kremenchug, some 500 km from the river mouth), Don (up to Pavlovsk) and Kuban’ with its
tributary Laba (some 300 km upstream). In the Danube River it was known from Silistra and Orechovo,
350 and 700 km upstream from its delta, respectively, and in the delta lakes Razelm and Yalpukh.
In comparison with eastern stock the western one was always less abundant and only single
specimens weighing mostly 3–4 kg were taken. Maximum recorded size of this species was 1100
mm and 24 kg. At present the population of this species generally dropped dramatically and only
few specimens are sometimes taken. In the main fishing grounds along the Georgian shores, where
formerly 9 tons were taken annually, the fishery for this species had to be completely stopped.
However, I must say that the exact status of the trout in the lower Danube is not known, in spite
that the local fishermen known this trout perfectly. Only few and generally small-sized specimens
are occasionally caught there (B&n&rescu 1964, Pavlov 1980) and Balon (1968, 1969) considered
them to be resident, non-migratory trouts washed downstream from the Danube river basin. According
to Suciu (pers.comm.) the Romanian trout catch in the Danube delta is about 100 individuals
annually and their length and weight varies from 400 to 520 mm Tl and 100–1800 g, respectively.
Trouts found in the middle Danube have a similar colouration as the sea trouts and lake trouts,
i.e. only black spots are scattered on their dorsum and flanks (Hol#ík 1969). The status of the sea
trout from the Black Sea was evaluated by Lelek (1987) as endangered and by Pavlov et al. (1994)
as vulnerable. However, the situation in the Danube basin is more critical, so the actual conservation
status of this species at the western range of its area may be evaluated as critically endangered.
The reasons of its decline are not properly known but it is certainly the combination of
overfishing and the deteriorated environmental conditions, including pollution, construction works,
and damming of streams where this species spawns. A list of conservation measures proposed by
Lelek (1987) includes environmental protection, fishing restrictions, establishing of hatcheries and
also intensive research focused on the population ecology of this species. Russian authors (Pavlov
et al.1994) recommend also the the exclusive use of the local brown trout stock in hatcheries.
Most Alpine and pre-Alpine lakes in Austria are inhabited by the lake form – the lake trout
Salmo labrax (m. lacustris). This form resembles the sea trout in form and also colouration,
however it is much larger as it grows up to 31 kg. Its ecology is similar to that of both the sea trout
and brown trout, but details are not well known. Sexual maturity is reached and first spawning
performed only when fish attains 500 mm in the Tl, i.e. as 4 (3+) and 5 (4+) years old males and
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females, respectively. In late autumn the lake trout migrates into lake tributaries, sometimes even
into the outlets to spawn and afterwards it returns to the lake. In rare occasions the spawning was
also observed in the lake itself, but usually close to the mouth of tributaries. In some localities the
spawning with the resident brook trout was observed, but hybrids are said to display lower growth
rate and they attain much smaller size than the lake trout (Hochleithner 1989). Two forms of the lake
trout were recognized in particular lakes. They differs in the size, colour and the diet as well. Small,
light-coloured and sexually immature fish called Schwebforelle, Silberforelle, Silberlachs or Jugendform
inhabits upper layers of the water column, dwells close to shores and feeds on the
airborn insects or small fishes. Large, dark and adult specimens called Grundforelle inhabit pelagial
and are confined to depths up to 40 m below surface. They are typical predators preying on
fish. Present conservation status of the lake trout in Austrian lakes was described as disastrous by
Hochleithner (1989). This author made an attempt to assess its actual situation and tried to check
populations inhabiting particular Austrian lakes. However, his data are based on reports of various
persons, they lack quantitative figures dealing with population abundance and fish catch statistics.
Due to this the following review is only a very rough and somewhat subjective simplification.
From 14 lakes of the Danube basin the lake trout catches and recorded individual maximum or mean
weights decreased in six (43%), seems to remain the same in four (28%) and improved in one (7%)
lake. In three lakes, however, (Fuschlsee, Traunsee and Hallstättersee; 21%), the lake trout is now
extinct. Individual weight of the lake trout catches from these lakes, recorded mostly till the first
half of this century, attained up to 31 kg. As noted by Einsele (1959) 20 kg specimens were not rare.
At present, however, the catches of individuals over 10 kg are very rare and the mean weight
recorded during last years varied mostly from 6–8 kg. Both Hochleithner (l. c.) and Jagdsch (pers.
comm.) claimed that reason of the decline of the lake trout is a combination of overfishing and
environmental changes, including alteration or damage of the spawning grounds by construction
works, and weirs and dams which have cut access to spawning grounds, and pollution and
eutrofication which kill the fertilised eggs and hatching alevins, as well as the food resources for
juveniles. Both Hochleithner and Jagdsch also stressed the wrong management which contributed
to the decreasing trend of the lake trout stock in the Alpine and pre-Alpine lakes. With the aim
to improve the population density of this species, foreign, (mainly Danish strains) of the sea trout
(i.e. Salmo trutta [m. trutta]) reported to be the lake trout, were imported to Austria and stocked
into the lakes. The imported trouts are accused to hybridise with the true, native, lake trouts. Their
hybrids are noted, for both their lower growth rate and individual weight, but no data were introduced
to support this assertion.
This, however, does not indicate the introgression of the imported trout of the Atlantic origin.
Largiader & Scholl (1995) found that indigeneous genetic variation in the brown trout populations
is still predominant in the Danube basin in Switzerland despite stocking. The low impact of foreign
brown trout strains was also found also elsewhere as documented by Arias et al. (1995) in Spain,
Beaudou et al. (1994, 1995), Poteaux et al. (1998) in France and by Ráb (pers. comm.) in former
Czechoslovakia. The same seems to be valid for the genus Salvelinus (Kummer & Jungwirth 1999).
According to Jagdsch (pers. comm.) the present managing practice in Austria improved. Stocking
activity is now based on the native lake trout progeny, and the Grundlsee and the Attersee
lakes provide the original brood stocks for hatcheries. According to the Red List of endangered
fishes in Austria the situation seemed to have improved during past decade. While in 1983 the lake
trout was listed among endangered fish species (Hacker 1983) in 1989 it had moved to the vulnerable
category (Herzig-Straschil 1994). In Switzerland, the lake trout now occurs in only one lake of
6 lakes of the Danube basin populated by fish and the species is evaluated as critically endangered
(Pedroli et al.1991).
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There is one more species of the genus Salmo described from the Alpian lakes. It is Salmo
schiefermuelleri Bloch, 1784, known under vernacular name Mayforelle, sometimes also Silberforelle.
This is a mysterious fish, whose vernacular name is derived from the observation that it
might be seen close to the lake surface only in spring, while during most of the year it dwells in
greater depths. This form was known to inhabit the lakes Attersee, Traunsee and Fuschlersee
(Heckel & Kner 1854). It was reported to grow to the same large size as the deep-water lake trout or
Grundforelle, i.e. 20–30 kg. According to Heckel (1851) and Heckel & Kner (1858), the difference
between the deep-water Grundforelle and the Mayforelle is in the head shape, coloration, more
deciduous scales, white eggs which would be of the same size as the millet seeds (while in the
Grundforelle they are of the size of the pea seeds) and hardships to keep it alive. Systematic status
of this trout is not solved yet. Haempel (1930) thought that the Mayforelle is only a sterile form of
the lake trout. Berg (1932, 1948) considered it as a synonym of the Danubian basin lake trout –
Salmo trutta labrax according to his nomenclature – while Balon (1968) supposed it to be the
infraspecies i.e. ecological form of the species Salmo trutta. According to Kottelat (1997) Salmo
schieffermuelleri is a valid species. With respect to data by Heckel & Kner (1858) and Haempel
(1930) dealing with small eggs and/or its reproductive sterility, I suppose that the Mayforelle are
those specimens of the adult lake trout (Grundforelle) which left out spawning for one or two
years. Such a situation is known in various species of fishes including coregonids and salmonids
occurring in northern regions or in deep lakes (Kennedy 1953, Grainger 1953, Gullestad 1973,
Hol#ík et al. 1988). The exact taxonomy of this trout remains to be solved. However, the original
stock seems to be extinct (Balon 1968, Kottelat 1997).
Genus Salvelinus Richardson, 1836
The systematics and nomenclature of the charrs, genus Salvelinus are even more obscure than
those of the trouts. As pointed out by Kottelat (1997), here the main problem is the existence of a
number of various stocks, which are variously called, and even were described as species, subspecies,
forms, ecomorphs, etc. Moreover, possible transition of charrs originating from sympatric
speciation from one form to other in the ontogeny, and, in some cases reproductively isolated and
morphologically, ecologically and genetically different forms (Alekseyev et al. 1999, Dynes et al.
1999) complicate the situation and taxonomic status of particular stocks. Berg (1932) recognized 25
valid species in Europe, afterwards he (Berg 1948) lumped them into two. One is Salvelinus alpinus
(Linnaeus, 1758) occurring in the lakes of Finland, Sveden (lake Vättern), southern Norway, in
Alpine lakes (lakes in the watersheds of the rivers Danube, Rhine, Rhône), lakes on the British
Isles, Orkney and Shetland islands, in Iceland, Kola peninsula, Karelia and in the lakes Onega and
Ladoga. The second charr species is Salvelinus lepechini (Gmelin, 1780) inhabiting some lakes in
Finland, Sweden (lakes Mälaren and Vennern) and southern Norway. Ladiges & Vogt (1979) recognised
only one species, Salvelinus alpinus (Linnaeus, 1758) with 24 subspecies. According to
Kottelat (1997) who extensively reviewed all available data, there are 23 valid species in Europe: 7
in England, Wales and Scotland, 6 in Ireland, 3 in Iceland, 2 in the Alps and 4 different species in
Scandinavia, in the Onega and Ladoga lakes, in the Shetland, Orkney and Faroe islands, respectively.
In the Danube river watershed Kottelat (l.c.) distinguishes Salvelinus umbla and Salvelinus
profundus. The name Salvelinus umbla has nomenclatorial priority over the name S. salvelinus
and S. alpinus salvelinus (Linnaeus, (1758) (Kottelat 1997), which was traditionally used just
for charrs of lakes in Alps.
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Salvelinus umbla (Linnaeus, 1758)
The Alpine charr, was described from various Alpine and pre-Alpine lakes, among them also
Mondsee, Traunsee, Königssee and lakes of the Salzkammergut region in Austria, all in the watershed
of the Danube river. Three different forms were known to inhabit them: (1) the dwarf deepwater
Tiefseesaibling (also known under the names Schwarzreuter and/or Hungersaibling) attaining
170–250 mm in Tl, inhabiting the deep parts of the lakes and feeding on zooplankton, (2) the
Normalsaibling feeding on zooplankton and zoobenthos, and (3) the Wildfangsaibling which is
the largest (attaining 650–750 mm in Tl and 3–5 kg in weight), predatory and occurs in two forms,
the yellow and white one (Buresch 1925, Haempel 1924, 1930, Berg 1932, Ladiges & Vogt 1965).
Heckel & Kner (1858) and Schindler (1940) supposed that the Normalsaibling and the Wildfangsaibling
are only different age classes of the same species. Kottelat (1997) suggests that in these
lakes were at least two species, the deep-water and the “normal” one, which were morphologically
distinct, and with different parental ancestry and descent. There is no doubt that the deep-water
Tiefseesaibling belongs to different species Salvelinus profundus as it will be shown below.
Morphology and ecology of these forms in Alpine lakes of the Danube basin are not properly
known and the more or less complete information for the “normal” form are those by Dörfel (1974)
from the Lake Constance (Bodensee; the Rhine River Basin). He found that it dwells in depth not
exceeding 60 m. The diet is composed of the chironomid pupae (75%), copepods (15%) and
chironomid larvae (10%). Spawning periods in first half of December and sexual maturation sets at
age 3 years. Sexual dimorphism is well developed and in addition to more bright colouration, males
display also a kype of their lower jaw. Maximum size was 278–441 mm Tl at 5 years. Fecundity of
this form varied from 900 to 2000 eggs. Concerning the present status of this charr species in
Austria, the following information are mostly from Wanzenböck (pers.comm.) and Jagdsch
(pers.comm.). Indigenous populations in Austria still exist in 16 lakes. In two additional lakes
(Irrsee in Upper Austria, Zellersee in the Salzburg county) the Alpine charr became extinct in the
1960ies following eutrophication and in the medieval time because of mining, respectively. A
restocking program of the Zellersee was established in 1985. In the lakes Grundlsee, Altausser-See
and also in the Attersee, this charr is still the commercially most important species together with
whitefish (Coregonus spp.). In other lakes as in Mondsee and Wolfgangsee, the stocks of charr
are much smaller and a following general pattern is observed: increasing eutrophication is accompanied
with decreasing density of the charr, the dominant position of which is replaced by the
whitefish. This phenomenon when salmonids are replaced by coregonids is well known also from
other oligotrophic lakes in Eurasia and North America (see e.g., Colby et al.1977, Reshetnikov 1980,
Hol#ík et al.1989). Wanzenböck suggests that competition between char and whitefish played also
an important role, since alternating trends in the stock developments are obvious: overfishing of
charrs led to increase in whitefish stock and vice versa. Both authors point out that the most
serious problem might be the mixing of stocks from different lakes leading to the genetic erosion of
different forms adapted to the specific conditions of particular lakes. Stocking material sometimes
does not come from nearby lakes but also outside Austria, even from Scandinavia and Canada
(Jagdsch 1987). With re-oligotrophication of many other lakes (e.g., Mondsee) signs of improvement
of the charr stock have become obvious in recent years. In many small lakes in the high Alps,
the charr have been introduced since the Middle Ages in habitats in which the fish can just
survive, forming dwarf forms. But because these are not primary habitats for the species, its
disappearance from some of these lakes cannot be regarded as the sign of danger. According to
Wanzenböck the situation of native charrs in Austria is generally not bad, but Jagdsch (pers.
comm.) estimates the char catch much lower than l kg.ha –1 . According to Maitland (1995) who
sampled information by questionnaire in December 1993, 15 Austrian lakes still have indigenous
182

charr population, 13 lakes have indigenous plus introduced stocks from elsewhere and there are
150 lakes with introduced charr. This species is still economically important and the Austrian charr
catch in 1993 was 30 000 specimens taken by anglers and 35 metric tons, fished commercially.
Nevertheless, this species is regarded as endangered (see also Hacker 1983 and Herzig-Straschil
1994).
Salvelinus profundus Schillinger, 1901
It is another mysterious salmonid species reported from the Alpine lakes. It was formally named by
Schillinger (1901) as Salmo salvelinus var. profundus from the lakes Constance and Ammersee.
The proper and taxonomically correct description, however, has never been published. In spite of
this already Berg (1932), then Behnke (1972, 1980), Cavender (1980) and at present Kottelat (1997)
consider it to be a valid species. Its characteristic features are blunt and steeply downward bent
snout, almost inferior mouth, large eyes and large teeth. This species has a lower number of gill
rakers (l9–27, M=22.3) than the “Normalsaibling” (25–31, 0=27.7) and also its structure is different
(Dörfel 1974, Behnke 1980, Cavender 1980). Its colouration is similar to that of the whitefish,
without any spots, and it does not change in the period of spawning. There is no sexual dimorphism.
Its ecology was studied by Brenner (1980) in Atterssee. He found that it dwells in depths
from 40 to 130 m, both sexes became sexually mature in the third year of life, the main spawning
period is between July and the beginning of November. However, sexually mature specimens of
both sexes were caught, and monthly fertilisation tests were positive throughout the year, demonstrating
that this charr reproduces all year round. The main spawning grounds in Attersee were at
depths between 40 and 60 m and consisted of gravel with grains of 1.5–2.5 cm in diameter. The egg
number of females 145–205 mm in Tl varied from 220–260 and decreased with increasing female
size. The food of this charr consisted mainly of crustacean zooplankton, but fish eggs and remainders
of fish were also found in its intestines. The growth of females and males appeared to be the
same and the largest specimens were 6 years old and 160–190 mm Tl. It is of interest that 55 years
ago the growth rate of this charr in the same lake was better, as 4 years old specimens reached 250
mm in Tl (Buresch 1925). It is worthwile to mention also that Brenner (1980) found in Attersee only
this deep-water form but other two forms had disappeared, reportedly due to overfishing. The
same situation probably also happened in other Alpine lakes. From other sources we know, that
this deep-water charr may be found at depths around 100 m together with a whitefish (Coregonus
spp.). When taken on the surface it becomes flatulent. Its maximum size is 150–175 mm. The female
is able to spawn at the size of 100 mm Tl. Spawning period is in December and January. Data
gathered by Dörfel (1974) from the Bodensee (Rhine basin) are similar and convincingly show
morphological and ecological differences between the “normal” and a deep-water form allowing to
admit the species status also to the latter. As Kottelat (l.c.) noted, it should urgently be investigated
if the deepwater populations (of which several seem extinct) from various Alpine lakes are
conspecific or not. It is of interest that in Transbaikalian lake Davatchan the deepwater dwarf charr
occurs, resembling S.profundus in many characters. As pointed out by (Alekseyev et al. 1999) this
form represents an interesting example of parallel evolution in deepwater mountain lakes.
Genus Hucho (Linnaeus, 1758)
Hucho hucho (Linnaeus, 1758)
The present distribution of this species is only a part of the earlier one. Historical records show
that the huchen was quite common in almost all rivers of the Danubian watershed and this species
inhabited almost 12 000 km of rivers in Europe. However, after 1945 the situation changed and by
183

