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Actinocyclus normanii (Bacillariophyceae), in some rivers and

lakes in Europe

Article · January 1990

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K.T. Kiss, R. Le Cohu, M. Coste, S.J. Genkal & V. Houk, 111-123,
Ouvrage dédié ti H. Germain, Koeltz 1990.

Actinocyclus normanii (Bacillariophyceae),

in some rivers and lakes in Europe. Morphological
examinations and quantitative relations.
Actinocyclus normanii (Bacillariophyceae) des lacs et rivieres d'Europe.
Morphologie et relations quantitatives.
by

K.T. KISS,
Hungarian Res. Station, Hungarian Acad. of Sciences. Göd Jávorka S.u.14. 2131 Hungary.
René LE COHU,
Lab. d'Hydrobiologie, Univ. P. Sabatier.Ll B Route de Narbonne, 31062 Toulouse, France.
Michel COSTE,
Cemagref-Qepp. 50 Avenue de Verdun Gazinet B.P.3, 33610 Cestas France.
S.1. GENKAL,
Instit. of Biology of InIand Waters. Acad. of Sciences of U.S.S.R.152742 Borok.
Jaroslav ,Nekouz. U.S.S .R.
V. HOUK,
Waterworks of Prague, U.H.T. Podolská 15. 14700 Praha 4. Czechoslovakia.

Abstract: Actinocy clus normanii was colleeted from sever al rivers and lakes in
Czechoslovakia, East Germany, France, Hungary, the Soviet Union and in the Caspian Sea.
In some cases (like in the Danube river), the species was found in great number representing a
large biomass in summer. Actinocyclus normanii is thought to indicate heavy pollution and
advanced eutrophication in the great lakes in North America. The present study aims at
giving newelements of information about the morphological features. Actinocyclus
normanii (Greg.) Hustedt f. normanii and A. normanii f. subsalsa (Juhl, Dannf.) Hustedt.
cannot be separated by the size because of the continuum observed between them; in the same
way, they cannot be distinguished by their ecology, as smaller and larger cells can be
observed in both fresh and sea water.
Key-words: Bacillariophyceae, Actinocyclus, morphology, ultrastructure, ecology.

Résumé: Actinocyclus normanii a été recensé dans plusieurs lacs et r ivieres de

Tchécoslovaquie Allemagne de l'Est, France, Hongrie et Union Soviétique ainsi que dans la
mer Caspienne. 11 représente parfcis comme dans le Danube une part importante du
potamoplancton estiva!. Dans les Grands Lacs d'Amérique du Nord cette espece est considérée
comme indicatrice de fortes pollutions ou ou tout au moins d'une eutrophisation marquée. La
présente étude tente d'apporter des éléments nouveaux concemant la morphologie dc l'espece
li l'aide dc la microscopie photonique et électronique. Actinocyclus norman ii (Greg.) Hust. f .
normanii et A. normanii f. subsalsa (Juhl, Dannf.) Hust. ne peuvent étre dissociées li partir
des eriteres de taille ou de leurs exigences écologiques car toutes les formes transitoires entre
ces deux taxons ont été observées aussi bien en milieu continental que marin.
Mcts-clés : Bacillariophyceae, Actinocyclus, morphologie, ultrastructure, ecologie.

III
 Introduction

Several papers reported on Actinocyclus normanii in North Europe and North America
in the last lD-15 years. Its expansion was thought to be in connection with pollution
and the increase of eutrophication, especially in North America (Stoermer & Yang
1969). Although its presence in Europe has been known for more than 100 years
(Juhlin-Dannfelt 1882), information about its distribution remains limited. The most
detailed morphological study was carried out by Hasle (1977) , and valuable additional
data can be obtained from the works by Bekher & Swale (1979) and Klee & Steinberg
(1987).

Actinocyclus normanii has been recorded in many European rivers and lakes during our
investigations. Besides the morphological study we would like to show a comparison of
different populations and their quantitative relations.

Material and methods

Samples were colleeted from:

- Czechoslovakia: Svratka river (1986 August) near Brno; River Vltava at Prague
(1987 September); Nové Mlyny Reservoir on River Dyje(1987-1989); River Danube at
Bratislava (1988 November), River Váh at Kolárovo (1989 Septcrnber).
- France: Dordogne river downstream Liboume, Fensch (East France) and in the
lower part of the River Oise.
- East Germany : River Spree (1989 August); Müggelsee (1988 April) near Berlin.
- Hungary : Danube river, entire stretch (in the main arm in many old branches,
1988-1989).
- The Soviet Union: Volga river near Astrachan, Caspian Sea (approximately 100
km south of the Volga delta - two sampling points: a) 44°43' N. latitudc, 47"52' E.
longitude; b) 440)6' N. latitude, 47"56' E. longitude.

