Studies on Eurytemora

Proceedings of the Eurytemora conference,

St. Petersburg, 2019

By
Natalia M. Sukhikh, Sami Souissi, Gesche Winkler
and Victor R. Alekseev (Editors)

CRUSTACEANA MONOGRAPHS, 23

LEIDEN | BOSTON

For use by the Author only | © 2021 Larysa Samchyshyna et al.

 CONTENTS

Note on Original Publication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

BIOGEOGRAPHY
VICTOR R. ALEKSEEV & NATALIA M. SUKHIKH, On time and place of
origin of continental calanoid families: a hypothesis . . . . . . . . . . . . . . 5
V. I. LAZAREVA, Distribution of Eurytemora caspica Sukhikh & Alek-
seev, 2013 (Copepoda, Calanoida) in the water reservoirs of the
Volga and Don river basins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
LARYSA SAMCHYSHYNA, YULIA GROMOVA & KATERYNA ZORINA-
SAKHAROVA, Recent distribution of Eurytemora velox (Lilljeborg,
1853) (Copepoda, Calanoida) in brackish and fresh waters of
Ukraine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
VASIL V. VEZHNAVETS & ANASTASIYA G. LITVINOVA, Representatives
of the genus Eurytemora Giesbrecht, 1881 (Calanoida, Temoridae)
in aquatic ecosystems of Belarus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

MORPHOLOGY
ELENA FEFILOVA, NATALIA SUKHIKH, EKATERINA ABRAMOVA &
ILYA VELEGZHANINOV, About the systematics of Palaearctic Eu-
rytemora (Copepoda, Calanoida) based on morphological analysis,
with focus on Eurytemora gracilicauda Akatova, 1949 . . . . . . . . . . . 59
DMITRY LAJUS, NATALIA SUKHIKH & VICTOR ALEKSEEV, Stochastic
phenotypic variation: empirical results and potential use in Eury-
temora research (Copepoda, Calanoida) . . . . . . . . . . . . . . . . . . . . . . . . . 77
ŁUKASZ SŁUGOCKI, Variability of mandible shape in the freshwa-
ter glacial relict Eurytemora lacustris (Poppe, 1887) (Copepoda,
Calanoida, Temoridae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

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vi CONTENTS

ANISSA SOUISSI & SAMI SOUISSI, Abnormalities in shape and size

of Eurytemora affinis (Copepoda, Calanoida) and its eggs under
different environmental conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

ECOLOGY
JORY CABROL, RÉJEAN TREMBLAY & GESCHE WINKLER, Differen-
tial eco-physiological performances of two pseudocryptic species
of the Eurytemora affinis complex (Copepoda, Calanoida) in the St.
Lawrence estuarine transition zone: a reciprocal transplant experi-
ment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
ELENA G. KRUPA, The ecological preferences of Eurytemora affinis
(Poppe, 1880) in the water bodies of Kazakhstan (Central Asia) and
some notes about Eurytemora caspica Sukhikh & Alekseev, 2013 165
LAURI KUISMANEN, LOUISE FORSBLOM, JONNA ENGSTRÖM-ÖST,
ULF BÅMSTEDT & OLIVIER GLIPPA, Salinity effects on egg pro-
duction, hatching, and survival of Eurytemora affinis (Copepoda,
Calanoida) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
ASTRA LABUCE, SOLVITA STRAKE & INTA DIMANTE-DEIMANTO-
VICA , Eurytemora affinis (Poppe, 1880) (Copepoda, Calanoida) in
the Gulf of Riga, Baltic Sea — elemental composition and diurnal
vertical migration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
ALEXANDER W. TIMPE & BART T. DE STASIO, Comparison of respira-
tion rate and electron transport system (ETS) enzyme-mediated re-
duction assay of the invasive copepod Eurytemora carolleeae Alek-
seev & Souissi, 2011 (Calanoida, Temoridae) in Green Bay, WI,
U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
YAN-GUO WANG, LI-CHUN TSENG, ROU-XIN SUN, ZHI-YONG LIU,
MAO LIN & JIANG-SHIOU HWANG, Effects of the China Coastal
Current on the community structure of planktonic copepods in early
spring, with notes on Eurytemora pacifica Sato, 1913 in the western
Taiwan Strait . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247