the end of 1980s it disappeared from 39%, became rare in 28% and is now common in only about
33% of its former distribution. The present situation in those countries we have information from,
is as follows. In Switzerland huchen occurred in the Inn River in the Engadin valley, but now is
extinct as during past 50 years was not recorded there (Pedroli et al.1991). In Germany the huchen
still occurs in the Danube and Iller in the vicinity of Ulm, but is rare and maintained by continuous
stocking (Berg et al. 1989, Harsányi & Aschenbrenner 1994, Rösch pers.comm.). Jungwirth
(pers.comm.) reported, that in Austria the huchen status is better than 10–15 years ago. In the
rivers Drau, Mur and Pielach there is about 150 km of sections with very good huchen status and
natural reproduction. There is also extensive stocking of 1, 2 and 3 years old juveniles amounting
annually to 10,000–20,000, 5,000–10,000 and 1,000 specimens, respectively. This production is
used to stocking of the rivers Danube, Mur, Drau, Pielach as well as the Inn and the Enns. Huchen
catch statistics are not available. Jungwirth only mentioned, that in the rivers Drau and Mur annual
catches are 50–60 and 30 large huchens, respectively. In Slovenia the rivers Mura (=Mur), Drava
(=Drau), Sava, Savinja, Krka and Kolpa were known as the good huchen rivers. At present the
huchen is extinct in the Mura and in other rivers the sections inhabited by the species are remarkably
shorter than before. Also the population density of the huchen significantly decreased. In the
Krka river, for instance, 60–65 mating huchens in the Soteska spawning ground could be counted
in the past but today the number of spawners dropped to 5–6. Pov' & Sket (1990) evaluate its
status in the water bodies of Slovenia as endangered and vulnerable. The huchen stock in the
rivers is maintained by stocking (Pov' & Sket 1990, Skalin 1994). We have no information on the
situation in other countries of the former Yugoslavia, but it is hardly better than elsewhere. From
Romania we have reports by Miron (1994) and B&n&rescu (pers.comm.). According to the first
author, there were formerly 16 rivers in Romania with the huchen stock. At present their number
dropped to three, i.e. 18.8% of the former number. During 1961–1979, 13 other rivers were stocked
with huchen but acclimatization (probably also naturalisation) appeared only in 6 (46%), the negative
one in 4 (31%) and there are no data for other three rivers (23%). According to B&n&rescu the
huchen in Romania underwent a numerical decline, but is not in danger of extinction. The huchen
is also present in some man-made lakes as is the Bicaz Lake ( on Bistricza River) its spawning
however, is limited to tributary streams. An extensive stocking program was adopted, but its
success depends on the water quality of the rivers and especially on hydraulic engineering, as the
program for damming Romanian rivers seems to expands significantly. In Slovakia, the range of
this species has considerably decreased too (Hol#ík 1990, Hol#ík et al. 1988). Some fifty years ago
the huchen inhabited more than 1000 km of rivers in Slovakia. At present the huchen disappeared
from 48% of the length of all streams in which it occurred in the past, it is rare in 12% and common
in 40% of the total length of the streams. Analysis of historical records indicates that the population
density of the huchen was formerly relatively high, supporting catches of up to 20 huchen of
3–30 kg in weight within a day at the same place. Recent observations in Slovakian rivers indicate
that in spite of some protective measures like the legal restrictions, involving the bag and size
limits and the closed season along with stocking, establishment of a reserve, the huchen in Slovakia
is steadily decreasing. As it follows from the statistical data the catches of huchen shows a
decreasing trend in numbers: the mean catch in periods 1954–1969, 1970–1978, 1979–1989 and in
1990–1995 was 154, 112, 92 and 54 specimens of the huchen, respectively. The increasing mean
individual weight in these periods shows the reverse trend as it rose from 5.35 kg to 5.94, 6.67 and
7.3 kg, respectively. This indicates, that the population of huchen in the rivers of Slovakia becomes
older. In other words both the natural reproduction of the huchen and its stocking are not sufficient
enough to compensate its mortality. Recently Hol#ík (1997) suggested that if the observed
decreasing trend continues, in 2017 the last huchen will be taken in Slovakia. Status of the huchen
184

in Poland was recently reported by Witkowski & Kowalewski (1994). Here the natural occurrence
of the huchen was limited to two streams only, the Czarna Orawa and Czadeczka, belonging to the
Danube river basin. Its density there was rather low as both represented the upper range of its
natural occurrence. At the beginning of the 50’s the huchen disappeared from the Czadeczka
stream because of the pollution and after the construction of the Orava dam in the territory of
Slovakia and subsequent construction of weirs in the Czarna Orawa river the occurrence of the
huchen in the Czarna Orawa was reduced to 8–10 km only. Although the Polish stocking of the
Dunajec, Poprad, Skawa, Sola and Raba rivers was successful and despite the occurrence of the
huchen in about 300 km of streams, the general decrease of this valuable species in the Danube
basin could not be reversed. Generally, the present occurrence of the huchen is no longer continuous
in many places, as it was in the past, and these are often but isolated localities. As in other
salmonids reasons for the decline is the denaturalization of the huchen habitat caused by industrial
development and, later, by intensive large-scale agriculture. Destructive factors include the
canalisation of rivers, construction of dams, the release of industrial waste waters and municipal
sewage, eutrophication, deforestation and expansion of arable land. Overfishing, including poaching,
is only a secondary factor, as fishing for the huchen requires some experience and special
equipment. This factor had more effect before World War II, when legal restrictions were very
loose, allowing anglers to catch unlimited bags of partly immature fish, and the fishing season
lasted 9 months. Considering all these facts the present status of this valuable fish has to be
evaluated as critically endangered.
A c k n o w l e d g e m e n t s
This paper would never been completed without the generous help of P. B&n&rescu (Bucarest, Romania), R. Suciu
(Tulcea, Romania), A. Jagdsch (Scharfling, Austria), T. Jungwirth (Vienna, Austria), R. Rösch (Langenargen,
Germany), and J. Wanzenböck (Mondsee, Austria) who supplied information on the recent status of particular
species of salmonids in their countries. M. Kottelat (Cornol, Switzerland), and P. Ráb (Lib!chov, Czech Republic)
are warmly acknowledged for their critical comments, suggestions and notes, and K. Lohniský (Hradec Králové,
Czech Republic) for supply of some literature. I am grateful to J. Fott, D. Král (Praha, Czech Republic) and an
anonymous reviewer for their comments.
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188

Acta Soc. Zool. Bohem. 66: 189–203, 2002
ISSN 1211-376X
Effects of acid atmospheric deposition on chemistry and benthic
macroinvertebrates of forest streams in the Brdy Mts (Czech Republic)
Jakub HORECKÝ 1) , Evžen STUCHLÍK 1) , Pavel CHVOJKA 2) , Peter BITUŠÍK 3) ,
Marek LIŠKA 4) , Petra PŠENÁKOVÁ 1) & Jan ŠPA"EK 5)
1)
Department of Hydrobiology, Charles University, Vini#ná 7, CZ–128 44 Praha 2, Czech Republic;
e-mail: horecky@natur.cuni.cz
2)
Department of Entomology, National Museum, Kunratice 1, CZ–148 00 Praha 4, Czech Republic
3)
Faculty of Ecology and Environmental Sciences, Technical University in Zvolen, Kolpašská 9B,
SK–969 01 Banská Štiavnica, Slovakia
4)
Vltava River Authority, Hole#kova 8, CZ–150 24 Praha 5, Czech Republic
5)
Labe River Authority, Víta Nejedlého 951, CZ–500 03 Hradec Králové, Czech Republic
Received May "0, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. Water chemistry and macroinvertebrates of 8 streams in the Brdy Mts (Central Bohemia)
differing in pH value were studied during a synoptic survey in December 1997. Together 73 macroinvertebrate
taxa were identified, varying in number from 9–25 at sites of critical pH (3.8–4.3), contrary to 25–28 taxa
at non-acidified sites (pH around 6.7). The fauna of streams was dominated by Chironomidae (Diptera) and
Plecoptera, Trichoptera were less abundant. In three strongly acidified streams the predominance of
Plecoptera exceeded 50% of all macroinvertebrates. The absence of some aquatic insects (like e.g.
Ephemeroptera, some Trichoptera, most Diptera, and some Plecoptera) at sites with pH < 4.8 is discussed.
The streams were classified according to their macroinvertebrate communities, with use of the Jaccard
index of similarity. Symposiocladius lignicola (Kieffer, 1915) (Diptera: Chironomidae) is reported from
Bohemia for the first time.
Acidification, running waters, winter season, water chemistry, benthic fauna, macroinvertebrates,
Czech Republic, Palaearctic region
INTRODUCTION
The majority of the regional knowledge concerning processes of anthropogenic surface water
acidification and its effects on biota is associated with the studies in the mountains on the borders
of the former Czechoslovakia. The high mountain lakes in the Tatra Mts (Kopá#ek et al. 2000), the
forest lakes in the Bohemian Forest (Veselý et al. 1998, Vrba et al. 2002), mountain drinking water
reservoirs in the Jizera Mts (Stuchlík et al. 1997) and some streams in the Ore Mts (Krám & Hruška
1994) have been intensively studied for more than 20 years, providing detailed information about
the impact of acid deposition on freshwater ecosystems. The response of plankton communities to
pH changes was investigated, including the response to oligotrophication and the toxic role of
aluminium in particular (Fott et al. 1994).
The Brdy Mts situated in Central Bohemia have been historically known as an area rich in large
peat bogs, but their sensitivity to acid deposition was not elucidated until a detailed water chemistry
survey performed in 1984 (Veselý & Majer 1996). Surface waters of this area, represented
mostly by brooks and drinking water reservoirs, exhibit mean pH values lower than 5.5, indicating
This contribution is dedicated to Doc. RNDr. Jaroslav Hrbá!ek DrSc., upon the occasion of the 80th anniversary
of his birthday ("2th May "92").
189

that the Brdy Mts are among the regions of the Czech Republic which were the most intensively
impacted by acid atmospheric deposition (Veselý & Majer 1998).
While the effect of acidification on the macroinvertebrates of running waters in Europe and
North America is generally well documented (e.g., Hall & Ide 1987, Weatherley et al. 1989, Mulholland
et al. 1992, Herrmann et al. 1993, Guerold et al. 1995, Lien et al. 1996, Orendt 1998, Szcz*sny
1998), there is a lack of such information from the Czech Republic and only several limited studies
have been published so far (R+ži#ková 1998, Scheibová & Helešic 1999).
Our knowledge on the benthic invertebrate fauna of the Brdy Mts is rather fragmentary, as the
possibility of field research has been limited due to the existence of a military area in this region
since 1926. One of the first records of the Brdy Mts freshwater organisms were presented at the
Fig. 1. Stream system of the Brdy Mts with the sampling sites. ⊗ – sampling site, triangle – Nepomuk precipitation
station.
190

eginning of the 20 th century by Roubal & Štorkán (1924). Basic information on the fauna of
Ephemeroptera, Plecoptera and Trichoptera were obtained thanks to long-term research of several
streams in the Brdy Mts (mainly at lower altitudes) in the late 1950s (K%elinová 1962, Novák 1962,
pers. comm., Landa & Soldán 1989) and only a single site was sampled in the mid-1990s again
(Soldán et al. 1998). Results of a hydrobiological study directed to small streams in the central part
of the Brdy Mts including a brief list of macroinvertebrate genera were published by Pivni#ka et al.
(1993).
The main aim of this pilot study is to recognise the impact of acid atmospheric deposition on
water chemistry, species composition and diversity of benthic fauna in several brooks of these
mountains .
SITES AND METHODS
Study area
The Brdy Mts (49° 32’ N, 13° 42’ E – 49° 50’ N, 14° 05’ E) represent the largest forested complex in the central
part of the Czech Republic being situated in the Labe basin, about 50 km SW of Prague (Fig. 1). The maximum
altitude of the mountain range is 864.9 m a. s. l. and the altitude of our study sites ranges from 470–685 m a. s. l.
(according to base maps 1:50 000 – Mašek 1990). The Brdy Mts are covered mostly by spruce (Picea abies)
forests and peat bogs are frequent in spring areas.
The geology of the Brdy Mts is dominated by cambric sandstone, conglomerates, and quartzites. These rocks,
with a low content of calcium and other nutrients, form nutrient-poor brown soils that tend to form high moor
bogs at the highest flat altitudes. Brown soils relatively rich in nutrients occur locally, especially in the north-west
mountain part where late Variscan granodiorites and diabase vulcanites encroach. The southern part contains
Middle Proterozoic shales and greywackes with numerous phthanitic intercalations (Pet%í#ek & Dejmal 1998,
Havlí#ek 1986, Mašek, 1990).
Eight sites comparable in vegetation but varying in stream size, pH and altitude were selected on the following
brooks: Voldušský (VOL), "ervený (CER), Mourový (MOU), Reserva (RES), T%ítrubecký (TRI), Kotelský
(KOT), Smolivecký (SMO) and Litavka (LIT) (Tab. 1, Fig. 1). Although spruce forest covers most of the
catchments of all brooks, none of them is typical brown water. The geology of catchments of the studied brooks,
with the exception of Voldušský brook and Kotelský brook, is similar to the general characteristics of the Brdy
Mts. Intercalations of shales and larger areas of basalt must be emphasised in the case of Voldušský brook and
Kotelský brook, respectively. The sampling site at "ervený brook is situated below a small fishpond, and,
consequently, some liming effect may occur since the upper part of the catchment is strongly acidified (Stuchlík
unpubl. data). Occasional liming is performed in the catchment of Litavka brook due to water quality concerns in
a drinking water reservoir closely downstream the sampling site (Kulina 2000). The lowest discharge during the
sampling time was registered at Smolivecký brook, which is of special character, since its catchment was altered
by forest-drainage measures.
Sampling and analytical methods
The investigation was carried out on 13 th December 1997. Water samples were taken prior to invertebrate ones.
All analyses were conducted in the analytical laboratory of the Hydrobiological station “Velký Pálenec” near
Blatná (Faculty of Science, Charles University). Conductivity was measured by conductometer CDM (Radiometer,
France), pH was recorded at the beginning of determination of alkalinity by Gran titration on the automatic
titrator TIM 900 (Radiometer, France), and the concentration of major ions was analysed by ion chromatography
(Thermo Separation Products, USA). The laboratory participates in regular international intercalibration
runs organised by NIVA, Oslo, Norway and the AQUACON project by CNR-ISE, Verbania-Pallanza, Italy. Precipitation
chemistry was based on data from the Nepomuk precipitation station operated by the Institute for Forest
Ecosystems Research (IFER) and published in the Tabular Survey by the Czech Hydrometeorological Institute
(Fiala et al. 1999).
Benthic macroinvertebrates were collected by kick sampling (net mesh size 0.5 mm) for 5 minutes per site
from riffles with stony or stony-gravely substratum within a 10-m stretch, and preserved in 80 % alcohol.
Invertebrates were identified to species or generic level, if possible, according to keys by Hrab! (1979), Rozkošný
(1980), Wiederholm (1983), Losos (1996), Reusch & Oosterbroek (1997), Thomas (1997), Waringer & Graf
(1997), Krno (1998), Zahrádková & Soldán (1998), Bitušík (2000) and Knoz (2001).
191

Tab. 1. Sampling site description, water chemistry of studied brooks at the sampling sites (the left part of the table, 13 th December 1997) and precipitation
chemistry from the Nepomuk precipitation station (annual weighted mean 1997/98, Fiala et al. 1999)
Category III Category II Category I
Brook Reserva T%ítrubecký Litavka Smolivecký Kotelský Mourový "ervený Voldušský Nepomuk
Code RES TRI LIT SMO KOT MOU CER VOL wet only bulk throughfall
Altitude m a. s. l. 610 570 650 685 665 510 470 480 860
Latitude N 49°42’ 27” 49°41’ 53” 49°39’ 50” 49°34’ 24” 49°35’ 41” 49°45’ 00” 49°46’ 53” 49°48’ 40” 49° 39’ 33’’
Longitude E 13°49’ 13” 13°48’ 30” 13°53’ 17” 13°45’ 45” 13°46’ 22” 13°50’ 00” 13°53’ 11” 13°39’ 56” 13° 49’ 09”
River basin Klabava Klabava Litavka Lomnice Skalice Litavka Litavka Klabava
Max. width m 3 3 2 1 1 2 3 2
Dist. from source km 4.3 5.6 2.1 1.5 0.5 4.7 7.5 1.7
Precipitation mm y –1 762 752 595
K20 µS cm –1 107.3 83.8 83.7 78.8 79.9 64.8 68.9 165.6 19.2 18.6 67.2
pH 3.9 4.2 4.3 4.4 4.9 5.3 6.6 6.7 4.6 4.6 4.1
Alk µeq l –1 –130.6 –70.5 –54.0 –41.4 –6.2 7.1 120.3 203.7 – – –
NH4 – -N µg l –1 0 0 0 0 0 0 0 0 397 498 576
Ca 2+ mg l –1 4.28 3.27 3.68 4.12 5.82 5.78 6.11 15.67 0.13 0.21 1.02
Mg 2+ mg l –1 1.20 1.42 1.97 2.62 3.38 2.14 2.84 12.69 0.02 0.04 0.18
Na + mg l –1 1.06 1.15 1.69 2.06 2.17 1.47 1.99 3.38 0.13 0.20 0.60
K + mg l –1 1.08 1.17 1.41 1.72 1.39 1.20 1.05 1.50 0.05 0.16 2.76
F – µg l –1 125 121 112 140 145 128 162 112 20 20 50
Cl – mg l –1 1.88 2.20 1.52 2.07 2.14 1.86 2.45 2.56 0.23 0.35 1.44
NO2 – -N µg l –1 6 2 2 6 4 7 2 1 0 0 0
NO3 – -N µg l –1 997 843 242 445 452 836 388 432 431 452 876
SO4 2– mg l –1 20.90 23.12 20.18 23.90 22.10 24.68 22.43 17.58 1.60 1.87 8.15
Tab. 2. Total (N) and relative (%) abundance of organisms of 8 sampling sites in the Brdy Mts found on 13 th December 1997. 1) Nemoura cf. cambrica
(Stephens, 1836); 2) Nemoura cf. uncinata Despax, 1934
RES TRI LIT SMO KOT MOU CER VOL
Reserva T%ítrubecký Litavka Smolivecký Kotelský Mourový "ervený Voldušský
N % N % N % N % N % N % N % N %
OLIGOCHAETA
Bythonomus lemani (Grube, 1879) " 0.4 " 0.4
Lumbriculus sp. " 1.5
Stylodrilus sp. " 1.5 4 13.8 " 0.4
Lumbriculidae gen. sp. 2 3.0
Naididae gen. sp. " 1.5 " 0.4
Eiseniella tetraedra (Savigny, 1826) " 0.4
Oligochaeta gen. sp. 2 4.5
EPHEMEROPTERA
Baetis rhodani Pictet, 1845 " 0.4 2 0.7 " 0.2 " 0.3
192