Samples were colleeted from ncar the surface layer and fixed with Lugol solution. In
some samples, phytoplankton cells were counted using the Utermöhl method (Lund et
al. 1958).

The morphological investigation of A. normanii was perforrned under light (LM) and
electron (EM) microscopes. Sediménted plankton samples wcrc digested with hydrogen
peroxide and washed with distilled water. For LM. examination, the cleaned sarnples
were mounted in Pleurax resin (R.I.=1.75 - 1.77). The usual methods were used for EM
studies. An üPTüN inverted microscope, a ZEISS Amplival and Jenával-contrast
microscope, a NACHET NS400 microscope were used for LM studies, a TESLA BS
500, a lEüL lEMlOO S, HITACHI H-300 transmission electron microscope and a
lEüL lSM 25S, lSM35 scanning electron microscope for TEM and SEM studies.

The structural elernents of the valves were measured and analysed by methods based on
the works of Genkal (1977) and Genkal & Kuzmin (1979); the terminology used
follows that suggested by Anonymous (1975) and Ross et al. (1979).

112
 Results

Morphological investigations

The relatively large cell size and the numerous discoid chloroplasts are the most
noticeable features of the living cells of A. norman ii under L.M. (Figs. 1,4,7).
Thcrefore it is easy to identify the species and make accurate qualitative analysis even in
undigested samples.

In the cleaned samples, the frustule shows a concave or convex val ve face in the LM.
The valve face is divided into sectors. The longest areolae delimiting a sector arc radial,
the others which are decreasing in length are parallel to them (Figs. 6,9,21). There arc
several rimoportulae at regular intervals on the valve mantle (Fig. 3). The
pseudonodulus, which is characteristic 10 this genus and is a scgregating criterion from
Coscinodiscus, is not visible with LM. Il is very difficult 10 investigate A. normanil
under TEM as the electron rays can hardly or not al ali penctrate through the thick
sil icon wall of the frustule (Fig.9).

The frustule'structure can be studied beuer by SEM. The diameter is 13-66 11m, the
length of the pervalvar axis ll-23 11m (Table 1). The ratio of the two is between 2.1-
1.0, generally 1.6-1.3. The ratio is between 1.2-1.0 in cells before a division, and
between 2.1-1.9 after il. On the valve face, the areolae arc occludcd extemally by a
cribrum with very tiny pores (Fig.26); they are opened internally with a weil developcd
foramen (Fig. 16-21). Very few cells with undeveleped or poorly developed cribra were
observed (Fig. 13).6- 12 areolae in 10 11mwere general ly counted on the valve face, 11-
22 on the mantle; the number of the arcolae depends on the diametcr, the higher on the
small val ves (15-25 11m), the lower on the large ones (50-66 11m). On the valve mantlc,
areolae run down straight, parallel to the pervalvar axis and in quincunx arrangement
(Figs. 8,10,16,17,21,22,29).

The external opcnings of the rimoportulae are clearly visible on the mant!e's curved
outer surface. The pseudonodulus is located above the rimoportulae in the transition
zone between the val ve face and the mantle; it is larger than the extemal aperture of the
rimoportulae (Figs.24-25-27). The pseudonodulus inner openings can present several
ra ther large pores or can be partially occluded with only one pore. The most typical
morphological features of inner part of the frustule are the rimoportulae, 3-11 in
number, depending on diameter. Their inner tube is quite long, with a weil developed lip
at the end (Figs.28,29). The valvocopula and copula can be usually clearly seen in
girdie view under SEM.

Some morphometric and morphological characteristics arc distinct in the investigated

populations (Table 1). Specimens from River Spree and Müggelsee are square in girdJe
view, whereas the ones from the Danube arc rectangular. Moreover the size and volume
arc different, Individuals from the Danube have a larger average diameter and alonger
pervalvar axis; consequently, their average biomass is considerably more important
(close to 30%). This fact can be verified when biomass values are compared (fig. 30).