BIOLOGY IN TAXA RELATED TO Eurytemora

ELENA YU. NAUMOVA & VICTOR R. ALEKSEEV, How many gener-
ations does Epischura baikalensis (Copepoda, Calanoida) have in
Lake Baikal? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

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 CONTENTS vii

LI-CHUN TSENG, YAN-GUO WANG, GUANG-SHAN LIAN & JIANG-

SHIOU HWANG, A multi-year investigation of the Temoridae (Cope-
poda, Calanoida) assemblage succession within the interplay waters
of the northern South China Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
DENIS S. ZAVARZIN, Pseudodiaptomus japonicus Kikuchi, 1928 (Cope-
poda, Pseudodiaptomidae), a brackish-water copepod formerly
known as P. inopinus Burckhardt, 1913, on Sakhalin Island (Rus-
sian Far East) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
Taxonomic index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 309

First published as a Special Issue of Crustaceana. The bracketed

page numbers in this book refer to the page numbers in Crus-
taceana 93, 3-5 (2020). When citing from this book, refer to these
bracketed page numbers.

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 [When citing this volume, please refer to Crustaceana 93 (2020), 241-547]

Proceedings of the Eurytemora conference, St. Petersburg, 2019

RECENT DISTRIBUTION OF EURYTEMORA VELOX (LILLJEBORG, 1853)

(COPEPODA, CALANOIDA) IN BRACKISH AND FRESH WATERS OF
UKRAINE

BY

LARYSA SAMCHYSHYNA1,3,4 ), YULIA GROMOVA2 ) and

KATERYNA ZORINA-SAKHAROVA2 )
1 ) State Institution National Antarctic Scientific Center, Ministry of Education and Science of
Ukraine, Tarasa Shevchenka blvd. 16, Kyiv-01601, Ukraine5 )
2 ) Institute of Hydrobiology, National Academy of Sciences of Ukraine, Heroiv Stalingradu 12,
Kyiv-04210, Ukraine
3 ) Institute of Fisheries and Marine Ecology, Konsulska str., 8, Berdiansk-71118, Ukraine

ABSTRACT
The brackish water calanoid copepod Eurytemora velox has become a widespread species in
inland fresh waters of Ukraine. Nowadays it is one of the dominant crustacean species in the littoral
zooplankton of the main Ukrainian rivers: Dnieper, Danube and Pivdenny Buh. We present a survey
on E. velox localities in, mainly, fresh waters of Ukraine, data on some quite isolated populations of
E. velox in the Carpathian region, as well as new records of the species in the upper parts of some
rivers. These data will provide a more complete view of the inland distribution of E. velox in Ukraine.

RÉSUMÉ
Le copépode calanoïde d’eau saumâtre Eurytemora velox est devenue une espèce largement
répandue dans les eaux douces d’Ukraine. Aujourd’hui, c’est l’une des espèces de crustacés
dominante dans le zooplancton littoral des principaux fleuves d’Ukraine : Dniepr, Danube et
Pivdenny Buh. Nous présentons ici une étude de la présence de E. velox, dans principalement les eaux
douces d’Ukraine, des données sur des populations isolées de E. velox dans la région des Carpathes,
ainsi que de nouvelles mentions de l’espèce dans les cours supérieurs de quelques rivières. Ces
données fourniront une vue plus complète de la répartition intérieure de E. velox en Ukraine.

INTRODUCTION

The genus Eurytemora Giesbrecht, 1881 constitutes a highly evolvable genus

with respect to salinity (Lee & Gelembiuk, 2008). It is represented in Ukraine by

4 ) Corresponding author; e-mail: larysa.samchyshyna@gmail.com

5 ) ”Kyiv” was formerly also known as ”Kiev”.