Ecdyonurus torrentis Kimmins, 1942 " 0.4 " 0.2 4 1.2
Rhithrogena iridina (Kolenati, 1859) " 0.4
Leptophlebia vespertina (Linnaeus, 1758) 2 0.8 " 0.3 5 1.5
Paraleptophlebia submarginata (Stephens, 1835) 3 0.7
PLECOPTERA
Amphinemura borealis (Morton, 1894) 20 7.8 22 7.4
Amphinemura sp. juv. " 2.3 2 0.7 2 0.4
Nemoura avicularis Morton, 1894 2 0.8 8 1.8 2" 6.2
Nemoura cambrica (Stephens, 1836) 6 9.1 5 1.5
Nemoura flexuosa Aubert, 1949 " 0.2
Nemoura spp.
" )3 4.5
2 )3 6.8
2 )"2343.6
2 )3 1.0
2 )6013.3
" )25 7.3
Nemurella pictetii Klapálek, 1900 " 1.5 3 1.1 4 13.8 3 0.7
Protonemura auberti Illies, 1954 "3 19.7 25 56.8 77 27.3 "6 5.4
Protonemura meyeri (Pictet, 1841) 48 18.6 20 5.4
Protonemura sp. juv. " 0.2
Leuctra hippopus Kempny, 1899 " 0.4 "6 6.2 "7 5.7 37 8.2 55 16.1
Leuctra nigra (Olivier, 1811) 3 4.5 5 11.4 23 8.2 4 13.8 8 3.1 23 7.8 65 14.4 35 10.3
Leuctra prima Kempny, 1899 2 0.4
Leuctra rauscheri Aubert, 1957 "2 4.7 30 10.1
Diura bicaudata (Linnaeus, 1758) 6 9.1 " 2.3 " 0.4
Siphonoperla torrentium (Pictet, 1841) " 0.4
ODONATA
Cordulegaster annulatus (Latreille, 1805) " 0.3
MEGALOPTERA
Sialis fuliginosa Pictet, 1836 2 3.0 " 0.4 " 0.2 " 0.3
TRICHOPTERA
Rhyacophila tristis Pictet, 1834 3 4.5
Rhyacophila vulgaris Pictet, 1834 species group 2 4.5 " 0.4 2 0.7 " 0.2 " 0.3
Plectrocnemia conspersa (Curtis, 1834) 9 13.6 9 3.2 " 3.4 4 1.6 5 1.7 " 0.2 "0 2.9
Hydropsyche saxonica McLachlan, 1884 " 0.2
Hydropsyche sp. juv. 4 1.6 3 0.7
Drusus annulatus (Stephens, 1837) 5 7.6 2 4.5
Potamophylax sp. 4 1.2
Limnephilidae gen. sp. juv. " 1.5 2 0.7 " 0.4 " 0.2
Sericostoma sp. 2 0.8
Odontocerum albicorne (Scopoli, 1763) " 0.3
DIPTERA – excl. Chironomidae
Dicranota spp. 2 3.0 2 4.5 3 1.1 2 0.8 7 2.4 " 0.2
Pedicia (P.) cf. rivosa (Linnaeus, 1758) 3 1.1 " 3.4
193

Tab. 2. continuation
RES TRI LIT SMO KOT MOU CER VOL
Reserva T%ítrubecký Litavka Smolivecký Kotelský Mourový "ervený Voldušský
N % N % N % N % N % N % N % N %
Tipula sp. 3 0.9
Bezzia sp. 2 0.8
Prosimulium sp. " 0.4
Simulium (Nevermannia) spp. " 1.5 4 1.4 6 2.0 9 2.0 3 0.9
Simulium (S.) spp. 8 2.8 2 0.8 "0 3.4 "0 2.2
Ibisia marginata (Fabricius, 1781) 6 2.0 5 1.1
Wiedemannia sp. " 1.5 " 0.2
DIPTERA – Chironomidae
Apsectrotanypus trifascipennis (Zetterstedt, 1838) " 0.4
Conchapelopia sp. "2 4.7 2 0.4 3 0.9
Macropelopia sp. " 0.4 7 24.1
Trissopelopia sp. 2 0.4 " 0.3
Brillia modesta (Meigen, 1830) " 0 3.5 4 1.6 3 1.0 26 5.8 9 2.6
Heterotrissocladius marcidus (Walker, 1856) 5 17.2
Limnophyes sp. " 1.5
Rheocricotopus fuscipes (Kieffer, 1909) " 0.3
Symposiocladius lignicola (Kieffer, 1915) " 0.4
Tokunagaia cf. rectangularis (Goetghebuer, 1940) " 0.4 " 0.3
Tvetenia calvescens (Edwards, 1929) 3 1.2
Tvetenia discoloripes (Goetghebuer, 1936) " 0.4 " 0.3
Orthocladiinae gen. sp. " 0.4 "8 4.0
Micropsectra spp. 3 4.5 3 1.1 " 3.4 84 32.6 "22 41.2 "86 41.2 "4442.2
Polypedilum laetum (Maigen, 1818) species group " 0.4
Polypedilum scalaenum (Schrank, 1803) species group " 0.3
Tanytarsus sp. " 0.4 2 6.9
COLEOPTERA
Agabus sp. (larva) " 0.4
Deronectes platynotus (Germar, 1834) (imago) " 2.3
Helophorus sp. (larva) " 1.5
Helodes spp. (larvae) " 0.4 " 0.3 5 1.5
Elmis spp. (larvae) "7 6.6
Limnius sp. (larva) " 0.4
Tot. ind. (N) 66 44 282 29 258 296 452 341
194

Biological classification of the sites was based on the calculation of Jaccard’s index (Ludwig & Reynolds 1988).
Then, the Ward’s method of linkage was used to construct a dendrogram. Statistical analyses were performed using
STATISTICA software (StatSoft, Inc., USA).
RESULTS
Water chemistry
All studied sites represented poorly buffered soft water brooks very sensitive to acid atmospheric
deposition, with ion compositions dominated by sulphate, calcium and magnesium (Tab. 1). The
brooks were divided into three distinct categories according to their water chemistry (mainly pH,
alkalinity, and calcium and magnesium concentrations) as follows: Category I was defined as pH >
5.8 (mean Gran alkalinity of 162 µeq l –1 , and mean concentration of Ca 2+ and Mg 2+ of 10.8 and 5.33
mg l –1 , respectively). Category II was defined as pH 4.8–5.8 (mean alkalinity of 0 µeq l –1 , mean
concentration of Ca 2+ and Mg 2+ of 5.80 and 2.86 mg l –1 , respectively). Category III was defined as
pH

other groups of the order Diptera. Records of Symposiocladius lignicola, Rheocricotopus fuscipes,
Tvetenia discoloripes and T. calvescens are published from Bohemia for the first time. Considerable
density of Coleoptera was found at Kotelský brook only (see Tab. 2)
The dendrogram of site dissimilarities (Fig. 4) shows two distinct groups – the first one consists
of the four sites of category III (Litavka, Smolivecký, T%ítrubecký and Reserva brooks) and the
second one of the remaining sites that cannot be further distinguished.
DISCUSSION
The low pH and high concentrations of sulphate and DIN in all types of precipitation suggest the
atmospheric deposition to be the major source of acidity in the brooks studied. The observed
extremely low (

Tab. 3. Number of taxa per taxonomic group from Tab. 2
RES TRI LIT SMO KOT MOU CER VOL
Reserva T%ítrubecký Litavka Smolivecký Kotelský Mourový "ervený Voldušský
Oligochaeta 4 1 1 1 4 – – –
Ephemeroptera – – – – 4 2 3 3
Plecoptera 6 5 7 2 7 7 9 5
Odonata – – – – – – – 1
Megaloptera 1 – – – 1 – 1 1
Trichoptera 4 2 3 1 4 2 5 4
Diptera 5 1 12 5 10 6 10 10
Coleoptera 1 1 2 – 2 1 – 1
Suma 21 10 25 9 32 18 28 25
Even though aluminium was not analysed in this survey, our later detailed study on Litavka
brook chemistry confirmed the presence of high concentrations of reactive aluminium (up to 2000
µg l –1 ; Kulina 2000, Stuchlík et al. 2000). Labile aluminium, most toxic to water organisms, constituted
up to 70 % of that aluminium pool. This high ratio of labile to reactive aluminium in the category
Fig. 3. Relative numbers of macroinvertebrate taxonomical groups (%, OLI–Oligochaeta, EPH–Ephemeroptera,
PLE–Plecoptera, MEG–Megaloptera, TRI–Trichoptera, DIP–Diptera excluding Chironomidae, CHI–Chironomidae,
COL–Coleoptera). Site abbreviations as in Tab. 1.
197

III brooks clearly indicates that DOC concentrations are not high enough to ameliorate significantly
the toxic effects of aluminium on macroinvertebrates (Fott 1994).
Atmospheric input of sulphate and DIN in the Brdy Mts is within a typical range published for
the other forest regions in the Czech Republic (Fiala et al. 1999). The observed sulphate concentrations
in the brooks studied are substantially higher than could be expected from just an evapotranspiration
effect, suggesting some additional sulphate sources in their catchments. Such a situation
is typical for the ecosystems recovering from acidification and is associated with a wash-out
of sulphate accumulated in soils during previous decades of its elevated deposition (Alewell 2000,
Prechtel et al. 2001). This washing out may last several decades, as predicted for the geographically
adjacent mountain areas (Kopá#ek et al. 2001, Hruška et al. in press). In contrast to sulphate,
nitrate concentrations in the brooks are low, indicating that the forest ecosystem of the Brdy Mts
is probably in the early stage of nitrogen saturation according to Stoddard (1994). The elevated
export of aluminium from a catchment usually accompanies leaching of strong acid anions from the
acidified soils. Due to the dominant role of sulphate in the total pool of strong acid anions in the
brooks studied, the future extent of aluminium concentrations, as well as the associated toxicity
will predominantly reflect the fate of sulphur accumulated in soils. Consequently, the elevated
aluminium concentrations can be expected in the brooks also for the next several decades despite
a >80 % decrease in sulphate emission in the Central Europe (Kopá#ek et al. in press).
Comparable values of conservative ions (e.g., sulphate and chloride) in the brooks (Tab. 1)
suggest that even the limited data based on a single synoptic survey can provide a reasonable
basis for classification of the water acidity status.
The highest total abundance of water macroinvertebrates was recorded in all streams of categories
I and II and also in Litavka brook. The low numbers of individuals in Reserva and T%ítrubecký
brooks may be explained not only by the effects associated with acidification, but also by other
environmental factors, as the influence of acidification on aquatic macroinvertebrate abundance
has not been explicitly proven (Herrmann et al. 1993). Mulholland et al. (1992) consider the densities
of shredder and collector-gather macroinvertebrates to be correlated rather to benthic organic
matter than to pH. In the case of Smolivecký brook, the total abundance might also be poor due to
its drainage ditch character. Most of the collected organisms were insects, of which Plecoptera and
Diptera were the most abundant. A comparison of the composition of macroinvertebrate communities
of the streams studied in the Brdy Mts shows that number of invertebrate taxa is reduced as a
consequence of acidification. A decrease in species richness and diversity, and changes of composition
of functional feeding groups in acidified streams, are well documented effects of acidification
(e.g., Økland & Økland 1986, Szcz*sny 1990, Guerold et al. 1993, Herrmann et al. 1993,
Dangles & Guerold 2000). This feature results from the disappearance of Ephemeroptera, Mollusca
and Crustacea, and reduced richness of other groups (Diptera, Trichoptera, Coleoptera, Plecoptera,
see e. g., Raddum & Fjellheim 1984, Weatherley et al. 1989, Guerold et al. 1995, Szcz*sny 1998).
Differences in tolerance to acid waters can be found within stonefly (Plecoptera) larvae, among
which Siphonoperla torrentium is considered to be moderately sensitive (Braukmann 1994), as is
Diura bicaudata (occurrence at pH > 5 according to Fjellheim & Raddum 1990). Other authors
(Braukmann 1994, Soldán et al. 1998), however, consider the latter species as acid-tolerant. This
species was already reported from the Brdy Mts by K%elinová (1962) and Pivni#ka et al. (1993), and
found ourselves in 1997 in the most acid brooks: Reserva, T%ítrubecký and Litavka. Leuctra hippopus
and Nemoura avicularis appeared to be associated with waters of pH above 4.8 in the study
area, although they could be expected at lower pH values as well (Fjellheim & Raddum 1990);
Scheibová & Helešic (1999) found N. avicularis at a minimum pH of 5.5. Amphinemura borealis,
Nemurella pictetii, Leuctra nigra and L. rauscheri belong to the extremely acid-tolerant stone-
198

flies (Fjellheim & Raddum 1990, Braukmann 1994), which is in agreement with our results. The
occurrence of Amphinemura borealis and Diura bicaudata, vulnerable and endangered species
in the Czech Republic (Soldán et al. 1998), deserve attention from the faunistic point of view.
Heterotrissocladius marcidus, an acid-tolerant chironomid species (Orendt 1999), occurred at
relatively high abundance in acid Smolivecký brook. The presence of Apsectrotanypus trifascipennis
larva in Litavka brook (pH 4.3) is in contradiction with its extreme acid-sensitivity supposed
by Orendt (1999), though in this case it can be associated with an imbalance of the acidity state as
a consequence of liming. The increase of Micropsectra spp. abundance with increasing pH in
investigated streams might reflect its moderate sensitivity to acid conditions. The record of Symposiocladius
lignicola is interesting from the faunistic point of view; this species, so far known
from Moravia and Slovakia (Bitušík & Losos 1997), is reported from Bohemia for the first time. Due
to specific morphological features of its larva, this species is easily distinguishable from other
species of the subfamily Orthocladiinae (Bitušík 2000). The occurrence of larvae of the Polypedilum
(Tripodura) scalaenum group in Bohemia was shown only recently (Chalupová & Mat!-
na 2002). Other species recorded not listed in Bitušík & Losos (1997), namely Rheocricotopus
fuscipes, Tvetenia discoloripes and T. calvescens, occur in the Czech Republic frequently (Mat!-
na, pers. comm.).
Fig. 4. Biological classification of sites based on Jaccard’s index of similarity; recorded water pH is indicated bellow.
Site abbreviations as in Tab. 1.
199