113
 Table 1. Data and measurements of the valve structure elements of 3
different A ctinocycius normanii populations
(Danube: 300 data, Pree-Müggelsee: 100 data; Caspian sea: 50 data).

Danube r iver d pa d/pa b

min-max 14-31 11-22 .89-1.7 2150-16600
x 21.94 16.65 1.30 6467
Sx .17 .13 .01 125.47
S 3.01 2.21 .24 2173
S2 9.03 4.89 0.06 4700000
CV 13.70 13.28 18.49 33.60

Spree- Müggelsee d pa d/pa b

min-max 13-25 14-23 .65-1.25 2120-8738
x 17.89 17.63 1.02 4594
Sx .26 .18 .01 220.40
S 2.60 1.86 .13 2215
S2 6.79 3.45 0.02 4900000
CV 14.49 10.48 12.66 47.98

Caspian Sea d nlp na

min-max 31-60 5-11 7-9
x 43.8 7.7 7.9
Sx 0.9 0.2 0.1
S 7.0 1.5 0.7
CV 15.9 19.4 8.8

d: diameterof valve in um: pa: length of pervalvar axis in um; d/pa: ratio diameter/
pervalvar axis; b:biomass in Jlm3; nlp: number of labiate process; x: mean value; na:
areolae in 1O~; Sx: error of mean value; S: corrected scatter; S2: variance; ev: variation
coefficient; n: number of data.

Quantitative relations :
We have accurate and regular data concerning the quantitative relations of A. normanii
from the 1988 vegetation period in the Danube near Budapest. Based on these data the
species appeared in considerable number (11 thousand ind.drrr '), at the middie of June
(Table 2)_

By the middie of July the species increased in number (550 thousand ind. drrr"), and
began to decline after that. After the mid-October, we found it in occasional samples
only (few thousand ind. dm-). As it is a large species, few individuals represent a
proportionally greater biomass. It is worth mentioning that in the summer of 1989, we
found the species only in a few samples above 20 thousand ind.zdm>. Before 1988, the
species was not coUected from the Hungarian section of the river.

114
 Table 2: Quantitative relations of phytoplankton of the river
Danube at Göd (algal counts ind/ml)

1988 AlI ind. Actinocyclus Centrales Pennales Chloroph yceae others

VI.15. 10901 II 7117 275 3072 437

VL22. 22710 25 15fJJ7 275 6322 506
VI.29. 4019 40 29475 1373 8798 551
VII. 7. 42301 549 32037 550 9347 367
VII.13. 37634 366 27827 826 8706 275
VIL20. 9720 153 6012 61 3493 154
VII.27. 57%5 122 38078 1098 17691 1098
VIII.3. 32095 183 21480 915 8785 915
VIII.lO. 31240 61 19222 488 104 1098
VIII.17. 34416 183 24837 610 8359 610
VIII.24. 4973 34 2859 170 1570 374
VIII.31. 6162 31 4241 152 1465 304
IX.7 5368 46 3569 79 1401 319
IX.l4. 7552 61 5675 151 1377 349
IX.2I. 4624 34 2459 135 11367 663
IX.29. 13186 23 8650 368 2771 1397
X.5. 17292 15 11718 806 3521 1247

When Actinocyclus was recorded in the highest number from the Danube, the river had
an invariant stream-flow regime (Fig.31). The dominant species in the phytoplankton -
as usual at summertime - were centric diatoms, and Chlorococcal algae. Among the
Centrales besides Actinocyclus normanii, Stephanodiscus hantzschii f tenuis (Hust.)
HAk. & Stoermer, S. invisiuuus Hohn & Hellerman, S. minutulus (Kütz.)Cleve &
MöIler occurred in large numbers (lOS to 106 ind. drrr ') . The amounts of Actinocyclus
in Czechoslovakian waters were different in 1986-1989. In the Nové Mlyny Reservoir
the number of ce Ils reached 1500 thousand dm·3 in late summer 1986-89, in the River
Svratka this amount reached several hundreds dm' in August 1986 (after Dr. P.Marvan)
and in other localities this species was observed only unfrequently or rarely. It was
present in small arnount in the River Volga, the Caspian Sea and in Freneh Waters. The
species was abundant in River Spree and Müggelsee, it is the characteristic species of
the waterblooms in the growing season.