© Larysa Samchyshyna et al., 2021 | DOI:10.1163/9789004465565_005

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36 [276] LARYSA SAMCHYSHYNA, YULIA GROMOVA & KATERYNA ZORINA-SAKHAROVA

four species and one subspecies (Samchyshyna, 2011). One of these, Eurytemora
velox (Lilljeborg, 1853) is widely distributed in the brackish waters from the Arctic
Ocean and the Baltic Sea shore to south-eastern Europe, the Sea of Azov, and the
Caspian Sea (Vranovský, 1994). E. velox is believed to be a generalist species,
which constantly broadens its area of distribution, occupies new ecological niches,
and colonizes not only the saline and brackish water bodies, but freshwater habitats
away from the marine coasts as well (Pandourski & Evtimova, 2006).
In Ukraine, the geographic range of E. velox extends beyond the coastal waters
of the Black Sea and Sea of Azov and the lower streams of the main Ukrainian
rivers. The species inhabits a variety of water bodies from hypersaline salt marshes
to fresh waters (Lyashenko et al., 2018). In the Middle Dnieper River and its
tributaries, the species was found in the second half of the 20th century by many
authors (Melnikov, 1948; Monchenko, 1974; Polishchuk et al., 1978; etc.). The
species is widely distributed, especially in streamlets, channels, marshes, ponds,
and lakes of the Danube Delta, in waters with a large range of salinities (Spandl,
1926; Markovsky, 1955; Polishchuk, 1974; Parchuk, 1985; Kharchenko et al.,
2005; Zorina-Sakharova et al., 2014).
In this paper, we present a survey on the distribution of E. velox in, mainly,
fresh waters of Ukraine, data on some quite isolated populations of E. velox in the
Carpathian region, as well as new records of the species in the upper parts of rivers;
then, where possible, data on the species’ abundance are given.

MATERIAL AND METHODS

Zooplankton samples were collected using a hand-thrown, small plankton tow

net with a 100 μm mesh, during various sampling campaigns of both the Institute
of Hydrobiology (IHB AN) and the Institute of Zoology NASU (IZAN), in
1997-2019. The samples were preserved in 4% formaldehyde solution. Part of the
processed samples from earlier years (1956-1983) originate from the depository
of the IZAN and were collected by Prof. Vladyslav Monchenko and Dr. V.
Polishchuk. Samples from the Danube Delta were collected as a part of the IHB
AN project “Climatogenic restructuring of hydrobiont communities and its impact
on ecological situation and bioproductivity of EU cross-border rivers in Ukraine”.
Samples from Khmelnitsky nuclear power plant cooling pond were collected in
2014-2018 as a part of a multi-year monitoring programme led by Prof. O. Protasov
(IHB AN).

RESULTS

The abundance of Eurytemora velox in the Vita River varied from 7 up to

106 ind/m3 in 1997-1998 and from 30 up to 300 ind/m3 in 2011-2014, depending

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 EURYTEMORA VELOX IN UKRAINE [277] 37

on the season and on the speed of the water current. The abundance of E. velox in
this river reached 420 ind/m3 in 2016.
At the Ukrainian part of the Danube Delta, E. velox was recorded in the Kiliya
(= Chilia) Arm with 10-340 ind/m3 (2011, 2018, 2019) and in the following
Kiliya Delta branches: Starostambulskyi (= Stambulsky) with 10-1060 ind/m3
(2009); Ochakovskyi, 10-160 ind/m3 (2009, 2011, 2012); Bystryi, 10-400 ind/m3
(2009, 2010, 2011); Tsyganka, 40 ind/m3 (2009); Vostochnyi, 10-280 ind/m3
(2009, 2011); and some channels and lakes of the Small Tataru and Ermakov
Islands, 10-40 ind/m3 (2018). E. velox is wide spread also in the saline lagoons
of the Kiliya Delta, e.g., Solonyi Kut, 80-130 ind/m3 (2011); and Badika Kut,
40-80 ind/m3 (2011). Finally, we recorded species at the following freshwater
bodies of the delta: Anankin Kut Lake, 10-120 ind/m3 (2009, 2011); and the
lagoons Potapiv Kut, 120 ind/m3 (2009); Deliukiv Kut, 60-120 ind/m3 (2009,
2011); and Bystryi Kut, 90-150 ind/m3 (2011, 2014).
There are also some new records of E. velox in the middle and upper parts
of several Ukrainian rivers (fig. 1). In 2000 the species was found in the middle
part of the Salgir River (Crimean peninsula); in 2001 and 2014-2018 in some
areas of the Horyn River basin (33-50 ind/m3 in the cooling pond of Khmelnitsky
nuclear power plant in 2014) and at the Dniester River near Kamenetc-Podolsky.
In 2018-2019 the species was discovered in the Olexandriiske water reservoir and