Rhyacophila larvae (Trichoptera) are generally regarded as acid-tolerant, with the exception of
R. tristis which is considered more sensitive (Guerold et al. 1993, 1995, Braukmann 1994), preferring
waters with pH 5.4–7.4 (Novák 1962). Surprisingly, R. tristis was found in Reserva brook at pH 3.9.
Plectrocnemia conpersa exhibits a great tolerance to acidification as has been well documented
earlier (e. g., Fjellheim & Raddum 1990, Scheibová & Helešic 1999). This most abundant caddisfly
species of streams in the Brdy Mts occurs independent of acidity. Drusus annulatus was recorded
only in streams of category III; larvae of this common mountain species of the Czech Republic also
inhabit streams with low pH in other acidified regions (Horecký unpubl. data). This species was
reported as acid-tolerant e.g. from the Vosges Mts (Guerold et al. 1995), although Szcz*sny (1990,
1998) did not find it in acidified streams of the West Carpathians.
The majority of Ephemeroptera species are strongly acid-sensitive and therefore become extinct
in waters below pH 5–5.5 (Økland & Økland 1986, Herrmann et al. 1993, Havas & Rosseland
1995). In the investigated streams, mayflies were recorded at sites with pH > 4.8. Baetis rhodani,
considered a potential indicator species of non-acidified waters (Larsen et al. 1996) and extremely
acid-sensitive (e.g. Scheibová & Helešic 1999), was found at pH around 5 in the Brdy Mts, which
better correspond to Engblom & Lingdell (1984), who mention this species as moderately acidtolerant
showing somewhat lower pH tolerance limits (4.5). On the other hand, we recorded Leptophlebia
vespertina only in streams of categories I and II although this species is known as
extremely acid-tolerant (Larsen et al. 1996) and Engblom & Lingdell (1984) report survival of this
species up to pH 3.5. Its absence at sites with the lowest pH might be the consequence of unfavourable
flow conditions there, since L. vespertina larvae prefer habitats with low current speed or
pools (Soldán et al. 1998).
The permanent fauna is represented only by Oligochaeta found in streams with pH < 5. Since
Oligochaeta belong to acid-tolerant organisms often forming the major component of the bottom
fauna under acidic conditions (Weatherley et al. 1989, Guerold et al. 1995), their absence or low
abundance can reflect the absence of fine sediments at the sampled sites or the sampling procedure.
Aquatic Mollusca and Crustacea, generally considered acid-sensitive (e. g., Økland & Økland
1986), were not found at the studied sites, but occur in more alkaline waters of the Brdy Mts
(Roubal & Štorkán 1924, Pivni#ka et al. 1993, Horecký unpubl. data).
The biological classification corresponds well with characteristics based on water chemistry
data distinguishing the group of streams highly impacted by atmospheric deposition (category
III). Although point liming at the Litavka catchment results in episodic extremes of water pH up to
neutral or alkaline values, the site still appears to be unfavourable for mayflies, either due to pH
imbalance (Kulina 2000, Stuchlík et al. 2000) or low minimum pH. On the other hand, relatively low
differences in species composition of the remaining sites (categories I and II) probably represent
a consequence of episodic acidification. Our results confirm the importance of biological assays,
especially in the case of non-frequent water sampling.
A c k n o w l e d g e m e n t s
The authors wish to thank David Hardekopf for revision of the English text, Tomáš Soldán (Institute of
Entomology, Academy of Sciences of the Czech Republic, "eské Bud!jovice) for the revision of Ephemeroptera
larvae, and Josef Mat!na and Ji%í Kopá#ek (Hydrobiological Institute, Academy of Sciences of the Czech Republic,
"eské Bud!jovice), who made valuable comments on early draft of the manuscript.
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Acta Soc. Zool. Bohem. 66: 204, 2002
ISSN 1211-376X
204
BOOK REVIEW
BUSH A. O., FERNÁNDEZ J. C., ESCH G. W. & SEED J. R: Parasitism: The diversity and ecology of animal
parasites. Cambridge: Cambridge University Press, 2001. IX+566 pages. Format 191×245 mm. Soft cover. Price
Lstg 29.95. ISBN 0-521-66447-0
The authors are accredited experts affiliated with the universities and scientific societies in USA and Canada. As
stated in the preface, the present book is derived from another, entitled A functional biology of parasitism
(Chapman & Hall, 1993). The authors provide fundamental insights into the parasitism – the most prevalent,
ubiquitous life style among organisms. Introduced are data that deal with the diversity of major eukaryotic
parasites of animals considering the impact they have on their hosts. Special attention is centred upon ecological
relationships of the host-parasite systems. Not omitted is basic biochemical, molecular and immunological piece
of information relevant to parasitism. The volume is composed of 16 chapters divided into sections (subchapters
– in number from 3 up to 12). Chapter 1 is intended to give the introduction to parasitology while discussing
parasitism in historical perspective, symbiotic relationships, kinds of parasites, kinds of hosts, and ecology in an
environmental context. Immunological, pathological, and biochemical data related to parasitism are outlined in
chapter 2. Consequent thematic block of seven chapters (3 through 9; 265 pages = 47%) provides access to the
systematics of protozoans and metazoans. In chapter 3 examined are three protozoan phyla: the Sarcomastigophora,
Apicomplexa and Ciliophora. It is important to be mentioned that the systematics of the Protozoa is
under constant revision and with the use of molecular procedures will continue to be revised for the foreseeable
future. Newer classifications introduce different schemes and do not include the group Sarcomastigophora.
Microsporidians are now considered to be Fungi. Chapter 4 is devoted to flatworms Platyhelminthes comprising
seven taxonomical groups. It is stated here that flatworms present a diverse group showing great variability in
anatomy, life history, size, and habitat – they are likely ancestors of all the metazoan phyla. Systematic
classification of these parasites has changed significantly in the last decade or two because of morphological
information made available by electron microscopy, the application of cladistic techniques, and molecular
systematics. The classification outlined here is based on studies of Brooks and other authors in the ninethies of the
past century. Chapter 5 focuses on roundworms Nematoda. It is stressed that nematodes are now recognized as
perhaps the most significant metazoan parasites associated with humans. Recent advances in molecular systematics
(18S rDNA sequences) may clarify the phylogenetic relationships in nematodes, nonetheless new proposals
await further verification. The classification scheme used here is one of convenience based on morphological data
more readily available than nucleotide sequences. In subsequent chapters 6 through 8 acanthocephalans, pentastomids
and arthropods are analyzed. In addition to the major phyla of parasitic organisms discussed in previous
chapters, in chapter 9 attention is centred upon other animal phyla that are parasitic in some stage in their life
cycle. With the application of the newer molecular techniques, it is now recognized that myxosporidians Myxozoa,
formerly classified among the Sporozoa, are not unicellular protozoans but a metazoan group. Subsequent
chapters 10 through 16 take into consideration selected general phenomena of parasitism. Chapters 10 and 11
analyse population concepts and factors influencing parasite populations while giving general definitions, and
describing environmental factors, distributional patterns, dynamics of population growths, density-dependent
and -independent factors, regulation of parasite populations. Chapter 12 provides access to the influence of
parasites on host populations: the Crofton’s approach (analysis of host-parasite systems), aggregation and
regulation, epidemiological implications, non-genetic and genetic predispositions. In chapter 13 looked at is the
ability of parasites to complete their life cycle. Chapter 14 gives an overview of parasite communities. In chapter
15 elucidated are biogeographical events and processes, geographical distribution of parasites and diverse aspects
of ecology and applied biogeography. Final chapter 16 illuminates biological evolution. In conclusion, there is an
extensive glossary (14 pages) of key terms relevant to topics discussed. Accentuated on grey background, there
are separate textual boxes containing supplemental discussions and topical observations to the current text. The
volume is extensively illustrated by a wealth of figures numbered separately in each chapter and composed of
schematic line drawings and photographs. Featured are individual parasitic organisms, views of external and
internal structural parts, various developmental stages, life cycles, functional interactions and metabolic pathways,
diverse cladograms and evolutionary trees, miscellaneous diagrams, geographical maps, and selected pathological
conditions in parasitized humans and animals. Moreover, tabular overviews summarizing data given in the
textual part are numerous and detailed. Written in a user-friendly style, this monograph offers an original and
most informative approach to the study of animal parasites of medical, veterinary as well as general zoological
interest.
Jind%ich Jíra

Acta Soc. Zool. Bohem. 66: 205–212, 2002
ISSN 1211-376X
Crustaceans (Crustacea: Cladocera, Copepoda) of the Morava River
Alluvium on the Slovak Territory
Marta ILLYOVÁ 1) & František KUBÍ"EK 2)
1)
Institute of Zoology, Department of Hydrobiology, Slovak Academy of Science, Dúbravská cesta 9,
SK–824 06 Bratislava, Slovakia; e-mail: marta.illyova@savba.sk
2)
Faculty of Science, Masaryk University, Kotlá%ská 2, CZ–611 37 Brno, Czech Republic
Received July 2, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. The paper presents the first comprehensive survey of 58 Cladocera species and 38 Copepoda
taxa from 15 localities of various types of arms, temporary pools and oxbows of the Morava River
inundation on the Slovakian territory. Among the species found, the cladocerans Bosmina coregoni Baird,
1875, Daphnia parvula Fordyce, 1901, D. ambigua Scourfield, 1946, Pleuroxus denticulatus Birge, 1879,
Moina weismanni Ishikawa, 1896 and the calanoid Eurytemora velox (Lilljeborg, 1853) are supposed to be
either invaders or introduced species.Cladocerans Bosmina longirostris (O. F. Müller, 1776), Moina micrura
Kurz, 1874, M. weismanni Ishikawa, 1896, the genus Daphnia and copepods of the genera Eudiaptomus
and Acanthocyclops prevailed in relatively deep side arms situated in the inundation. Phytophilous
assemblages of Daphnia curvirostris Eylman, 1887, Megafenestra aurita (Fischer, 1849), Mixodiaptomus
kupelwieseri (Brehm, 1907), genera Simocephalus, Scapholeberis, andAcroperus dominated in waters of
temporary character and shallow dead arms covered with macrovegetation.
Distribution, wetlands, floodplain, pools, Crustacea, Morava River, Palaearctic region
INTRODUCTION
Aquatic alluvial biotopes are often being terrestrialized. It is caused either by natural ageing or by
anthropogenic intervention. The Slovak part of the Morava River floodplain, especially the inundated
area, is the territory where the diversity of original aquatic biotopes has been preserved.
Several recent studies (Adámek & Sukop 1992, Kopecký & Koudelková 1997, Sukop & Kopecký
1999, Kopecký et al. 1999) have been published on the Cladocera and Copepoda fauna in the
upper section of the Morava River and its tributary Dyje. There is a lack of data on Cladocera and
Copepoda species composition in the lower section of the Morava River on the Slovak territory,
except rare Crustacea in temporary pools (Brtek 1976 and 1992). This lack is caused by prohibited
access to portion of the Morava River that forms the border between Slovakia and Austria over a
long period of time. Systematic monitoring of the study area began only in 1994 thanks to project
on revitalisation of cut-off meanders (Št!rba 1995, Lisický et al. 1997). The result of this project is
the first list of Cladocera and Copepoda species in the Morava alluvium (Illyová 1999).
Our study focuses on different types of standing waters of the Morava River floodplain. Our
goal is to explain the occurrence of cladocerans and copepods in the Slovak portion of the Morava
River.
We dedicate this contribution to Doc. RNDr. Jaroslav Hrbá!ek DrSc., upon the occasion of the 80th anniversary
of his birthday ("2th May "92").
205

Tab. 1. Description of study sites
number of locality name of locality river km cadastre
1 meander XVIII. 65.6–66.0 Moravský Sv. Ján
2 meander XVII. 65.3–65.5 Moravský Sv. Ján
3 meander XVI. 63.5–64.0 Moravský Sv. Ján
4 meander VII. 18.9–20.6 Vysoká pri Morave
5 meander II. 11.7–12.3 Vysoká pri Morave
6 Šrek 11.5–14.5 Stupava
7 Nová Kakvica 22.8–24.7 Vysoká pri Morave
8 Rudavné jazero 51.5–52.1 Malé Leváre
9 Lep0a 57.0–57.9 Malé Leváre
10 Bezodné 13.0 Vysoká p.M.
11 Oblaz 27.0 Záhorská Ves
12 Štokrzí 49.0–50.0 Gajary
13 Bu#any 61.0 Moravský Sv. Ján
14 Lantov 62.0–64.0 Moravský Sv. Ján
15 Pod Devínom 0.5 Devínska Nová Ves
STUDY AREA
Our 15 study localities (Table 1) are situated between the Moravský Svätý Ján village and the Devín village, or
river km 66.0 and km 0.5, respectively, in the Záhorie lowland, Slovakia. They can be divided according to their
distance from the main stream (sensu Roux et al. 1982) into three groups:
1. The former meanders of the Morava River (Localities 1, 2, 3, 4 and 5). They were permanently connected
with the main channel during high discharge and correspond to the parapotamon type according to Roux et. al
(1982) and Ward et al. (1995). Their depth is about 2 m. Locality 1 is in the advanced stage of terrestrialisation
(Št!rba et al. 1995). Its littoral zone contain floating macrophytes (Spirodela polyrrhiza, Lemna sp.) and
submersed and emergent macrophytes (Ceratophyllum demersum and Glyceria sp.), respectively.
2. Dead meanders of the Morava River (Localities 6, 7, 8 and 9) and meadow pool (15). Localities of this
group are connected to the main channel only during inundation of the floodplain. They correspond to plesiopotamon
of Roux et al. (1982). These localities can be further divided into subgroups:
(2a) Relatively deep meanders (Localities 6–9). Their length remarkably exceeds their width and the maximum
depth reaches about 2 m. Macrophytes Ceratophyllum demersum, Polygonum amphibium, Potamogeton
sp., Nuphar lutea, Phragmites australis, Spirodela polyrrhiza and Lemna minor are common in this subgroup.
(2b) A permanent floodplain meadow pool (Locality 15). It has a round shape, and maximum depth of 1 m. The
pool is often dry in late summer and autumn. Its shore is overgrown by Phragmites australis.
3. Old meanders of Morava (Localities 10, 11, 12, 13 and 14). They are isolated from the main stream by a
main dam. They correspond to paleopotamon of Roux et al. (1982). Old meanders can be divided into subgroups:
(3a) Shallow temporary pools (Localities 11, 12, 13 and 14). Their depth is less than 1 m and the length exceeds
width. The maximum water level is achieved during the spring, or autumn. Prevailing vegetation at these localities
contains Rorippa amphibia, Schoenoplectus lacustris, Glyceria maxima and Typha sp. Tn the locality 13 at
Bu#any Hottonia palustris and Stratiotes aloides occured, until the pool became cattle crossing. When the water
became turbid and smelled of liquid manure, the vegetation contained only Spirodela and Lemna.
(3b) Relatively deep pool (Locality 10). It has an oval shape and 2.5 m depth. The original meander was most
probably deepened by the gravel excavation. Its littoral zone contains Ceratophyllum demersum, Batrachium
sp., Rorripa amphibia, Carex gracilis and Schoenoplectus sp.
METHODS
Samples of the cladoceran and copepod assemblages were taken using plankton net with 130–140 µm mesh. They
were preserved in 4% formaldehyde. Medial zone samples were taken using vertical tows from boat or oblique tows
from bank moved from bottom to surface. Littoral zone samples were taken as well.
206

Altogether 271 samples were analysed.
Cladocera samples were collected from the medial and littoral zone at 34 sampling sites of 12 localities (1–9,
12, 13 and 14) during August–September, 1994.
Altogether 105 oxbow samples were taken at 15 sampling sites of localities 1–5 as part of the meander
revitalisation project of Št!rba et al. (1995) during 1994–95. Both Cladocera and Copepoda were studied.
Samplings took place during May, July, August, October and December in 1994 and February and May in 1995.
In total, 75 samples of Cladocera and Copepoda were collected from the littoral zone of 5 oxbows during 1995.
They were taken at 5 sampling sites of localities 1–5. Samples were collected as part of the project of Lisický et
al. (1997). Sampling took place in May, July and October.
Altogether 70 Cladocera and Copepoda samples were collected from the medial and littoral zone of 10
localities during 2001. They were taken at 20 sampling sites of localities 6–15. Samples from localities 6, 9, 10
and 13 were collected during May–October. Locality 7 was sampled during May, June, September and October.
Locality 11 was sampled during May and July–September. Localities 12 and 14 were sampled during May,
September and October and Locality 15 was sampled during May–July. Locality 8 was sampled only once in May.
RESULTS
In total, 58 Cladocera species and 38 Copepoda taxa were found in the study area (Tabs 2 and 3).
They belong to three different types of environment: (1) former meanders of the Morava River, (2)
dead meanders and meadow pool inside-dikes, and (3) old meanders and oxbow outside-dikes.
The former meanders of the Morava River
The highest number of Cladocera species, 46 taxa, was found in side arms. They represent 79% of
all species found. The species Bosmina longirostris, Moina micrura and M. weismanni prevailed
in the medial zone. They occurred in mass at some localities. Other pelagic species Daphnia
galeata, D. ambigua, D. longispina and D. parvula were present almost at all localities. Littoral
Cladocera fauna was also abundant. Sida crystallina, Pleuroxus aduncus, Graptoleberis testudinaria
were the most frequent (Table 2). The Copepoda taxocoenosis consisted of 28 taxa, amounting
to 74% of all Copepoda species found. Eudiaptomus gracilis, Diaptomus castor and cyclopoids
Acanthocyclops robustus, A. vernalis and Cyclops vicinus were the most frequent ones
among pelagic calanoids species. Eucyclops serrulatus and Macrocyclops albidus prevailed
among phytophilous species.
Cladocera Bosmina coregoni, Pleuroxus denticulatus, Daphnia parvula, D. ambigua, Moina
weismanni and the calanoid Eurytemora velox were found from invaders.
Dead meanders and meadow pool inside-dikes
We found 36 Cladocera species in this arm type. There were species of genera Diaphanosoma,
Daphnia, Moina and Bosmina longirostris present in this environment. The meadow pool Pod
Devínom (Loc. 15) forms the exception. It recorded a significant prevalence of littoral species
Daphnia curvirostris, Megafenestra aurita, Simocephalus vetulus and S. congener (Table 2). The
Copepoda taxocoenosis included 20 species. Invaders Daphnia ambigua and Eurytemora velox
were found.
Old meanders and oxbow outside-dikes
In this arm type we found 36 Cladocera and 22 Copepoda. Littoral species prevailed (81%) in the
Crustacea taxocoenosis. The typical species were Daphnia curvirostris, Mixodiaptomus kupelwieseri
and Megacyclops viridis. Other common species were Daphnia pulex, D. pulicaria, Oxyurella
tenuicaudis, Canthocamptus staphylinus, Paracyclops poppei and Attheyella (B.) trispinosa.
From invader species only E. velox was found in the arm Bezodné (Loc. 10).
207

Tab. 2. Species composition and presence (+) of cladocerans (Crustacea, Cladocera) of water bodies of the Morava
river floodplain in 1994–1995 and 2001
locality number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
taxon
Ctenopoda
Sididae
Sida crystallina (O. F. Müller, 1776) + + + + + + + + +
Diaphanosoma brachyurum (Liévin, 1848) + + + + + +
Diaphanosoma mongolianum Ueno, 1938 +
Diaphanosoma orghidani Negrea, 1982 + + + + +
Anomopoda
Bosminidae
Bosmina coregoni Baird, 1875 + +
Bosmina longirostris (O. F. Müller, 1776) + + + + + + + + + + + + +
Chydoridae
Eurycercus lamellatus (O. F. Müller, 1776) + + + +
Pseudochydorus globosus (Baird, 1843) + + + +
Chydorus sphaericus (O. F. Müller, 1776) + + + + + + + + + + + + + +
Graptoleberis testudinaria (Fischer, 1848) + + + + + + + + + +
Pleuroxus truncatus (O. F. Müller, 1776) + + + + + + + +
Pleuroxus laevis Sars, 1862 + +
Pleuroxus trigonellus (O. F. Müller, 1776) +
Pleuroxus aduncus (Jurine, 1820) + + + + + + + + + + + + + +
Pleuroxus denticulatus Birge, 1879 +
Leydigia leydigii (Schoedler, 1862) + + + +
Dunhevedia crassa King, 1853 + +
Alonella nana (Baird, 1843)
Alonella excisa (Fischer, 1854) + + + + + + + +
Alonella exigua (Lilljeborg, 1853) + +
Alona guttata G. O. Sars, 1862 + + + + + + + + +
Alona costata G. O. Sars, 1862 +
Alona rectangula G. O. Sars, 1862 + + + + + + + + +
Alona quadrangularis (O. F. Müller, 1776) + +
Alona affinis (Leydig, 1860) + + + +
Oxyurella tenuicaudis (Sars, 1862) +
Tretocephala ambigua (Lilljeborg, 1900) + + +
Acroperus harpae (Baird, 1836) + + + +
Acroperus neglectus (Lilljeborg, 1900) + + + + + +
Daphnidae
Ceriodaphnia reticulata (Jurine, 1820) + + + + + + + + +
Ceriodaphnia pulchella G. O. Sars, 1862 + + + + + + + + + + +
Ceriodaphnia megops G. O. Sars, 1862 + + + + +
Ceriodaphnia laticaudata P. E. Müller, 1867 + +
Ceriodaphnia rotunda Sars, 1862 + + +
Ceriodaphnia quandrangula (O. F. Müller, 1785) + + + + + +
Simocephalus serrulatus (Koch, 1841) +
Simocephalus vetulus (O. F. Muller, 1776) + + + + + + + + + + + + + + +
Simocephalus exspinosus (Koch, 1841) + + + + + + + +
Simocephalus congener Schoedler, 1858 + + + + +
Daphnia pulex Leydig, 1860 +
Daphnia pulicaria Forbes, 1893 + +
Daphnia curvirostris Eylmann, 1887 + + + +
Daphnia parvula Fordyce, 1901 + + + +
Daphnia ambigua Scourfield, 1946 + + + + + +
Daphnia longispina O. F. Müller, 1785 + + + + + + + + + + +
208