Discussion

The morphological characteristics of the studied populations were basically the same as
mentioned in literature (Bekher & Swale 1979, Hasle 1977, Klee & Steinberg 1987).
Stoermer & Yang (1969) found short colonies in Lake Michigan consisting of maxi-
mum 10 individuals besides single cells. Hasle (1977) noted solitary cells and cells in
pairs. We also noticed paired cells remaining together after division (Fig.7).

115
Aceording to Hasle (1977) the pervalvar axis is half or a littIe more than half the
diameter (diameter/pervalvar axis - d/pa = 2). We could find the same data in Beleher &
Swale's work (1979). This ratio is certainly higher in specimens colleeted from the
Danube. It is, using mean data of speci mens from River Spree and Müggelsee, twice as
big as the figures mentioned above. Large part of the population is square in girdle view
and does not have a disk like shape.

The distinction of the two forms made by Hustedt (1957), was based on the diffcrences
in diameter. Several authors discussed the validity of this separation (Hasle 1977,
Beleher & Swale 1979, Klee & Steinberg 1987). They added that A. normanii
i.normanii which has larger cells prefcrs sea water, while the smallcr A. normanii f.
subsalsa is usually to be found in fresh water, so they can be considered as ecotypes. In
Beleher & Swale's opinion, the separation of the two forms is not weil founded. Our
investigations confirm this opinion (Table 3). In Hustedt' separation, the two forms
overlap each other in cell size; in other words, thcre is a continuous transition from one
to one form to the other. Our data are consistent with this observation. As a matter of
fact, population from River Vltava, Rivcr Spree and Müggelsee can rank in the
subsalsa size range. However, speci mens from the Volga and the Caspian sea show the
entire size range between f. subsalsa and f. normanii. This also tends to pro ve that the
separation of the two forms as ecotypes is not correct, as we found cells in the River
Volga with a diameter 65-66 um, and in the Caspian Sea with a diamcter 30-66 um
(salinity 3.2 - 4.5 %). The sea populations may consist of larger cells, and fresh water
populations of smaller ones, but it does not seem appropriate to scparate them on such
a basis.

Table 3. Valve diameter in um of Actinocyclus normanii.

Act. f. subsalsa Act. f. normanii

Hustedt (1925,1957) 16-44 30-110
Hasle (1979) 16-44 42-67
Klee & Steinberg (1987) 20-25*
Beleher & Swale (1979) 24-58
our results (2 mixed forms)
Danube 13-31
River Vltava 14-32
River Spree, Müggelsee 13-25
River Volga 35-66
Caspian Sea 31-60
(* measured by us on the basc of micrographs from different papers).

A. normanii was a very rare species in Hungary. Hajós (1979) described it from fossils,
and Szemes (1972) observed it in one case from the Danube; its invasive occurrence was
not noticed until recently. Its presence in the 1988 summer phytoplankton of the
Danube was surprising not only because it was not seen in the previous year, but also
because it was present in a considerable number. Its number was 1.3 % of alI

116
phytoplankton and 1.7% of Centrales in the beginning of July. This means that the
amount of Amormanii carne just after the abundances of Skeletonema potamos (Weber)
Hasle, Stephanodiscus hantzschii f.tenuis and S. invisitatus. If we compare the
biomass values, we find that A. normanii represented II% of ali phytoplankton and/or
14% of Centrales species, so A.normanii preceded the above mentioned species.

Amormanii is found in same amount and ratio in other waters. It fonned 3-3.5% of ali
diatoms in Main river between 1972-73 (Lange-Bertalot 1974). Holland & Claflin
(1975) reported a maximum figure of 90-130.103 ind/dm" in the Green Bay of Lake
Michigan in June 1970.

It is difficult to explain why A.normanii appeared invasively in the Danube. As Hohn

(1969), Stoenner (1978), Stoenner & Kreis (1980) and Stoenner & Yang (1%9) had
already noticed, the species proliferated in the Great Lakes in North America, where
there was high poJlution and extensive eutrophication. In relation to this fact,
Anormanii is considered as indieater of water poJlution or eutrophication. No increase
of water pollution and eutrophication was observed in the Danube recently and it can not
account for the apparition of this species at that time. Indeed the potential trophic level
(N. P. supply) of the Danube has made the river hypertrophic for years, and the actual
trophic level in the vegetation period is eu-polytrophic (100-300 ug chlorophyll a / drn '
; Kiss 1985). A. normanii should have been prcscnt in large numbers in the water for
years, and we would have noticed its presence, as we have been making regular
investigation of the phytoplankton on the Danube for three decades.