Fig. 1. Map depicting collection sites of Eurytemora velox (Lilljeborg, 1853) in fresh and brackish
waters in Ukraine. Circles represent sites where samples from existing collections were processed
(1953-1981). Triangles represent personal observations (1998-2019). Rhombi represent literature
data.

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38 [278] LARYSA SAMCHYSHYNA, YULIA GROMOVA & KATERYNA ZORINA-SAKHAROVA

in the Pivdenny Buh River. In the region of the Eastern Carpathian Foothills of
Ukraine, E. velox was recorded in 2001 in the pond between Drogobych and
Boryslav, while according to stored samples from 1976-1978, deposited at IZAN,
the species had earlier been absent in that watershed.
Between other records from processed archived samples are: Dnieper River
near Kyiv [= Kiev] and Kaniv (1953-2019), Prypiat River (in 1963, 5 ind/m3 ,
1982, 2000-2003, in 2010-2014, 10 ind/m3 ), Kahovsky water reservoir (1962),
Dnieper-Bug estuarine area (1966), Black Sea Biosphere Reserve (1983), Staro-
Krymske water reservoir (1974), Symferopilske water reservoir (2000), Inhuletc
River (1973), Holtva River (1956), Vorona River (1956), Desna River (1956,
2017), Psel River (1997-1998, 10-600 ind/m3 ), Vovcha River (1971), Teteriv River
(2000), Stryzhen River (2000), Kuhurluy Lake (1966, 1981), Kytai Lake (1966),
Katlabuh Lake (1963, 1981), Kagul Lake (1964), Berezansly Lake (1979), and
lower part of Prut River (1965). Some additional old records in the Danube Delta
are from the Zavodnitske branch (1966), Zhebriiansky Lagoon (1966), Shabash
Kut Lagoon (1965), Taranov Kut Lagoon (1965), Lazorkin Kut Lake (1965) and
Poludenoe branch (1965).

DISCUSSION

The idea of a strong relationship between the level of penetration of inverte-

brates into continental waters and the adaptation to fresh water on the one hand,
and the ancientness of the taxon, on the other, was expressed by Robert Gurney
(Gurney, 1931). The family Temoridae includes four genera, of which only one is
marine: Temora Baird, 1850. The other three are brackish and freshwater genera:
Eurytemora Giesbrecht, 1881, Heterocope Sars, 1863 and Epischura Forbes, 1882,
of which Eurytemora at all events and in all aspects appears to be a relatively re-
cent immigrant into fresh water. This genus is considered as a neo-limnetic form,
according to the classification of Starobogatov (1970).
Siewerth (1926) found Eurytemora velox in Ukraine for the first time at
the mouth of the Seversky Donetc River. Later Markovsky (1928) revealed
the presence of the species in the Inhuletc River and considered this species
as a remnant of the Post-Pliocene marine basin. The Black Sea area has a
complex evolution from subduction to collisions during the closure of the Neo-
Tethys Ocean, and was exposed to multiple water-level fluctuations in successive
glaciations (Hippolyte et al., 2018). Markovsky assumed that E. velox, together
with E. affinis (Poppe, 1880), Heterocope caspia Sars, 1897 and Calanipeda
aquaedulcis Krichagin, 1873, are adaptive relicts that adapted well to changing
environmental conditions and will be extending their range over the relict area into