Tab. 2. continuation
locality number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
taxon
Daphnia galeata G. O. Sars, 1863 + + + + + + + + +
Daphnia cucullata G. O. Sars, 1862 + + + + +
Megafenestra aurita (Fischer, 1849) + + + + +
Scapholeberis rammneri
Dumont et Pensaert, 1983 + + + +
Scapholeberis mucronata (O. F. Müller, 1776) + + + + + + + + + + + + +
Moinidae
Moina micrura Kurz, 1874 + + + + + + + +
Moina weismanni Ishikawa, 1896 + + + + + + + + + +
Macrothricidae
Ilyocryptus sordidus (Liévin, 1848) + +
Ilyocryptus agilis Kurz, 1878 + + + +
Macrothrix laticornis (Jurine, 1820) + +
Haplopoda
Leptodora kindtii (Focke, 1844) + +
DISCUSSION
A relatively high number of Cladocera (58 taxa) and Copepoda (38 taxa) found in the area between
the 66 th and 0,5 th river kilometre confirms that the Morava floodplain is a relatively undisturbed
biotope (see Hudec 1999). The occurrence of genus Daphnia at our localities corresponds to their
natural habitats and the present status of their distribution (Hrbá#ek 1987). The Cladocera taxocoenosis
represents 60% of the 96 cladocerans species (Hudec 1998) occurring on the Slovak
territory. The number of planktonic crustaceans species found in the alluvium of Morava (96 taxa)
appears to be high also in comparison with findings of other authors who examined biotopes of
other inundation areas. This number is a little higher than 80 taxa of Terek & Obrdlík (1992), found
in several habitats of the Rhine River. It is also higher than 50 planktonic crustacean species of
Gulyás (1994), found in waters of the Sigetköz arms of the Danube inundation in Hungary.
Vranovský (1997) determined 30 Copepoda taxa from five Danube arms and two main stream
profiles on the Slovak territory. The harpacticoid Nitocra hibernica (Brady, 1880) was one of the
most dominant species. It was not recorded from the Morava alluvium. However, 80% of copepods
that Vranovský (1997) had found in Danube arms were found also in Morava arms.
During the extensive recent faunistic research of ten Danubian side arms, 72 Cladocera and 25
Copepoda taxa were found between Dobrohoš1 and "í#ov villages (river km 1841–1081) (Illyová
1996). Danubian arm species, which were not found in the Morava floodplain, are the rare cladoceran
Anchistropus emarginatus Sars, 1862 and the benthic cladocerans Disparalona rostrata (Koch,
1840), Macrothrix hirsuticornis (Norman et Brady, 1867) and Pleuroxus uncinatus (Baird, 1850).
Adámek & Sukop (1992) found approximately the same number of Cladocera (53 taxa) and
Copepoda (41 taxa) in Morava waters to the north of our study area.
Part of the Morava floodplain, namely the confluence of Morava and Dyje rivers, was studied
in the project on aquatic invertebrates in the biosphere reserve Pálava in Czech Republic (Opravilová
et al. 1999). Sukop & Kopecký (1999) found 29 Cladocera species in this area. We found all of them
in our study area, except of Daphnia magna Straus, 1820. Only Eudiaptomus vulgaris (Schmeil,
1898) from 17 copepod taxa determined by Kopecký et al. (1999) was not present in our study area.
209

Tab. 3. Species composition and presence (+) of copepods (Crustacea, Copepoda) of selected habitats of Morava
river floodplain in 1994–1995 and 2001
locality number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
taxon
Calanoida
Temoridae
Eurytemora velox (Lilljeborg, 1853) + + +
Diaptomidae
Arctodiaptomus bacillifer (Brady, 1880) + +
Diaptomus castor (Jurine, 1820) + + + + +
Eudiaptomus gracilis (Sars, 1863) + + + + + + + + + + +
Eudiaptomus zachariasi (Poppe, 1886) + + + +
Mixodiaptomus kupelwieseri (Brehm, 1907) + + + +
Cyclopoida
Cyclopidae
Eucyclopinae
Macrocyclops fuscus (Jurine, 1820) + +
Macrocyclops distinctus (Richard, 1887) +
Macrocyclops albidus (Jurine, 1820) + + + + + + + + + + +
Eucyclops macrurus (Sars, 1863) +
Eucyclops speratus (Lilljeborg, 1901) + + + + +
Eucyclops serrulatus (Fischer, 1851) + + + + + + + + + + + +
E. serrulatus var. proximus (Lilljeborg, 1901) +
Tropocyclops prasinus (Fischer, 1860) +
Mesocyclops leuckarti (Claus, 1857) + + + + + + + + +
Paracyclops fimbriatus (Fischer, 1853) + +
Paracyclops poppei (Rehberg, 1880) +
Cyclopinae
Metacyclops gracilis (Lilljeborg, 1853) + +
Cryptocyclops bicolor (Sars, 1863) + + + + +
Microcyclops varicans (Sars, 1863) + + +
Megacyclops viridis (Jurine, 1820) + + + + + + + +
Megacyclops gigas (Claus, 1857) + +
Megacyclops latipes (Lowndes, 1927) + +
Acanthocyclops robustus (Sars, 1863) + + + + + + + + + + + +
Acanthocyclops vernalis (Fischer, 1853) + + + + +
Diacyclops bicuspidatus (Claus, 1857) + + + + + + +
Diacyclops bisetosus (Rehmberg, 1880) + + + +
Cyclops furcifer Claus, 1857 + + + +
Cyclops insignis Fischer, 1857 +
Cyclops strenuus Fischer, 1851 + + + + + + + + +
Cyclops vicinus Uljanin, 1875 + + + + + + + + + +
Thermocyclops crassus (Fischer, 1853) + + + + + + + + + + + +
Thermocyclops dybowskii (Lande, 1890) +
Thermocyclops oithonoides (Sars, 1863) + + + + + + + + + +
Ectocyclops phaleratus (Koch, 1838) + +
Harpacticoida
Attheyella (B.) trispinosa (Brady, 1880) +
Bryocamptus (B.) minutus (Claus, 1863) + + + +
Canthocamptus staphylinus (Jurine, 1820) + + + + + + + + + + + +
We assume that invaders drifted into the Morava inundation either from the South Moravia
lenitic biotopes (D. parvula, D. ambigua, M. weismanni and B. coregoni) or expanded from the
Danube inundation area (P. denticulatus and E. velox); both alternatives are possible. For example,
the species Bosmina coregoni probably drifted from the Nové Mlýny dam where is was
210

determined by Adámek & Sukop (1992). However, the species could migrate also from the Danube
River where Vranovský (1974) and Hudec (1989) found it in the main stream and "í#ov Arm,
respectively. The direction in which the species Daphnia parvula migrated into the Morava
alluvium remains questionable. The species was frequent in Morava side arms in 1994–1995 and
occurred also in the K%ivé jezero lake plankton (Sukop & Kopecký 1999) in the South Moravia
region. Other authors do not mention Daphnia parvula in the Morava alluvium. It has not been
found in the Danube inundation either.
Hudec (1998) assumes that Daphnia parvula migrates to Slovakia from the Morava and Danube
river basins similarly as B. coregoni and D. ambigua. Invader cladocerans could get to this
region from the water systems of the South Moravia. This could relate to the fact that water birds,
potential carriers, migrate in larger numbers than before after the construction of the Nové Mlýny
Dam. The species Eurytemora velox penetrated into the Morava inundation most probably from
the Danube where it was the dominant species in mid nineties at several localities (Vranovský
1997). The species Pleuroxus denticulatus spread in the Danube inundation with the same velocity.
It was recorded for the first time in 1992 (Terek 1997) but it occurred almost in the whole Slovak
Danube section in mid nineties (Hudec & Illyová 1998).
Crustaceoplankton taxocoenoses in various types of arms, oxbows and temporary pools reflected
different character of these environments. Pond assemblage prevailed in water bodies with
the stable water level. Small Cladocera species and small-size plankton (unpublished data) indicated
high densities of planktivorous fish (Hrbá#ek 1962). Phytophilous crustaceans inhabited the
littoral overgrown with macrovegetation. The assemblage of phytophilous and benthic species,
Tretocephala ambigua, Daphnia curvirostris, Mixodiaptomus kupelwieseri, Megacyclops viridis,
Paracyclops poppei, Thermocyclops dybowskii, Attheyella (B.) trispinosa, in old meanders and
pools with abundant macrovegetation occurred. Kopecký & Koudelková (1997) found the similar
species composition in two pools of the Morava River floodplain. The crustaceans taxocoenosis
(Cyclops strenuus, Mixodiaptomus kupelwieseri and Daphnia curvirostris), found in periodic
pools and temporary arms, corresponds to associations described from the Záhorie and Podunajská
lowlands (Št!rba 1988), with exception of missing D. magna.
Protection of the original aquatic ecosystems undisturbed by anthropogenic impact is the
inevitable condition of preservation of aquatic fauna biodiversity in the Morava inundation. The
uniqueness of this area is supported by the presence of rare species such as Hemidiaptomus
hungaricus Kiefer, 1933 or Heterocope saliens (Lilljeborg, 1863) in the Záhorie lowland (Brtek
1953). Other crustaceans worthy of protection are Lepidurus apus Linnaeus, 1758 and Siphonophanes
grubii (Dybowski, 1860) occurring in snow and flood pools (Lukáš 2000).
A c k n o w l e d g e m e n t s
The research was supported by the grant No. 1/8200/01 from the Slovak Grant Agency for Science VEGA.
Authors thank to Igor Hudec for the provision of his determined Cladocera material from project of Otakar
Št!rba during 1994–95, and to Otakar Št!rba for his kind permission to use data from the unpublished report. We
are grateful to reviewers for their constructive criticism of an earlier version of this paper. We also thank Michal
Nem#ok for his linguistic help with the English manuscript.
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212

Acta Soc. Zool. Bohem. 66: 213–220, 2002
ISSN 1211-376X
Moina (Crustacea: Anomopoda, Moinidae) in the Czech Republic: a review
Adam PETRUSEK
Department of Hydrobiology, Charles University, Vini#ná 7, CZ–128 44 Prague 2, Czech Republic;
e-mail: petrusek@cesnet.cz
Received April "0, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. Current state of knowledge about general characteristics and ecology of the Central European
species of Moina Baird, 1850 is summarised. Five species present in the Czech Republic or in the immediate
vicinity can be divided into two groups according to their ecology: Two of them – Moina macrocopa
(Straus, 1819) and M. brachiata (Jurine, 1820) – have larger body size and they are common in small
ephemeral localities or highly eutrophic village ponds in the lowlands of the whole country, whereas
M. micrura Kurz, 1874 and M. weismanni Ishikawa, 1896 are generally smaller and live in larger permanent
waters. M. micrura is a native species, described from a locality in Central Bohemia. M. weismanni is likely
a Far East invader but it has been present in the region for several decades already, at least since early
1930’s. The last species, M. ephemeralis Hudec, 1997, was described from a fishpond in Slovakia only very
recently; however, it had been probably overlooked in the past due to the similarity of parthenogenetic
females as well as usage of misleading identification keys.
Ecology, review, Cladocera, Moina, Czech Republic, Palaearctic region
Moina Baird, 1850 (Crustacea: Anomopoda: Moinidae) is a cladoceran genus closely related to
Daphniidae (Goulden 1968, Fryer 1995). They are opportunistic cladocerans typically occurring in
temporary pools, saline lakes or other waters with extreme conditions (e. g., with high temperature
fluctuations). The genus has a worldwide distribution (except Antarctica). The majority of the
species can be found in areas with a relatively warm climate, from tropical to temperate zones.
There are only very few exceptional reports of moinids from subarctic or alpine regions (Šrámek-
Hušek 1962, Flössner 2000).
All Moina are small or medium-sized cladocerans. The body length of adult parthenogenetic
females varies between 0.5 and 1.8 mm, depending on the species. Males are always smaller than
females. The appearance, functional morphology and presumed phylogeny of Moina are described
in details in the monograph by Goulden (1968). In this complete revision of the genus, the
author pointed out some specific morphological characteristics (e. g., morphology of trunk limbs,
head shape, antennules) and peculiarities in sexual cycle and development of Moina. In his opinion,
these specific characters were important enough to separate two genera, Moina and Moinodaphnia
Herrick, 1887, from the rest of Daphniidae. He assigned these genera to the newly described
family Moinidae. This separation of the family has been recently subject to criticism as a
result of both morphological analyses (e. g., Fryer 1995) and DNA studies (Schwenk et al. 1998).
However, additional sequence data from three Moina species suggest that this genus is divergent
from other daphnid genera and may constitute a well-defined group (Petrusek, unpublished data).
Several species of Moina have considerable economic importance. Most of the species have a
short generation time, fast population growth and they are easy to keep in culture. There is an
This contribution is dedicated to Doc. RNDr. Jaroslav Hrbá!ek DrSc., upon the occasion of the 80th anniversary
of his birthday ("2th May "92").
213

increasing interest in production of Moina as a cheap and nutritionally valuable fish food for
aquaculture (e. g., Alam et al. 1993, Tamaru et al. 1997). Moina is also routinely used as a test animal
in toxicological assays (e. g., Krishnan & Chockalingam 1989, Wong 1989). In addition to wellknown
Daphnia Müller, 1785, Moina has been used as a convenient model organism for studies of
cladoceran physiology, ontogenesis, genetics etc. The findings about its biology formed substantial
parts of some influential “classical” papers (e. g., Weismann 1877, Banta 1939).
Special features of the sexual cycle of Moina
Moina reproduces by the cyclic parthenogenesis as most of cladocerans do. The typical reproductive
cycle of daphniid cladoceran (and its impact on the genetic structure of populations) is
described and discussed in many reviews (e.g., Mort 1991, De Meester 1996). Although Moina
reproduction follows the same general pattern as Daphnia, there are some differences arising from
peculiarities in its ontogeny.
The parthenogenetic females of Moina possess a special placenta-like structure (“Nährboden”
sensu Weismann 1877; “marsupial organ” sensu Smirnov 1976) that presumably serves to nourish
the developing embryos in the brood pouch (Weismann 1877, Rammner 1933, Goulden 1968).
Organs with a similar function have independently developed also in other cladoceran taxa – in all
families of Onychopoda and in Penilia avirostris Dana, 1849 (Ctenopoda: Sididae) (Makrushin
1985). The yolk content of parthenogenetic eggs is reduced in the taxa possessing placenta
(Rammner 1933, Makrushin 1985) and the eggs are not able to develop out of the brood pouch of
the mother. This adaptation could possibly lead to increased brood sizes and/or faster egg production.
No further details on function and physiology of the placenta of moinids are available.
The presence of the placenta apparently prevents the parthenogenetic females of Moina from
forming of an ephippium. This slightly changes the sexual phase of the reproductive cycle of
Moina. Although any parthenogenetic female can switch to sexual reproduction in Daphnia or
other Anomopoda, it is not the case of Moina. Males can develop in a brood of any parthenogenetic
female of Moina but the sexual eggs and ephippia can only be produced by freshly matured
females (Goulden 1968).
The sexual females of Moina can continue to produce sexual eggs and ephippia throughout
their life. The new ephippium is formed immediately after the deposition of the previous one and
the females are ready for the next mating. This allows one female to produce several genetically
diverse resting eggs during the sexual phase of reproduction, even in species where only one
sexual egg is deposited into the ephippium. However, once a sexual female switches to parthenogenetic
reproduction, this switch is irreversible.
One moult is necessary between the deposition of the ephippium and the first parthenogenetic
brood in at least several Moina species (Goulden 1968; pers. obs.). Goulden suggested that this
post-sexual instar was necessary for placenta formation.
Moina in Europe and in the Czech Republic
Eight species of Moina have been recorded from the European continent. Moina lipini Smirnov,
1976 was described from a fishpond area near Moscow (Russia) and there are no reports of this
species from Central or Western Europe.
Moina salina Daday, 1888 has been described from Transylvania and occurs in saline waters of
the Old World. (The name M. mongolica Daday, 1901 is still widely in use, however, it is only a
junior synonym of the same species (Negrea 1984).) In Europe it is present in Eastern European
lowlands and in the Mediterranean region – e.g., on the Balkan Peninsula (Stephanides 1948,
Negrea 1984), in Spain (Alonso 1996), and on Sardinia (Margaritora 1971). This species does not
214

occur in the Czech Republic, the nearest localities with its presence are in southern Hungary (Forró
1988, Hrbá#ek et al. 1978).
Moina affinis Birge, 1893 is a North American invader. It has been repeatedly recorded in Italian
rice fields in the last 40 years. First findings were published by Moroni (1962); for more recent data
see Margaritora et al. (1987) or Leoni et al. (1999).
The remaining five species of European Moina can be found on the territory of the Czech
Republic and Slovakia. These are Moina macrocopa, M. brachiata, M. micrura, M. weismanni,
and M. ephemeralis. The differential diagnosis of these species has been given in papers of
Hudec (1988, 1989, 1997).
Our species of Moina can be divided into two groups according to their preferred habitat and
corresponding ecological traits. The facts about their ecology and habitat preference given below
describe the situation in Central Europe.
Moina brachiata and Moina macrocopa – species of puddles and hypertrophic pools
Two species, Moina brachiata and M. macrocopa belong to a group of “large” Moina species –
their parthenogenetic females can grow up to 1.5 mm and their mean size is usually over 1 mm
(Goulden 1968). They are characteristic inhabitants of small, usually ephemeral water bodies in the
temperate zone. Both species prefer lower altitudes, they are rarely found higher than 450 m a.s.l.
in Central Europe (Hudec 1989, pers. obs.). They sometimes co-occur in the same water bodies
(Petrusek 2000).
Moina brachiata (Jurine, 1820) is the type species of the genus. It is an Old World species,
widely distributed in Europe, Africa and Central Asia (for details on distribution of all species, see
Goulden 1968, Smirnov 1976, Hrbá#ek et al. 1978, Flössner 2000).
Moina macrocopa (Straus, 1819)* is widespread mostly in Europe, North Africa (Algeria), Middle
East, Eastern Asia and North America. The populations from Paleotropic and Palearctic regions
do not differ substantially in their morphology. They belong to the typical form, designated by in
Goulden (1968) as the subspecies M. m. macrocopa (Straus 1819). There is only one isolated
report of M. m. macrocopa from the Neotropic region (Paggi 1997) and that could have been a case
of the recent species introduction.
In North America, M. macrocopa occurs in the United States (Goulden 1968) and Mexico (F.
Martinéz-Jerónimo, pers. comm). However, animals from these populations differ from the typical
form by the pattern of ephippial surface and setation of the carapace rim. They were described as
the new subspecies M. m. americana Goulden, 1968. The actual extent of the divergence between
the two subspecies is not known yet.
There are two basic types of habitats strongly preferred by M. brachiata and M. macrocopa.
Small village ponds or sewage ponds are examples of the first type. These are usually permanent
water bodies, highly eutrophic and with excess of food supply. However, the conditions during the
season become unfavourable for most of the zooplankton species. The water temperatures can be
relatively high in summer and there are often periods of low oxygen content (especially if the water
surface is covered with floating plants such as Lemna, or detritus). Although fish may be present,
their predation pressure is not very important factor structuring zooplankton communities in those
ponds. Moina usually co-occurs with some large Daphnia species (D. magna Straus, 1820, D. pulex
Leydig, 1860) in such habitats (Šrámek-Hušek 1941).
* Alonso (1996) suggested that the name Moina macrocopus should be used instead because of linguistic and
historical reasons. Although this suggestion follows the ICZN rules, I still use the name M. macrocopa. The taxon
is widespread and the name macrocopa has been in use continuously for more than ninety years. I believe it should
be assigned the status of nomen conservandum to avoid confusion.
215