Acknowledgements

We wish to thank Dr. S. Hoeg and Dr. 1. Köhler (Inst. Geogr. GeoecoJ. Hydrobiol. Lab. East
Germany ), Dr. E. Labunskaya (Centre of Hydrometeorology, V.S.S.R.) for making material
available for this investigátion from the River Spree, Müggelsee, River Volga and Caspian
Sea. We thank in particular Dr. E.F. Stoermer (Great Lakes Res. Div. V.S.A.) for providing
very interesting and important personal informations about A. normanil in the Great Lakes
of North America. Many thanks to Dr. P. Marvan (Bot. Inst. Czechosl. Acad. Sci. Trcboii
Czecho slovakia) for providing data on A. normanil from River Svratka, Nové Mlyny
Reservoir and River Váh.

References

ANONYMOVS. (1975). Proposals for a standardization of diatom terminology and

diagnoses. Nova Hedwigia, Beih. 53, 323-354.
BELCHER, J.H. & SWALE, E.M.F. (1979). English freshwater records of Actinocyc/us
normanii (Greg.) Hustedt (Bacillariophyceae). Br. PhycoL J. 14, 225-229.
GENKAL, S.1. (1977). K metodike podschota nekotorih taksonomicheski znachirnih
strukturnih elementov stvorki u diatomovih vedoroslej sem. Thalasssiosiraceae Lebour
emend Hasle (Bacillariophyta).[(On counting of some taxonomically significant structural
elements of valves in the diatom algae of the family Thalassiosiraceae Lebour emend Hasle
(Bacillariophyta).]. Bot. Zhurn. 62, 848-851.
GENKAL, SJ. & KVZMIN, G.V. (1979). Biometricheskiy an aliz osnovnih strukturnih
elementov stvorki u vidov roda Stephanodiscus Ehr. (Bacillariophyta). [Biometric analysis

117
of main structural elements of valves in the species of genus Stephanodiscus Ehr ,
(Bacillariophyta)]. Bot Zhurn, 64, 1237-1244.
HAJÓS, M. (1969). Die Diatomeen der rniozanen Ablagerungen des Mátravorlandes.
Geologica Hungarica, Seria Paleontologica 37.
HASLE, G.R. (1977). Morphology and taxonomy of Actinocyclus normanii Lsub salsa
(Bacillariophyceae). Phycologia 16, 321-328.
HOHN, M.H. (1969). Qualitative and quantitative analyses of plankton diatoms. Bull. Ohio
Biological Survey. 3, 1-208.
HOLLAND, R.E. & CLAFLIN, L.W. (1975). Horizontal distribution of planktonic diatoms in
GreenBay, mid-July 1970. Limnol. Oceanogr.20, 365-378.
HUSTEDT, F. (1957). Die diatomeenflora des flusssystems der Weser im Gebiet der Hansestadt
Bremen. Abh Naturw YerBremen s (3), 181-440
á

JUHLIN-DANNFELDT, H. (1882). on the Diatoms of the Baltic Sea. Bih. svensk, Vetensk.
Akad. Handl. 6,1-52
KISS, K.T. (1984). Changes of trophity conditions in the riv er Danube at GÖd. Danubiala
HungaricaX(W). Annls.UnivSci Budapest., Sect Biol, 24-26, 47-59.
KLEE, R. & STEINBERG, C. (1987). Kieselaigen Bayerischer Gewasser. Loseblattsammlung.
Bayer. Landesamt für Wasserwirshaft München. 4/87, 225 p.
LANGE-BERTALOT, H. (1974). Das Phytoplankton im unteren Main unter dem Einfluss
starker Abwasserbelastung. Cour. Forsch. Inst. Senckenberg 12, 1-88.
ROSS, R., COX, EJ., KARAYEVA, NJ., MANN, D.G., PADDOCK, T.B.B., SIMONSEN, R. &
SIMS, P.A. (1979). An amended terminology for the siliceous components of the diatom
cell. Nova Hedwigia beih. 64, 513-533.
STOERMER, E.F. (1978). Phytoplankton as indicators of water quality in the Laurentian
Great Lakes. Trans Am. Microse. Soc. 99, 2-16.
STOERMER, E.F. & KREIS, R.G. (1980). Phytoplankton composition and abundance in
Southern Lake Huron. U.S. Environmental Protection Agency, Duluth, Minnesota. USEPA
Report WEPA-600/3-80-061, 384 p.
STOERMER, E.F. & YANG, J.J. (1969). Plankton diatom assemblages in Lake Michigan.
Univ. Michigan Great Lakes Res. Div., Spec. Rep. 47,1-268.
SZEMES, G. (1964). Untersuchungen über das Phytoplankton der ungarischen Donaustrecke
in Sommermonaten. (Danubiala Hungarica XXV). Annls Univ. Sci. Budapest. Sect. Biol. 7,
169-199.

