Scientific Papers. Series D. Animal Science. Vol. LXIII, No. 1, 2020
ISSN 2285-5750; ISSN CD-ROM 2285-5769; ISSN Online 2393-2260; ISSN-L 2285-5750
Alosa immaculata Bennet, 1835: A SHORT REVIEW OF THE SPECIES
AND ITS BIOLOGY
Mihaela MOCANU, Lucian OPREA, Anca Nicoleta CORDELI (SĂVESCU), Mirela CREȚU
“Dunărea de Jos” University of Galați, 47 Domnească Street, 800008, Galați, Romania
Corresponding author email: mocanumihaela64@gmail.com
Abstract
Currently, in the continental and marine waters of the Earth, there are over 33,400 species of fish. Over time,
naturalists, and specialists in ichthyology have proposed several variants of systematic classification of fish. Thanks to
new methods and advanced scientific research equipment, 300-500 new species are discovered every year. This
explains the fact that in just 20 years (1998-2017), 7,436 species were described. In 2004, the scientific organization
Integrated Taxonomic Information System published ITIS Report, in which a complete taxonomic hierarchy of the genus
Alosa is presented. Alosa immaculata (Pontic shad), is an important fish species in the ecological system of the
Danube-Delta-Black Sea, with a significant economic value due to the high catches and to the nutritional qualities of
meat. The knowledge of the essential elements regarding the biology and the exploitation of the species contributes to
ensuring the necessary information for the conservation of the species and the management of the stocks. The current
level of knowledge on the biology of the Pontic shad is known from the researches of the Romanian and Russian
specialists from the years 1960-1970, but in the last 30 years, numerous changes have occurred in the environmental
and exploitation conditions of the species. In this context, the aim of this paper is to present as updated information as
possible regarding the biology and exploitation of the Pontic shad, under the new environmental conditions, to meet the
current objective of species conservation in the requirements of the sustainable exploitation of the stocks.
Key words: Alosa immaculata (Pontic shad), biology, stocks.
phylogenetic relationships within the genus
Alosa, resulting in systematic and taxonomic
uncertainty, which may undermine the
establishment of adequate conservation
measures (Faria et al., 2006).
The genus Alosa is present in the Northern
hemisphere of the Earth. Many of the species
representing this genus are in the Atlantic,
Mediterranean, Black, and Caspian. In the
Danube River and the Black Sea, are found
Alosa immaculata (Bennett, 1838) - Pontic
shad, Alosa tanaica (Grimm, 1901) - Azov
shad, synonymous with subspecies Alosa
caspia nordmanni (Antipa, 1906), and Alosa
maeotica (Grimm, 1901) - Black Sea shad
(Năvodaru et al., 2014).
According to IUCN Red List 2008, Alosa
immaculata is a vulnerable species, and the
current threat of the species is overfishing, at
sea, and in the rivers during the migration runs,
which is causing a population decline of
unknown levels (Figure 1).
INTRODUCTION
The Danube River covers a length of 2,857 km.
Human activities have significantly influenced
the flow of the Danube through flood
prevention, navigation activities. The most
important dams in the Danube at a distance of
943 km from the river, the Iron Gate I, and the
Iron Gate II have formed an accumulation lake,
which in turn represents obstacles for migratory
fish species and Pontic shad.
Besides the negative impact of the dams and
the regulation of the river flow, the common
stocks of Pontic shad in the region of the Lower
Danube have been affected by overfishing and
the pollution. Some of the obstacles
encountered in the standard effective
management of these fish stocks may be the
lack of harmonization and coordination of
management.
Clupeidae is one of the world’s most
commercially essential families of fish. Despite
their importance, little is known about the
516
Figure 1. The geographic range of Alosa immaculata
(IUCN, Red List 2008; Freyhof & Kottelat, 2008)
varying depths and great distances from the
coast of Ukraine, Romania, and Bulgaria
(Bănărescu, 1964).
The Alosa immaculata is a marine, migratory,
anadromous species, migrate for spawning
from the Black Sea to the Danube River,
and the average weight is around 240 g and the
total length of 31 cm (Lenhardt et al., 2016)
(Figure 2).
In the European Union, according to the Annex
II of the Habitats Directive 92/43/EEC, the
Pontic shad is a protected species, requiring
protection under the Natura 2000 Network and
site management, which is under its ecological
requirements. Also, the species is listed in
Annex IV, which obliges the Member States to
ensure that their exploitation is compatible with
maintaining a favorable conservation status. At
the national level, the species is protected by
the GEO no. 57/2007.