Ephemeral pools (such as rainpools or ditches; further on called “puddles”) are examples of the
second habitat characteristic for Moina brachiata or M. macrocopa. Typical puddle is a small
water body, with a diameter of one to several meters and maximum depth often not exceeding 20 cm,
often drying up several times during the summer season. The volume and duration of water is
dependent mostly on the rain frequency and exposition to the sunlight and wind. It is usually
either dry or completely frozen during the winter. Puddle localities are unstable, they can deteriorate
and/or disappear completely during few seasons; so the Moina population does not find
suitable conditions at the site anymore.
Ephemeral habitats inhabited by M. macrocopa and M. brachiata are usually found in open,
unshaded areas (pers. obs.), and therefore they are often considerably warmed up during the day.
The temperature differences between day and night in puddles can rise up to 15° C (Maier et al.
1998). Water level fluctuations in puddles induce abrupt changes of both biotic and abiotic conditions
– salinity, seston (and food) concentration etc. (Hronešová 1992). Changes of water volume
also directly affect the density of populations.
Puddles often harbour a high diversity of invertebrates, including other cladocerans (e.g.,
Daphnia obtusa Kurz, 1875), ostracods, copepods and insects (pers. obs.). The most important
predators possibly affecting Moina populations in this type of habitat probably belong to the
latter two groups. Other crustaceans typical for ephemeral habitats, Anostraca, Notostraca, and
Conchostraca, may also co-occur with Moina (Maier et al. 1998, pers. obs.).
Puddle species of Moina can tolerate high water temperatures (Maier 1993) and elevated content
of dissolved substances. This tolerance allows them to colonise water bodies where the
extreme environmental conditions limit the presence of their potential zooplankton competitors
(e.g., cattle manure runoff with high organic content). M. brachiata is also one of few cladoceran
inhabitants of sodic waters in Austria, Hungary and Serbia, occurring both in temporary pools and
permanent “lakes” (Metz & Forró 1989). Similar saline habitats are uncommon in the Czech Republic.
One such locality (with an abundant presence of M. brachiata) is a swamp adjacent to the
fishpond Nesyt near Lednice in South Moravia (pers. obs.).
Both M. brachiata and M. macrocopa can reach densities over 10,000 ind×l –1 (Maier 1993,
Khalaf & Shihab 1979) in favourable conditions. The highest abundance usually corresponds with
periods of low water level in temporary pools, when population growth and drying out of the
habitat have synergistic effect. Such crowding lowers dramatically the available food supply,
consequently the decrease of ingestion stops population growth and encourages sexual reproduction
(D’Abramo 1980), preparing the Moina population for possible drying of the habitat.
Each of the two puddle species can be found occasionally in fishponds. This occurs in periods
when conditions in the pond approach those experienced in the temporary waters – for example
within few weeks after the flooding of the area. The population of Moina can reach a brief peak of
abundance in such cases. However, neither M. brachiata nor M. macrocopa regularly occur in
permanent water bodies with the presence of fish or large Daphnia species.
Moina micrura, M. weismanni and M. ephemeralis – species of ponds
These three species are representatives of smaller cladocerans of the “shallow lake” habitat and
they share together many characteristics. Localities where these Moina species occur include
fishponds, dead river arms and oxbow lakes, flooded sandpits and similar water bodies (Hudec
1988, 1997, I. P%ikryl – unpublished data), especially in warmer areas. Planktivorous fish commonly
prevail in such waters and shift the composition of crustacean zooplankton towards small cladocerans.
Adult females of all three Moina species from permanent water bodies have relatively small
body size, usually not exceeding 1 mm. M. ephemeralis is larger than the other two species – the
216

largest specimens measured up to 1.36 mm (Hudec 1997). Their populations are intermittent and no
adults survive over winter. Populations of Moina often appear in short-time peaks of abundance
only, usually in spring or autumn (Hudec 1997, I. P%ikryl – unpublished data). M. micrura can
occasionally reach densities up to 8,000 indx.l –1 during these peaks (Flössner 2000). If prevailing
for longer time, Moina populations have usually low densities and occur together with other
genera of small pelagic cladocerans (e.g. Bosmina Baird, 1846, some Daphnia species, Diaphanosoma
S. Fischer, 1850 etc.). Two or even all three species can be occasionally found together on the
same locality (Hudec 1997, Petrusek 2000).
Moina micrura Kurz, 1874 was until recently the only species of Moina with a small body,
recognised from the Central European region. It is a typical example of the unresolved taxonomy
within the genus. It was described from a pond near Malešov (near Kutná Hora, Central Bohemia).
The type specimens are not available and the type locality no longer exists – the pond had been
dried up and the area forested. The original description did not include enough details and probably
was not widely known; so several other similar species have been described during following
decades from different parts of the world. More information about morphology of M. micrura
sensu stricto was added later in the report of the presence of this species in Bohemia (Šrámek-
Hušek 1940). However, the detailed redescription of M. micrura from another Czech locality remained
unpublished (Šrámek-Hušek, unpublished manuscript). The morphological variability of
populations of small-sized Moina is so high that seven species names were declared invalid and
synonymised with M. micrura in the revision of the genus by Goulden (1968); some species were
included also later (Smirnov 1976, Sharma & Sharma 1990).
Moina micrura is considered to be a cosmopolitan species with extensive morphological and
ecological plasticity, occurring in a wide range of different habitats. M. micrura sensu lato (Goulden
1968) has been recorded in various habitats in most parts of the world, with the exception of cold
regions. Partly because of acknowledged variability of the species, any population of small Moina
found on the territory of former Czechoslovakia had been previously identified as M. micrura or
invalid M. rectirostris (Leydig, 1860). The results of the experimental hybridisation of clones of
M. micrura sensu lato of different origin and the sequence comparison of the mitochondrial gene
for 12S rRNA show that M. micrura is clearly a complex of sibling species (Petrusek 2000). However,
the Central European populations belong undoubtedly to Moina micrura sensu stricto.
Two more species with similar morphology and ecology, M. weismanni and M. ephemeralis,
occur in Central Europe. The most important species-specific characters can be found on the
sexual individuals. The morphology of ephippial females allows an easy identification of the
species. Small differences in morphology of parthenogenetic females can explain the fact that
M. weismanni (and probably also M. ephemeralis) often escaped the detection. When a sample of
the population does not contain sexual animals, the correct species assignment can be a difficult
task. M. weismanni and M. ephemeralis are absent from most of the identification keys of European
cladocerans (including recent publications – e.g., Flössner 2000). The use of older publications
(e.g., Šrámek-Hušek 1954, Šrámek-Hušek et al. 1962) that include invalid species names
(M. rectirostris) and incorrect descriptions of valid species (M. brachiata) still adds to the taxonomical
confusion.
Moina weismanni Ishikawa, 1896 is thought to be a Far East invader, brought to Europe with the
cultivation of rice (Margaritora et al. 1987, Hudec 1990). It is widespread in South-East Asia –
Japan, China, Indonesia, and Indian subcontinent (Smirnov 1976). Since the differential diagnosis
of the species (e.g., in Hudec 1990) had become widely known, it has been recorded in various
localities in both the Czech Republic and Slovakia (Hudec 1988, I. P%ikryl – unpublished data).
217

However, Moina weismanni has been apparently a part of the Central European fauna for much
longer than previously thought. The occurrence of this species in Bohemia is documented back to
1931. There is a well-preserved sample of a rich M. weismanni population, including males and
ephippial females, collected by Šrámek-Hušek in a village pond in Bohumile# (Pardubice district)
on August 14, 1931. The animals were identified as M. rectirostris (invalid name, which had been
used for several different Moina species) (Šrámek-Hušek 1941), therefore the presence of this
species in the old sample remained unnoticed until recently (V. Ko%ínek, unpublished results).
Vojtek (1958) recorded the presence of M. weismanni in Southern Moravia (four localities near
Znojmo) in the years 1947–1948. Although he called the animal M. micrura, he described well the
morphology of sexual females. The identity of the species is beyond question thanks to his precise
drawing of the ephippial surface.
The presence of this species has been also recorded in several areas of Central Asia and Russia
(Mirabdullaev 1992), on the Balkan peninsula (Petkovski 1991), in Hungary (Hudec 1990, Forró
1999), and in Belgium (Forró et al., in prep.). It seems that its current distribution is much wider than
previously thought. Either M. weismanni is quite an efficient invader, or its presence had been
overlooked for a long time. The detailed comparison of European and Far East populations of
M. weismanni, including molecular traits, would be helpful to validate the identity of this species
and the relationship among its populations.
Moina ephemeralis Hudec, 1997 is the rarest species of Moina living on the territory of the Czech
Republic and Slovakia. It was described recently from Southern Slovakia (carp ponds near Hrhov)
(Hudec 1997). Since then it has been found on several occasions in the floodplain of the lower
reach of the river Morava (I. Hudec, pers. comm.). The distribution of this species is unknown;
there are yet no records of this species from any other part of Europe.
Although Moina ephemeralis is one of only three European Moina species with ephippium
with two eggs (the other two are M. macrocopa and M. lipini), there are several reasons why it has
probably escaped attention for a long time. First, the life cycle of M. ephemeralis is very short
compared to other planktonic cladocerans (Hudec 1997), second, the correct identification of
populations without sexual individuals is rather difficult, as parthenogenetic females can be confounded
with those of M. weismanni. And the last, the use of misleading identification keys could
be also the reason.
Four species of Moina are included in the monograph on Czechoslovak cladocerans (Šrámek-
Hušek 1962). M. weismanni and M. ephemeralis are missing, but one more species, M. rectirostris
(Leydig, 1860) is listed. Goulden (1968) synonymised most of the occurrences of this taxon with
M. micrura, however, M. rectirostris sensu Šrámek-Hušek is equivalent to M. brachiata. Šrámek-
Hušek distinguished M. brachiata and M. rectirostris by the number of ephippial eggs. According
to his key, M. brachiata should have had ephippium with two eggs (which is incorrect) whereas
M. rectirostris with one egg only; M. ephemeralis would be therefore keyed out as M. brachiata.
This suggests that some records of M. brachiata sensu Šrámek-Hušek could have been actually
rare occurrences of M. ephemeralis.
A c k n o w l e d g e m e n t s
I would like to thank Vladimír Ko%ínek, Igor Hudec, and László Forró for sharing information and valuable
discussions. The research of cladoceran diversity (incl. Moina) is supported by Grant Agency of the Charles
University (project no. 146/2001) and Ministry of Education of the Czech Republic (project no. MSM113100004).
218

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Acta Soc. Zool. Bohem. 66: 221–230, 2002
ISSN 1211-376X
Two recent immigrants into Czech aquatic habitats: Daphnia ambigua
and D. parvula (Crustacea: Cladocera)
Magdaléna ŽOFKOVÁ, Vladimír KO$ÍNEK & Martin "ERNÝ
Department of Hydrobiology, Faculty of Science, Charles University, Vini#ná 7,
CZ–128 44 Praha 2, Czech Republic
Received May 8, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. Available data on the distribution of Daphnia ambigua Scourfield, 1947 and D. parvula Fordyce,
1901 in Czechia were summarized. We compare interspecific differences in the size distributions among
these two invasive and a common native species D. galeata G. O. Sars, 1863 as well as locality-specific (6
Czech and 1 North American) body-size differences for each of the exotic species. Both invaders are able
to attain maximal body size about 1.3 mm, a substantial overlap with the native D. galeata. In one
reservoir, D. galeata population dropped the mean body size down to similar values as are common in D.
ambigua. Size differences among localities are not significant due to the wide local range of values. Size
distribution frequencies in a locality are mostly symmetric suggesting that input of young stages into the
population in the sampling dates studied was already diminishing. The populations of invader species differ
in the timing of sexual reproduction; gamogenetic populations of D. parvula occur from September to
November, while those of D. ambigua were found only in May.
Biological invasions, zooplankton, Daphnia ambigua, Daphnia parvula, Daphnia galeata, size
distribution, gamogenetic populations, Czech Republic, Palaearctic region
INTRODUCTION
Two species of the genus Daphnia Müller, 1785 widely distributed in North America, has been
recently reported spreading across Europe invading various kinds of water bodies: Daphnia
ambigua, originally described from a Kew Garden reservoir in London, and D. parvula, from a fish
pond about 5 km south of Arapahoe, Nebraska. Starting from 1970, the former species was occasionally
reported from water bodies in continental Europe: Amoros (1972) mentioned a short
occurrence of D. ambigua in one fishpond in S.-E. of France, unfortunately without any further
details or drawings. Well-documented records from Belgium were published by Dumont (1974),
followed by Flössner & Kraus (1976), and Margaritora (1985). Daphnia parvula seems to be less
frequently recorded. The species appears in recent review of Spanish cladoceran fauna (Alonso
1996), and one of the figures of postabdominal claw of D. ambigua in Margaritora (1985) may
belong to D. parvula. Schrimpf & Steinberg (1982) reported the species from waters in neighbourhood
of Mannheim, Germany, and Petkowski (1990) found D. parvula in Macedonia. Hudec (1990,
1991) published data on the distribution of both the species in Slovakia. Weiler (poster communication,
symposium in Plön, Germany, 1999) found both species in lakes in neighbourhood of Bonn
(Germany). D. ambigua was found recently in a system of interconnected ponds in Belgium
(Michels et al. 2001.) German localities of both the species are summarised in Flössner (2000). In
This contribution is dedicated to Doc. RNDr. Jaroslav Hrbá!ek DrSc., upon the occasion of the 80th anniversary
of his birthday ("2th May "92")
221