118
Plate 1. Fig. 1. Living cell in valve view showing the many discoid plastises. Figs. 2,3.
The same valve in different focus. Fig. 2. focused to the plane of cribrae. Fig. 3. focused to
labiate processes. FigA. Living cell in girdIe view. Figs. 5,6. Valve face showing the
different areola pattem. Fig. 7. Dividing living cell in girdIe view. Fig. 8. Frustule in girdIe
view show ing valvocopula and the ligula of a copula. Fig. 9. TEM structure of valve face.
Fig. 1-8 LM. Fig. 9 TEM, scaIe: 10 um. Figs 1,4,7. specimens from Danube near Budapest.
Figs 2, 3, 6, 8. Danube at Bratislava. Fig.5 R. VItava at Prague. Fig.9. R. Spree near Berlin.

119
Plate 2. Fig. 10. Whole frustule with epicingulum from Müggelsee. Fig. ll. Who le frustule
from the Caspian Sea (sampling point b), with the openings of labiate processes (arrowed).
Fig. 12. Valve show ing the mantle's areola structure and the pseudonodulus (arrowed) - R.
Danube near Budapest. Fig. 13. Valve face with different areolation (the smaller part of
areolae have no vela) from old branch of R. Danube. Figs 14,15. Valve face of specimens
from R. Spree (pseudonodulus arrowed). Figs. 10-15. SEM, scale: 10 um.

120
Plate 3 . Figs. 16-21. Internal view of valve face, SEM, scale - 10 um, Fig. 16. Specimen
from the Caspian Sea (sampling point b) showing the characteristic areola pattern. Fig. 17.
Valve with 3 labiate processes from Müggelsee. Fig. 18. Broken valve from the R. Volga.
Figs. 19-21. Valves with 4,5,6 labiate processes from the R. Spree.

121
Plate 4 . Fig. 22. The inner part of a frustule with the valve and valvocopula. Fig. 23. SEM
view of a valvocopula. Fig. 24. The valve face- valve mantle junction with the opening of
the labiate process and pseudonodulus. Figs. 25, 27. Pseudonoduli with different structures.
Fig. 26. External view of the transition zone between valve face and valve mantle,
differently grouped pori of areolae and the opening of labiate process. Figs. 28-29. Side and
top views of labiate process. Fig. 28 shows some foramens elosed with a thin silica
membrane. Figs. 22-29 SEM. Figs. 22-23. scale: 10 um. Figs. 24-29. scale: 2 um,

122
 %
30,------------------------------------------------- ,
FIG.30 -o- DANUBE
...•. SPREE

20

10

O+-~~~_.~_.--~._~~~~~=T~~~~
o 2000 4000 6000 8000 10000 12000 14000 16000 18000
Biomass category of thousands (11m3)

40 FIG. 31

30

20

10

o .=:; .=:; -B .=:; .=:; .=:; .=:; "E ..c:

.=:; -v; .=:; ..c:
-v; ..c:
.=:;
.,.., N a- r- e<> <::> r-e <") ö r- ~ ;:;:; r-. ~ ;:::;
c(, .,.., [0
<u
'""<u N
<u ;>.. ;>.. '""c-, '""
;>.. '""ci) cs.. ~o.. ~o.. '""~ ~u t-
c: c: c: ci) ci) ci) ci) o.. <c
~ ~ ~ ~ ~ ~ ~ =>
A'
=>
<t:
=>
-c
=>
<t:
=>
<t:
<u
c/)
<u
c/)
<u
c/)
<u
c/)
o C

Plate 5 . Fig. 30. Biomass distribution of two A. normanii populations (each volume data
ranged to categories thousand's e.g. from 1500-2499 um ranged to 2000, from 2500-3499
ranged to 3000). Fig. 31. Changes of biomass of the main phytoplankton groups and
Actinocycius normanii in the R. Danube near Budapest (1988).

123

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