Pontic shad is a commercially important fish of
the Danube Delta and the countries of the
Lower Danube Region (Romania, Ukraine, and
Bulgaria) (Ciolac & Patriche, 2004).
In this context, the purpose of this paper is to
review some aspects regarding the biology and
current status of the Alosa immaculata
population along the Danube River and the
Black Sea.
Taxonomy.
Distribution.
Morphology.
Ecology and biology of the Alosa immaculata
Pallas (1811) made the first reports of the
Pontic shad, under the name of western
European shad, Clupeonela pilchardus. In the
year 1835, Bennett describes the species under
the name of Alosa immaculata - Pontic shad.
Alosa immaculata is an anadromous fish
species that inhabit the Black and Azov Sea, at
Figure 2. Alosa immaculata (Photo original)
517
The reproduction of Alosa immaculata takes
place in the Danube, up to 80 km, between
Brăila and Călarași county. Still, in the past,
reproductive shad migrate upriver up to 1000
km, reaching only the Iron Gates II (863.55
km). The current of the Danube carries larvae
and fry of Pontic shad to the sea (Cristea and
Cristea, 1958).
Sexual maturation occurs at the age of 3 and 4
years old, and only a few individuals spawn
two seasons (Năvodaru & Năstase, 2014;
Țiganov et al., 2016).
Usually, a Pontic shad spawns only once or
twice in its lifetime (Ciolac, 2004). Most
spawning in the Danube River occurs between
180 and 743 km upstream (Kolarov, 1985;
Schmutz, 2006). Individuals of two years have
a low contribution among breeders being
represented by individuals with growth and
maturation accelerated. Also, fish with ages
between 5 and 7 years are rarely found during
migration (Năvodaru, 1997).
The migration is dependent on water
temperature, and according to some authors
begins in February - March when water
temperatures reach 5-6˚C, with the highest rate
in April, at water temperatures of 9-13˚C
(Pavlov, 1953; Năstase et al., 2018). Năvodaru,
in 1996 and 1998, said that the Alosa
immaculata migration in the Danube River
begins in spring at water temperatures of 37.5˚C, peaks in April and May between 9-17˚C,
and is finished in June and July when water
temperatures reach 22-26˚C.
Another critical factor that affects the entry of
the Alosa immaculata into the Danube is
turbidity and water level. According to
Năvodaru (1997), higher water turbidity, and a
high water level in the Danube slow down fish
migration.
The sex structure of the migration population
can vary depending on the seasonality and
migration period. Several studies reported
female dominance with the aging and a
decrease of the males (Năvodaru, 1997;
Năvodaru and Năstase, 2014; Țiganov et al.,
2016).
Generally, the duration of the Danube
shad migration is estimated at 100-150 days
(Table 1).
Table 1. Size structure of Alosa immaculata in 2009, 2013, 2014 and 2016
Catch
area/year
Danube
River, 2009
Total weight
(g)
100-400 g
Average 276.72 g
The age of the
fish caught
Sex ratio
(M/F)
24-39 cm
Average 31.11 cm
-
-
Ibănescu et
al., 2016
Black Sea
coast, 2013
Spring
season
(March,
April),
summer
(June, July)
and autumn
(September)
163-442 g
Average 248 g
25.2-31.2 cm
Average 28.2 cm
2 years -16%
3 years - 42%
4 years - 37%
5 years - 5 %
0.62,
females
dominant
Țiganov et
al., 2016
Mouth of
Danube
River, 2014
January –
May
163- 422 g
Average 289 g
25.3 – 35.7 g
Average 31.1 cm
3 years – 46%
4 years – 50%
5 years - 4 %
Năvodaru
and
Năstase,
2014
February –
August
102-435
Average 236.8
22.7-36
Average 28.7 cm
Dominance
3-4 years
0.55,
64% females,
36% of
males
0.51,
females
dominant
Mouth of
Danube
River, 2016
Catch period
April - June
Total length (cm)
Generally, food consists of 70-75% of adult
fish (Engraulis, Clupeonella, Sprattus) in the
sea and Cyprinids in the Danube River, the rest
being shellfish (Crangon, Upogebia, Idotheia),
and other organisms according to their
abundance and availability (Bănărescu, 1964;
Cautiș et al., 1957).