Czechia, the second author found D. parvula in late seventies (1977) in a drinking water reservoir,
and D. ambigua has been regurarly recorded only from 1994 (I. P%ikryl, samples and personal
communication 1998).
The centre of geographic distribution of both the species is on American continent (Brooks
1957), and European localities are supposed to be secondary as result of recent invasion of both
the species. Careful re-examination of samples from various waters in Czechia revealed relatively
frequent occurrence of both the species in habitats like reservoirs, fishponds and pools. Such
situation is interesting from the ecological point of view. Newly arrived species has to found a
vacant niche and be successful in inter-specific competition with native species with similar environmental
requirements.
The aim of the present study is to give an overview of populations living in our region. The data
will be used for the further study of their phylogeography and competitive abilities.
MATERIAL AND METHODS
Beside re-examination of some of the preserved material offered by our colleagues, we have sampled prospective
habitats as well as those from which one of the species has been already reported. North American material was
obtained due to the kind assistance of colleagues from Canada, USA, Mexico and Cuba. Most of the material was
collected with the 100-µm-mesh size plankton net and preserved with 4% formalin solution. The following table
lists all the populations, which were at our disposal for the study. We omitted habitats where the species occurred
only sporadically and in low numbers.
Daphnia ambigua Scourfield, 1947
MATERIAL EXAMINED. Závist Pond, Skali#any, Blatná, Sep. 22. 1999; Padr1 Pond, Drásov, P%íbram, Sep., 23. 1999;
Novokest%anský Pond, Strakonice, Sep. 22. 1999; Nový Pond, W. of "ekanice, Blatná, Sep. 22. 1999; (all M.
Žofková legit). Zlivský Pond, Hluboká, "eské Bud!jovice, June 12. 1996; Zámecký Pond, Lednice, May 22. Aug.
22. 1996; $ežabinec, Písek, Aug. 17. 1998; Domin Pond, Vrbno at "eské Bud!jovice, June 12. 1996; V!žický
Pond, W. of Rovensko, June 8. Aug. 14. 1995; Skalský Pond, Protivín, Aug. 13. 1996; Jordánek Pond, Hrn#í%e,
Sep. 10. 1996; Komorní Pond, Turnov, Sep. 22. 1998; Žabokor Pond, Mnichovo Hradišt!, Sep. 1. 1994;
Žehu0ský Pond, Pod!brady, July 10. 1996. (all I. P%ikryl legit). Sand pit “Bagr” in park, "eské Bud!jovice, Sep.
2. 1997. (J. Se3a legit), Jul. 31. Aug. 15. Sep. 2. 1997, Jul. 2. 1998, (H. Luk#íková legit.); Orlík reservoir, Zvíkov,
May 12. 1999 (L. R+ži#ka legit). $ímov reservoir, "eské Bud!jovice, May 24. 2001, (J. Machá#ek J. legit);
village pond, Divišovice at Sedlec-Pr#ice, Jun. 29. 2001 (V. Ko%ínek legit). Channel, Pohansko, B%eclav, S.
Moravia, Jun. 21. 1999 (M. Omesová legit). Backwater, river Morava, Vysoká, Slovakia, May 27. 1994. (O.
Št!rba legit).
Comparative material from North and Central America
MATERIAL EXAMINED. Grenadier Pond, Ont., Canada, June 6. 1966 (H. Harvey legit); Upper Bass lake, Ont., Canada, June
1. 1965 (F. Rigler’s collection Univ. Toronto); Kashe lake, Muskoka Co., Ont., Canada, July 31. 1978 (M.J. Dadswell
legit); Johnson lake, Florida, USA, Jan. 19. 1994 (W.M. Lewis junior legit), Jan. 28. 1987 (D.G. Frey legit); Feb. 23.
1988 (G. Deevey legit); fish pond #6, Apr. 9., Apr. 28. 1968; fishpond #15, Apr. 9. 1968, Auburn, Alabama, USA, (all
J. Hrbá#ek legit); Laguna #1, El Dique, Cuba, March 5. 1968 (N. Martinez legit); Laguna Eduardo, prov. Habana,
Cuba, June 17. 1990 (J. Fott legit); Presa los Canos, Mexico, June 17. 1990 (M. Briano legit).
Daphnia parvula Fordyce, 1901
MATERIAL EXAMINED. Ruda Pond, T%ebo0, May 23. 1996 (I. P%ikryl legit); Sep. 28. 1999 (M. Žofková legit);
Dušákovský Pond, T%ebo0, May 23. 1996 (I. P%ikryl legit); Sep. 28. 1999 (M. Žofková legit); Velký Tisý Pond,
T%ebo0, June 26. July 17. 1996; Nový Spálený Pond, T%ebo0, May 23. 1996; Žehu0ský Pond, Pod!brady, July 10.
1996; P%esli#kový Pond, Nové Hrady, Sep. 11. 1996; Sousedský Pond, Chlum at T%ebo0, Aug. 14. 1996; Kotvice
Pond, Odra river, Nový Ji#ín, Aug. 29. 1995, N. Moravia (all I. P%ikryl legit); Božkov Pond, Mnichovice, Prague,
July 23. 1987, Oct. 29. 1989 (M. Pražáková legit); Dvorský Pond, Zbiroh, Sep. 19. 1994, Nov. 27. 1994, Oct.
29. 1997; fish pond in Zbiroh, Sep. 18. 1994 (all J. Vávra legit); Žabokor Pond, Mnichovo Hradišt!, Sep. 23.
1998; Novokest%anský Pond, Strakonice, Aug. 20. 31. 1998; small pond near Pa%ezská Lhota, Ji#ín, Sep. 23.
1998; fishpond S. of Branná, T%ebo0, Oct. 1. 1997; Buzický Pond, Blatná, Sep. 14. 1997; Nový Pond near
222

Fig. 1. Daphnia ambigua Scourfield: a – adult female, b – adult male, c – postabdominal claw of female, d – adult
male – antennule.
223

Fig. 2. Daphnia parvula Fordyce: a – adult female, b – adult male, c – postabdominal claw of a female, d –
immature male – antennule.
224

"ekanice, Strakonice, Aug. 30. 1998; $ežabinec Pond, Písek, Aug. 30. 1998; forest pond, Libínské sedlo, Prachatice,
July 17. 1997; Rokytnický Pond, Hrubá Skála, Sep. 23. 1998; Pýcha Pond, Skali#any, Blatná, Oct. 4. 1998;
fishpond in Podkost, Sobotka, Sep. 23. 1998, (all V. Ko%ínek legit); Skalka Pond, Drásov, P%íbram, Oct. 31. 1997 (K.
Urbanová legit); Hostiva% reservoir, Prague, July 8. 1980 (J. Fott legit), Aug. 14., Sep. 18., Oct. 2. 1981 (all E.
Hrušková legit); Vrchlice reservoir, Kutná Hora, June 21. 1977, (V. Gottwaldová legit); backwater Dlouhá, Upper
Lužnice river, N. Ves, Suchdol, Sep. 2. 1992 (J. Hrbá#ek legit); $ímov reservoir, Oct. 8. 1997 (J. Se3a legit); Pond
Bi%i#ka, Hradec Králové, Aug. 31. 1998 (M. Dewetter legit). Village pond in Sedlce, "eské Bud!jovice, Oct. 15. 2000
(L. Stará legit). Pond Jejkal Dolní, Vranov nad Dyjí, Jun. 8. 2001; pond Jejkal Horní, Jun. 8. 2001; "ervený pond,
W of Znojmo, Jun. 7. 2001; fish pond NW of Ješetice, Votice, Jul. 2. 2001, (all V. Ko%ínek legit). Pastvisko pond,
B%eclav, Jun. 3. 1993; Ost%icová pool, Jul. 8. 1993; Panenská pool, Jul. 8. 1993, K%ivé jezero reserve, S. Moravia (all
O. Skácelová legit); backwater at Devínská Nová Ves, Slovakia, May 27. 1994 (O. Št!rba legit).
Comparative material from North and Central America
MATERIAL EXAMINED. Lake Max, Turtle Park, Manitoba, Canada, Aug. 7. 1969; Lake Winnipeg, Manitoba, Canada,
July 9. 1969, (K. Patalas legit); Heart Lake, Ontario, Canada, May 28., Aug. 6., Nov. 5. 1965, (all F. Rigler’s
collection, Univ. Toronto); Teapot Pond, Ont., Canada, May 28. 1961; Toussaint Lake, Ont. Canada, June 2.
1965, (C.H. Harvey legit); Lake #302, Erickson, Manit., Canada, Oct. 13. 1971, (V. Ko%ínek legit); Lynx Lake,
B.C., Canada, July 15. 1969; Shell Lake, B.C., Canada, July 3. 1960, (all T. Northcote collection, Univ. B.C.);
Acueducto Holguin, Cuba, Apr. 23. 1966. (M. Legner legit).
RESULTS
Morphology
Daphnia ambigua. Females of Central European populations are mostly without any spine on top
of head or the spine is only faintly suggested in sub-cuticle tissues. We have found only two
habitats where females had remarkably pronounced apical horn on top of head. Both the habitats
are fishless pools with Chaoborus Lichtenstein, 1800 larvae present. Ocellus is seldom pigmented.
Other characters, including those of males, were found similar to North American populations. In
some of the ponds, the species co-occurs with D. parvula and the only reliable differentiation has
to be based on the difference in pectens on the postabdominal claw (Fig. 1).
Daphnia parvula. There was no difference in the morphology of females and males between
populations from our region and those from North America (Fig. 2).
Habitats
Populations of D. ambigua dwell mostly in shallow fishponds or in pools and backwaters along
rivers. Recently it was found in one reservoir ($ímov). On the other hand D. parvula probably
started to invade first such relatively young habitats as are reservoirs not more than several
decades after their construction. Only in nineties we have found the species in many fishponds.
Biology
The peak of the population densities in D. ambigua is reached mostly in August and at the beginning
of September. Nevertheless the populations start to occur already from May. Gamogenetic populations
were found only in May. Occurrence of males and ephippial females seems to be rare, we have
found it only in 4 habitats (a dug out flooded with water, a fish pond, one river backwater and a
reservoir). Besides populations from American continent, we have checked one locality reported
outside the supposed Euro-American range of the species: Japan (Ueno1960). The population from
a high mountain lake Mikuriga-ike (Toyama ken, Honshu, altitude 2405 m, Sep. 25. 1978, K. Okamoto
legit) belongs certainly to the “Daphnia longispina” group of species and morphology of females as
well as males of that population differs substantially from that of D. ambigua populations.
Maximal population density of D. parvula populations is reached mostly during the late summer:
second half of August and September. Only in few cases we have found dense populations in
225

spring (May). Males and ephippial females occur in the period September–October. The co-occurrence
with native species D. galeata G. O. Sars, 1863 is quite common in our samples.
Both the species are small. Body size according to Brooks (1957) is 0.7–1.0 mm for D. parvula and
1.0 mm for D. ambigua. As planktivorous fish dominate most of our habitats where the species are
dwelling, we supposed that the competitive advantage of invasive species is their small body size.
We compared the mean, minimal and maximal sizes of species studied with those of D. galeata from
a reservoir (high predation pressure of fish and Chaoborus larvae) and a fish pond (low fish predation
pressure – about 500 ind. of two years old carp) (Fig. 3). The high value of the mean body size
within 4 groups of populations was found in Daphnia parvula – over 0.9 mm, and the low one in
Daphnia galeata from a reservoir with presumably high predation pressure of planktivorous fish
(roach) – over 0.7 mm. Comparison of means and corresponding 95% conf. intervals of body size on
13 different localities and 12 different sampling dates (including populations from two randomly
selected Canadian lakes) revealed large interpopulation variability (Fig. 4). With few exceptions, the
size distribution within one habitat was nearly symmetric (Fig. 5). It means that in the time of sampling
the input of newly born was already decreasing. The largest individuals of both the species were
found on different localities in size class 1.3–1.4 mm. However, the native D. galeata is able to reach
the maximal body size in some fishponds to about 2.5 mm (Ko%ínek, pers. obs.).
Fig. 3. Comparison of the mean body length among three species – Daphnia parvula Fordyce, D. ambigua
Scourfield (respective populations treated jointly) and D. galeata G. O. Sars (two populations with different fish
predation pressure: high from Drásov reservoir at P%íbram, 1998; low from pond Smyslov at Blatná, 1968).
226

Fig. 4. Comparison of mean body length among particular populations of Daphnia parvula Fordyce and D.
ambigua Scourfield. Populations are numbered as follows: D. parvula: 1 – Skalka, 2 – Novokest%anský, 3 –
Žabakor, 4 – Pýcha, 5 – Heart lake, 6 – Ruda, 7 – Dušákovský; D. ambigua: 8 – Upper Bass lake, 9 – Skalský Aug.
1998, 10 – Skalský Sept. 1998, 11 – Novokest%anský, 12 – Nový, 13 – Padr1, 14 – Závist. More detailed labels are
to find in Fig. 5.
DISCUSSION AND CONCLUSIONS
– As the species composition of the zooplankton in Czech ponds and reservoirs was intensively
studied for several decades (Šrámek-Hušek 1962, Hrbá#ek 1962, Ko%ínek et al. 1986, Hrbá#ek 1984),
the probability that both the studied species had been overlooked is negligible. We suppose that
Daphnia parvula occurred in the region not earlier than in 1975, and D. ambigua not before 1990.
This is in good agreement with hypothesis that the species are slowly spreading from western part
of Europe eastwards and southwards.
– The competitive position of both the species within the local plankton community cannot be
judged from the data available. According to our results, there is an overlap with the size distribution
of native species D. galeata in attainable maximal body size. Factors like the size of the first
reproductive instar (primipara), natality and mortality within population and attained maximal size
of competitors co-occurring in one habitat are to be studied to ascertain the mechanisms allowing
successful establishment of invader populations.
227

228

Fig. 5. Size (body length) distribution within particular populations, locality and sampling dates are given. Daphnia
parvula Fordyce – left column, D. ambigua Scourfield – right column, overall mean measurements – top of the
graph.
– Differentiation of both the species from other females of native Daphnia species is based mainly
on the reduction of rostral part of head. Both species differ in the size of three pectens of spinules
on postabdominal claws: in D. ambigua all three are small and of the same length, in D. parvula
spinules in the middle pecten are longer than in remaining two (Ko%ínek 1988) . Spinules are
however delicate and not so robust like in the group of D. pulex. The presence of ocellus is not a
reliable character as the pigmentation is meagre in the European populations of both the species.
To differentiate the males, the tip of the antennular flagellum is the best diagnostic character.
A c k n o w l e d g e m e n t
The project was partly supported by the grant to the first author (project 105/1998) from Charles University and
partly from Czech Ministry of Education (Res. Project 3111-4). We are indebted to all colleagues who helped with
their own material.
229

REFERENCES
ALONSO M. 1996: Crustacea, Branchiopoda. Fauna Iberica 7. Madrid: Museo Nacional de Ciencias Naturales,
486 pp.
AMOROS C. 1973: Évolution des populations de Cladoc#res et Copépodes dans trois étangs piscicoles de la Dombes.
Ann. Limnol. 9: 135–155.
BROOKS J. L. 1957: The systematics of North American Daphnia. Mem. Conn. Acad. Arts Sci. 13: 1–180.
DUMONT H. J. 1974: Daphnia ambigua Scourfield, 1947 (Cladocera: Daphniidae) on the European continent. Biol.
Jb. Dodonaea 42: 112–116.
FLÖSSNER D. 2000: Die Haplopoda und Cladocera (ohne Bosminidae) Mitteleuropas. Leiden: Backhuys Publishers,
428 pp.
FLÖSSNER D. & KRAUS K. 1976: Zwei für Mitteleuropa neue Cladoceren-Arten (Daphnia ambigua Scourfield, 1946
und Daphnia parvula Fordyce, 1901) aus Süddeutschland. Crustaceana 30: 301–309.
FORDYCE C. 1901: The Cladocera of Nebraska. Trans. Amer. Microscop. Soc. 22: 119–174.
HRBÁ"EK J. 1962: Species composition and the amount of the zooplankton in the relation to the fish stock. Trans.
Czechoslov. Acad. Sci., Math.-Natur. Sci. Ser. 72(10): 1–14.
HRBÁ"EK J. 1984: Ecosystems of European man-made lakes. Pp.: 267–290. In: TAUB F. B. (ed.): Ecosystems of the
World. 23. Lakes and Reservoirs. Amsterdam: Elsevier Sc. Publ. B.V, 350 pp.
HUDEC I. 1990: Distribution and biology of species of the genus Daphnia, subgenus Daphnia (Cladocera, Daphniidae)
in Slovakia. 1 st part: D. obtusa, D. pulicaria, D. parvula. Biológia (Bratislava) 45: 491–499 (in Slovak, Engl.
abstr.)
HUDEC I. 1991: Occurrence and biology of the species of the genus Daphnia, subgenus Daphnia (Cladocera,
Daphniidae) in Slovakia. 3 rd part: D. galeata, D. cucullata. Biológia (Bratislava) 46: 129–138 (in Slovak, Engl.
abstr.)
KO$ÍNEK V., FOTT J., FUKSA J., LELLÁK J. & PRAŽÁKOVÁ M. 1987: Carp ponds of Central Europe. Pp: 29–62. In:
MICHAEL R. G. (ed.): Ecosystems of the World. 29. Managed Aquatic Ecosystems. Amsterdam: Elsevier Sc.
Publ. B. V., 166 pp.
KO$ÍNEK V. 1988: [New information on limnetic cladoceran species in Czechoslovakia]. Pp.: 70–72. In: Proceedings
of 8 th National conference of Czechoslovak Limnological Society, Chlum u T%ebon& Oct 3–7 (in Czech).
MARGARITORA F. G. & SPECCHI M. 1985: Cladocera. Fauna d’Italia. Bologna: Edizioni Calderini, 399 pp.
MICHELS E., COTTENIE K., NEYS L. & DE MEESTER L. 2001: Zooplankton on the move: first results on the
quantification of dispersal of zooplankton in a set of interconnected ponds. Hydrobiologia 442: 117–126.
PETKOVSKI S. 1990: Nachweise von Daphnia pulicaria Forbes, 1893 emend. Hrbá#ek (1959) und Daphnia parvula
Fordyce, 1901 in Jugoslawien (Crustacea, Cladocera, Anomopoda). Mitt. Hamb. Zool. Mus. Inst. 87: 261–272.
SCHRIMPF A. & STEINBERG C. 1982: Weitere Fundorte der für Süddeutschland neu nachgewiesenen Cladocere
Daphnia parvula Fordyce 1901 (Crustacea, Phyllopoda). Arch. Hydrobiol. 183: 372–381.
SCOURFIELD D. J. 1947: A short-spined Daphnia presumably belonging to the “longispina” group – D. ambigua n.
sp. J. Quekett microscop. Club 4(11): 127–131.
ŠRÁMEK-HUŠEK R., STRAŠKRABA M. & BRTEK J. 1962: Lupenonožci – Branchiopoda. Fauna $SSR "6 [Phyllopods
– Branchiopoda. Fauna of Czechoslovakia "6]. Praha: Nakl. "s. Akad. V!d, 470 pp (in Czech, with Russ. and
Germ. summ.).
UÉNO M. 1960: The daphnid inhabiting the alpine lakes on Mt. Tateyama. Jap. J. Limnol. 21: 293–306 (in
Japanese with Engl. summ.).
230