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 EURYTEMORA VELOX IN UKRAINE [279] 39

fresh waters. In spite of the fact that subsequent records of E. velox in Ukraine
were made in the range of marine fluctuations since the Last Glacial Maximum,
Sabaneev (1930) also supported the suggestion about future migration of this
species up along rivers.
Outside the ancient marine transgressions, E. velox were recorded many times
in successive years (Taran, 1931; Melnikov 1948). In the Middle Dnieper basin
near Kiev, the species was mentioned by many authors (Travyanko & Tseeb,
1967; Radzimovsky & Polishchuk, 1970; Samchishina, 2001; etc.). In the Dnieper
tributaries, the Psel, Desna, Prypiat, Holtva and Vorskla Rivers, this species was
mentioned by Monchenko (1974). That last-mentioned author concluded that the
absence of the species in the upper parts of the rivers indicates relatively recent
migration into fresh waters. In the Prypiat River basin, E. velox shows high
abundance in summer in the cooling pond of the Khmelnitsky nuclear power plant
on the Hnylyi Rih River, the inflow of the Horyn River (Protasov et al., 2011).
Elsewhere in the Eastern Europe, the species was also found, e.g., in Hungary
(Forró & Gulyás, 1992) and Slovakia (Vranovský, 1994) in 1991 for the first time,
and has recently largely expanded its range in those countries (Hudec & Illyová,
1998; Kiss et al., 2015). Simultaneously, in North America, another species of
the genus, Eurytemora carolleeae Alekseev & Souissi, 2011 (formerly considered
part of the Eurytemora affinis species complex) is, reportedly, recently invading the
Laurentian Great Lakes. It has been in the Great Lakes since the 1950s (Lee, 1999),
and is continuously being reported from new inland localities since that time. It
was commonly accepted that the species has spread by shipping traffic through
the discharge of ballast waters. Furthermore, the species recently is invading also
the Gulf of Finland in Europe (Sukhikh et al., 2019). Similar biological invasions
of Ponto-Caspian Eurytemora species are observed in the Volga River in Russia.
Thus, Eurytemora caspica Sukhikh & Alekseev, 2013 (also recently separated
from the E. affinis species complex) had invaded the Volga, Saratov and Kuibyshev
Reservoirs on Volga River until the 2010s. The species was next revealed in the
Kama Reservoir (2700 km removed from the Caspian Sea) for the first time in
2016, and has become a dominant species in the zooplankton in summertime in
most of those reservoirs (Lazareva, 2019).
We have assumed that the recent success of the natural invasion into fresh
water by the copepod E. velox in Ukraine can be interpreted as a result of a
mainly passive route (e.g., transit of undigested eggs in fish’s guts, shipping traffic,
etc.), however, the active way of range extension, i.e., by actively moving through
swimming might also be considered. Obviously, this oligohaline species is able to
cope well with new environments and with freshwater conditions, through being
equipped with effective adaptations for the required osmoregulation. We have
noticed euryoxybiontic abilities of the species in wintertime. Finally, acquiring

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40 [280] LARYSA SAMCHYSHYNA, YULIA GROMOVA & KATERYNA ZORINA-SAKHAROVA

the ability to survive in fresh water, might be a beginning of speciation for

populations entering the fresh waters, since contact between populations inhabiting
upper reaches of rivers may become limited. Accordingly, a scenario of allopatric
evolution might be starting to develop. Evidently, further research should be
conducted to explain and monitor this phenomenon, specifically also through using
molecular genetic tools.

ACKNOWLEDGEMENTS

The Khmelnitsky nuclear power plant cooling pond zooplankton samples

were investigated through courtesy of Prof. O. Protasov (IHB AN), and the
abundance data were kindly provided by the group of technical hydrobiology.
Sampling campaigns at the Danube Delta were funded by Program 6541230 (No.
0118U002287) of the IHB AN. Existing preserved samples were provided by the
depository of IZAN, for processing.

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First received 29 October 2019.

Final version accepted 25 November 2019.

For use by the Author only | © 2021 Larysa Samchyshyna et al.