Acta Soc. Zool. Bohem. 66: 231–234, 2002
ISSN 1211-376X
Anisus septemgyratus (Mollusca: Gastropoda) in the Czech Republic,
with notes to its anatomy
Luboš BERAN & Michal HORSÁK
1)
Koko%ínsko Protected Landscape Area Administration, "eská 149, CZ–276 01 M!lník;
e-mail: kokorinsko@proactive,cz, Czech Republic
2)
Department of Zoology and Ecology, Faculty of Science, Masaryk University, Kotlá%ská 2, CZ–611 37
Brno; e-mail: horsak@sci.muni.cz, Czech Republic
Received February "5, 200"; accepted October "6, 200"
Published November 4, 2002
Abstract. First specimens of Anisus septemgyratus (Rossmäessler, 1835) in the Czech Republic, were
found at three localities in South Moravia in flood-plain of the Dyje River in 1998. Dissection of about 30
specimens showed anatomical differences from A. leucostoma (Millet, 1813) and A. spirorbis (Linnaeus,
1758). Also habitats of this species seem to be different from those of A. leucostoma and A. spirobis.
Distribution, anatomy, Mollusca, Gastropoda, Anisus septemgyratus, Moravia, Palaearctic region
INTRODUCTION
Anisus septemgyratus (Rossmäessler, 1835) is known especially from East Europe (Ložek 1986,
Zhadin 1952) and its occurrence is documented also from West Siberia (Zhadin 1952). Some authors
(Hudec 1967,Glöer & Meier-Brook 1998) classified this taxon only as subspecies of A. leucostoma
(Millet, 1813) and other authors (Falkner 1989, Ložek 1986, Piechocki 1979) as species.
According to Ložek (1986) this taxon inhabits different habitats than A. leucostoma. A. septemgyratus
is known predominantly from large and not temporary pools, but A. leucostoma and A.
spirorbis (Linnaeus, 1758) are typical species of small temporary pools and wetlands. A distribution
of A. leucostoma is also wider (throughout the Palaearctic region) than A. septemgyratus (East
Europe, West Siberia) (Ložek 1986). The nearest locality is known from Slovakia (old arm of the
Hron River in Bánská Bystrica) (Ložek 1986). This species is considered as endangered or rare in
most of European countries.
MATERIAL AND METHODS
Molluscs were collected from vegetation or from water level by means of a sifter. Specimens for dissection were
drowned in water and later fixed in 70% ethanol. Dissected reproductive organs and shells are deposited in
collections of both authors. Sample of shells from second locality is deposited also in National Museum in Prague
and sample of fixed specimens is deposit in the collection of C. Meier-Brook (Germany) for further analysis.
RESULTS
First specimens were found in a pool near Nejdek (data are as follows, geographical coordinates,
code of the mapping square for faunistic mapping according to Buchar (1982), elevation above
sea-level (approximately), description of the locality, date of investigation, name of investigator –
LB – Luboš Beran, MH – Michal Horsák, LB+MH – both authors; 48°49’ 16,96” N, 16°46’ 52,04” E,
7166, 174 m, Nejdek, a pool /Bažina u Azantu/ on Nejdecké louky meadows between Nejdek and the
231

Fig. 1. Known distribution of Anisus septemgyratus (Rossmässler) in the Czech Republic.
Fig. 2. A shell of Anisus septemgyratus (Rossmässler) from locality B%eclav (restored pool in the Kan#í obora
floodplain forest, about 600 m N from the Dyje river is divided into two streams; lgt. M. Horsák, 25. IX. 1999,
height 0.9 mm, width 7.1 mm). Orig. M. Horsák.
232

Dyje River, 10. IX. 1998, LB+MH). Further specimens were found at two localities near B%eclav (48°
46’ 46,75” N, 16° 52’ 37,58” E, 7267, 158 m, B%eclav, restored pool in the Kan#í obora flood-plain
forest, about 600 m N from the point where the Dyje River is divided to two river streams, 10. IX.
1998, LB+MH, 25. IX. 1999, MH; 48°46’ 34,82” N, 16°52’ 37,76” E, 7267, 158 m, B%eclav, restored
pool in the Kan#í obora flood-plain forest, about 300 m N from the point where the Dyje River is
divided to two river streams, 11. IX. 1998, LB+MH). First locality is old and shallow pool, overgrown
by aquatic vegetation. Both other large (each about 0.5 ha) pools were recently (1991–96)
restored and are relatively fresh and with vegetation only near banks. Findings in larger pools
correspond with observation of Ložek (1986) in Slovakia and inhabiting biotopes are different to
biotopes which inhabits A. leucostoma and A. spirorbis. Especially in restored two pools was A.
septemgyratus very abundant and was eudominant mollusc. A occurrence of A. septemgyratus in
the Moravia is surprising and further research will be directed to study of population of this rare
species (especially in restored pools and their surroundings).
Figs 3–4. 3 – reproductive organs of Anisus septemgyratus (Rossmässler) from the same locality as in Fig. 2. AR
– bursa copulatrix, MR – penis retractor muscle, O – oviduct, Pr – prostate gland, PTd – penis sheat, PTp –
preputium, TR – truncus of bursa copulatrix, VD – vas deferens; scale: one segment = 1 mm. Orig. M. Horsák. 4 –
male copulating organs of Anisus septemgyratus (Rossmässler) from the same locality as in Fig. 2. MR – penis
retractor muscle, PTd – penis sheat, PTp – preputium; scale: one segment = 1 mm. Orig. M. Horsák.
233

Reproductive organs of about 30 specimen from second locality were dissected and results
were compared with Hudec (1967). Our most important results are as follows: (1) Prostate gland is
long and has many small folds. (2) Ratio between preputium and penis sheath is about 1:2. (3)
Bursa copulatrix is long and slim. First and third features are according to Hudec (1967) and also
our observations characteristic for A. septemgyratus and are different from A. leucostoma and A.
spirorbis. Second feature is according to Hudec (1967) characteristic for A. spirorbis and for A.
septemgyratus is documented ratio 1:1. This result is different to our observations and needs
further research.
According to our observation of inhabiting biotopes and especially of differences in reproductive
organs is probable that this taxon is separate species, but is suitable to carry out further
research on this taxon from other parts of Europe before final conclusion.
REFERENCES
BUCHAR J. 1982: Publication of faunistic data from Czechoslovakia. V&st. $s. Spole!. Zool. 46: 317–318.
FALKNER G. 1989: Binnenmollusken. Pp.: 112–280. In: FECHTER R. & FALKNER G. (eds): Weichtiere. Europäische
Meeres- und Binnenmolusken. München: Stenbachs Naturführer, Mosaik Verlag, 287 pp.
GLÖER P. & MEIER-BROOK C. 1998: Süsswassermollusken (Ein Bestimmungsschlüssel für die Bundesrepublik
Deutschland) "2. Auflage. Hamburg: Deutscher Jugendbund für Naturbeobachtung, 136 pp.
HUDEC V. 1967: Bemerkungen zur anatomie von arten aus der gattung Anisus Studer, 1820 aus slowakischen
populationen (Mollusca, Pulmonata). Biológia (Bratislava) 22: 345–363.
LOŽEK J. 1986: [From the Red Data Book of our molluscs – where does Anisus septemgyratus find refuge?]. Živa
34: 143 (in Czech).
PIECHOCKI 1979: Mieczaci (Mollusca). Slimaki (Gastropoda). Fauna Slodkow. Pol. 7, Warszava-Poznan: Pa4stwowe
wydawnictwo naukowe, 187 pp (in Polish).
ZHADIN V. I. 1952: Molljuski presnych i solonovatych vod SSSR [Molluscs of freshwater and saltwater of USSR].
Moskva-Leningrad: Izd. AN SSSR, 376 pp (in Russian).
234

Acta Soc. Zool. Bohem. 66: 235–239, 2002
ISSN 1211-376X
An immature stonefly from Lower Miocene of the Bílina mine
in northern Bohemia (Plecoptera: Perlidae)
Jakub PROKOP
Department of Zoology, Charles University, Vini#ná 7, CZ–128 44, Praha 2, Czech Republic and
Department of Palaeontology, Charles University, Albertov 6, CZ–128 43 Praha 2, Czech Republic;
e-mail: jprokop@natur.cuni.cz
Received May "8, 2002; accepted September 3, 2002
Published November 4, 2002
Abstract. The first record of an immature stonefly (Perla cf. burmeisteriana Claassen, 1936) from the
Lower Miocene of the Bílina mine in northern Bohemia (Czech Republic) is described and illustrated. Its
occurrence with supposed palaeoenvironmental aspect of fossilization is outlined.
Taxonomy, fossil, description, Insecta, Plecoptera, Perlidae, Tertiary, Lower Miocene, Czech
Republic, Central Europe
INTRODUCTION
Fossil stoneflies are extremely rare in Cenozoic fossil record and have not been found in Tertiary of
the Czech Republic until present. The oldest stoneflies are known from Early Permian. The family
diversity culminated in Jurassic and declined in Early Cretaceous. The general lack of fossil representatives
especially in Tertiary deposits is probably due to different living conditions preferring
running waters throughout their history. However, it is in contrary to a number of supposed lentic
taxa in Mesozoic (Sinitshenkova 1997). During Tertiary period, the palaeolakes were mainly oligotrophic
or stagnant waters. In these lakes, the running water elements were drifted by water
current, e. g., streams.
Only a few specimens of family Perlidae were found in fossil record. The oldest taxon is
Sinoperla abdominalis Ping, 1928 from the Early Cretaceous of China. Several larvae and adults
are described from Baltic amber (Eocene) and attributed to the recent genus Perla Geoffroy, 1762
(Carpenter 1992). A larva from the Oligocene of Southwestern Montana (USA) is attributed to the
genus ?Acroneuria Pictet, 1841.
The classical locality of Bílina mine (50° 34’ N, 13° 45’ E) is geographically situated in northwestern
Bohemia of the Czech Republic (see Fig. 1). From stratigraphical point of view it belongs to the
Lower Miocene (Eggenburgian/Ottnangian) of the Most Formation. The insect fauna is preserved
in the three fossiliferous horizons above coal seam (Clayey Superseam Horizon, Delta Sandy
Horizon, Lake Clayey Horizon) reflecting different sedimentary and palaeoenvironmental conditions
(Rajchl & Uli#ný 1999, Prokop 2002). The paleobotanical record and possible reconstruction
of vegetation on Clayey Superseam Horizon was previously outlined (Kva#ek 1998, Sakala 2000).
The nomenclature of larvae structures followed Theischinger (1991), Baumann (1987) and
Raušer (1980).
235

SYSTEMATIC PALAEONTOLOGY
Perlidae Latreille, 1802
Perla cf. burmeisteriana Claassen, 1936
(Figs 2, 3)
DESCRIPTION. Larval body flattened, overall length about 16.6 mm. Head with distinctly transverse
posterior occipital ridge. Pronotum rectangular (5.3 mm long and 2.1 mm wide) with clear marginal
groove separating marginal flanges; mesonotum (2.8 mm long and 6.2 mm wide) and metanotum
(2.6 mm long and 6.0 mm wide) of about same size with well-developed wing pads. Medial dorsal
suture distinctly present on all thoracic segments. Abdomen elongate, (9.2 mm long and 3.8 mm
wide) with ten visible dorsal segments; tenth segment shorter than ninth with medial apical tip
between basally thick cerci. A visible pattern of dark and light spot coloration.
DISCUSSION. The general habitus of large, dorsoventrally flattened body and dorsal color spot
pattern resembles family Perlidae, especially genus Perla. The shape of the pronotum is similar to
the recent abundant palaearctic species P. burmeisteriana Claassen, 1936. Nevertheless, a precise
classification is not possible because of the lack of the head and of the ventral branched gills on
thoracic segments.
Fig. 1. Geographical position of northwestern Bohemia within Europe (A), detailed map of the Most Basin (B),
1 – Bílina mine.
236

Lewis & Gundersen (1987) described ?Acroneura sp., an immature stonefly from Ruby River
Basin (Oligocene) of Southwestern Montana, USA. The specimen is very incomplete, smaller in
size and differs mainly from our specimen in the absence of the wing pads and a different abdominal
coloration. The nymphs of Perlidae described from the Baltic amber as Perla sp. have different
structure and shape of pronotum. They show thoracic gills that are not preserved in specimen
described here. The other fossil record of Perlidae is generally based on adult wing venation.
There is adult of Dominiperla antigua Stark et Lentz, 1992 from Dominican amber and Eoperlites
paradoxus Haupt, 1956, which is probably a fragment of wing of a Fulgoromorpha from the Eocene
of Geiseltales in Germany (Haupt 1956, Stark & Lentz 1992).
It is extremely difficult to determine Cenozoic fossils relative to modern taxa after the immature
stages. Thus, this specimen is left in open nomenclature.
Figs 2–3. Perla cf. burmeisteriana Claassen, specimen ZD0185 – photo and line drawing. Scale 3 mm.
237

MATERIAL EXAMINED. Specimen No. ZD0185 (Doly Bílina coll., Bílina, Czech Republic), a nearly complete larvae
(imprint) in dorsal view, posterior part of head, thorax, fragments of prothoracic and mesothoracic femora
present, abdomen with distinct segmentation and basal part of cerci preserved, the coloration pattern is also
preserved.
LOCALITY. Bílina mine near Bílina, Czech Republic.
AGE AND LAYER. Lower Miocene (Eggenburgian/Ottnangian), Most Formation, Clayey Superseam
Horizon.
CONCLUSIONS
The presence of this specimen indicates an aquatic environment. Stonefly nymphs are predators
that occur mainly in running water. The recent species preferably inhabit of small streams to large
rivers but they are found also in cold ponds and lakes. The family Perlidae comprises about 400
recent species that are widely distributed in the northern hemisphere with a few genera extending
into the southern hemisphere in Africa and South America. The major part of the genera is restricted
in the East Palaearctic (Baumann 1987, Nelson 1996). The history and living conditions of
modern Plecoptera started with the break up of Pangaea (Zwick 2000).
This fossil is the unique record of a stonefly in the Bilina paleolake. We can suspect that this
element is allochthonous and was probably transported into basin by an occasional flooding from
early uplands. This is supported by the fish fauna (M. Böhme pers. comm.), plant megafossils
(Pinus ornata Stemberk) and pollen analysis significantly confirmed from Lake Clayey Horizon.
The evidence of facultative foothills elements in Clayey Superseam Horizon is not clearly documented,
e. g., pollen analysis (Pinaceae). However, a stonefly is probably indicative of life apart
from back swamp or lake.
A c k n o w l e d g e m e n t s
The author is grateful to Zlatko Kva#ek (Charles University, Praha) and André Nel (Museum national d’Histoire
naturelle) for suggestive advices and to Zden!k Dvo%ák (Doly Bílina) for the loan of the material. My express
thanks goes to Miss Zuzana "adová (scientific illustrator, Charles University, Praha) for the magnificent habitual
drawing. The research was supported by grants of the Ministry of Schools J13/98-113100004 and GA"R 205/01/
0639.
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239

Acta Soc. Zool. Bohem. 66: 240, 2002
ISSN 1211-376X
BOOK REVIEW
SERVICE M.W. (ed.): Encyclopedia of Arthropod-transmitted Infections of Man and Domesticated
Animals. Wallingford, Oxon: CABI Publishing, a division of CAB International, 2001. XVI+579 pp. Format
170×240 mm. Hardback. Price Lstg 99.50 (USD 185.00). ISBN 0 85199 473 3
The editor is Emeritus Professor of Medical Entomology at the Liverpool School of Tropical Medicine. He has
been advised by the five international advisors – foremost authorities in the field: R. W. Ashford, C. H. Calisher,
B. F. Eldridge, T. V. Jones, and G. Wyatt. The project of this publication involved 88 authors of 19 countries:
physicians, veterinarians, parasitologists, entomolologists and zoologists. It is pointed up in the preface that this
is an encyclopedia, not a specialized medical or veterinary textbook, and so the aim has been to present up-todate
basic information on the transmission of a broad range of infections causing serious illnesses. Vertebrates,
including humans, become infected when they are bitten by an infected arthropod taking a blood meal. The
enormous increase in global travel is one of the major factors in the spread of infectious diseases, such as West Nile
virus which has emerged in temperate regions of Europe and North America. This encyclopedia contains 150
entries, describing arboviral, viral, bacterial, rickettsial and spirochaetal infections, protozoal and filarial infections.
Entries range from 100 words to 5000 words, they are arranged in alphabetical order starting with
aegyptianellosis and concluding with the Zika virus.
Arboviral and viral infections discussed include infections as the African horse sickness, African swine fever,
the Chikungunya virus, the Colorado tick fever, Ilesha virus, Ilheus virus, Issyk-Kul virus disease, Jamestown
Canyon virus, Japanese encephalitis, Keystone virus, haemorrhagic fevers, Powassan encephalitis, Rocio encephalitis,
Ross River virus, Venezulean equine encephalitis, vesicular stomatitis, West Nile virus, and many more.
Bacterial, rickettsial and spirochaetal infections incorporate plague, tularaemia, endemic typhus, epidemic typhus,
Rocky Mountain spotted fever, tick-borne typhuses, avian borreliosis, Lyme borreliosis, tick-borne and
louse-borne relapsing fevers.
Entries on protozoal and filarial infections are concerned with trypanosomes as aetiologic agents of African
sleeping sickness and the Chagas disease in humans, and Trypanosoma species in animals affecting the livestock
in tropical regions. Entries dedicated to parasitic diseases present the malaria in humans and birds, leishmaniases,
babesioses and theilerioses in humans, cattle, sheep and goats, equines and in dogs. Animal and human filarial
infections transmitted by bloodsucking arthropods are presented here by bancroftian and brugian filariasis,
dirofilariasis, mansonelliasis, onchocerciasis, parafilariasis, stephanofilariasis, and others.
Detailed entries provide insights into diverse arthropods – insects and arachnids, transmitting the named
infections. Emphasis is given to mosquitoes, black-flies, biting midges, eye-flies (Chloropidae), fleas, horse-flies,
mosquitoes, sand-flies, stable-flies, tse-tse flies (Glossinidae), to ticks and mites, and others.
Information on diseases include distribution, aetiological agent, transmission cycles, diagnosis, clinical symptoms,
treatment and control measures. Following each entry is a selected, mostly periodical or monographic
bibliography and more specialized textbooks relevant to the entries, to aid further reading on the topic. Entries
dealing with arthropods constitute a highlight of identification, biology, transmitted diseases and control.
The volume is extensively illustrated by schematic line drawings and black-and-white photographs: featured
are diagrams of natural cycles of transmission, adult forms and developmental stages of ectoparasitic vectors:
insects and arachnids, life cycles of causative agents and factors involved in pathogenesis, light and electron
microphotographs, genetic processes, maps of worldwide distribution of various infections, views of terrestrial
biomes, pathological and histopathological abnormalities in humans and animals, clinical conditions, skin lesions,
and more. In addition, there are numerous summary-type tables offering overviews of diverse microorganisms,
characteristics of symptoms, classification of drugs, dosages of insecticides, and more. This encyclopedia presents
a most comprehensive up-to-date handbook essential to anyone who is interested in finding out transmits of
infectious agent, where it is found and whether it can be cured or prevented. It covers a broad range of related
disciplines including human and veterinary medicine, epidemiology, epizootiology, immunology, microbiology,
virology, parasitology and entomology.
Jind%ich Jíra
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