References
Năstase et
al., 2016
Alosa immaculata has an elongated body,
compressed laterally, and the mouth is
terminal, broad, slightly oblique upwards. Well
developed adipose eyelid, leaving a narrow,
elliptical-shaped vertical opening. Laterally
compressed abdomen from the tip of the snout
to the base of anal. The hull is evident,
518
especially between ventral and anal. The dorsal
fin is located towards the middle of the body,
and the anal fin is located far behind. The
backside of the body is colored in blue-green,
silvery flanks, and sometimes white and
sometimes darker head, fins are colorless
(Antipa, 1909; Bănărescu, 1964).
Biological peculiarities of the species make it
vulnerable to different threats, but the major
ones are overfishing and loss of spawning
grounds.
In the context of fishery management that takes
ecological and ecosystem considerations,
protected areas are highly relevant.
Recent data on shads stocks, distribution,
population parameters, and genetics in the
Black Sea area urgently are needed for the
species conservation and management issues.
experimental fishing and from the commercial
fishing.
RESULTS AND DISCUSSIONS
In Romania, Alosa immaculata is a fish with
high economic and socio-cultural value for the
communities. Due to the highest nutritional
quality of Alosa immaculata (has a big content
of lipids 18-22% and a higher content of
soluble vitamins), this fish is usually consumed
in the Lent by the Christian population.
The fishery has a commercial value of about
1.5 million euros, with annual average catches
of 200-500 tons. According to Năvodaru and
Năstase (2014), Alosa immaculata has a
cyclical evolution of catches, with minimums
or maximums at 10-11 years, during the period
1960-1998, the minimum absolute was 200 t,
and the maximum of 2,400 t.
Between years 2007 and 2010, according to
data provided from FAO (2016) statistics
regarding the catches in the Danube and the
Black Sea, it is observed a dramatic decrease in
2014, of the level of catches in the Black Sea to
approximately of 2 tons, while catches from the
Danube in the last years are maintained around
400 tons per year (Figure 3).
Unfortunately, as much as the interest for the
exploitation of the species, it is higher, as well
as the danger of the drastic decline of the
Danube and Sea populations Black is bigger.
Also, the dramatic fall of Pontic shad migration
in the Danube river for reproduction is a real
issue that evidently should concern interested
persons and involved institutions, companies,
and organizations from both commercial and
ecological point of view.
In this context, there it is recommended some
management measures for the conservation of
the Alosa immaculata, such as the introduction
of bans, for example closing of certain fishing
areas and seasons or protection of fish by
regulating fishing methods and instruments for
the Black Sea and Danube (Țiganov et al.,
2016).
Knowledge of the essential elements of the
biology and exploitation of the species
contributes to ensuring necessary informatio for
species conservation and management of
stocks.
MATERIALS AND METHODS
The Danube is the most important stream in
Romania. The Danube sector under study is of
particular importance for fish populations, as it
is a central wetland type (Ibănescu et al., 2016).
Several representative areas for each habitat
type were chosen for fish sampling. The
evidence comes from scientific and commercial
fishing.
In order to study the biological parameters of
the pontic shad species from the collected
samples, samples were taken, which were
analyzed in the laboratory.
Special fishing gear was used for Danube
Pontic shad: driftnets and suitable boats
operated by fishermen. The material was
collected from different parts of the Danube, in
the spring season (March, April), summer
(June, July) and autumn (September) (Țiganov
et al., 2016).
The main physiological parameters recorded
for each individual were: total length,
individual mass, age, and sex. Weight was
determined in grams. The age was determined
by reading the annual growth rings on the
scales. The sex of each individual was
determined by dissection.
The study of the species Alosa immaculata
consisted in an ichthyological study of all the
samples collected during the studied period,
using both the samples collected from the
519
Figure 3. The level of catches of Alosa immaculata (Source FAO Statistics, 2016)
Control of fish stocks, including shad, is
achieved by the interaction of three key
compartments species - environmental exploitation.
For better management, further investigation
and more collaboration among countries in the
Lower Danube River Region is needed:
monitoring of stocks, studies on factors that
influence change in shares, molecular genetic
investigation of migrants, determination and
protection of spawning and nursery places in
the Danube River and its floodplains and delta
as well as the costal shelf of the Black Sea.
these river systems, as well as the adverse
effects on the species that recur in the lower
parts of this system are not very clear.
The monitoring of the migrating population of
Alosa immaculata into the Danube River
deserves further empirical study. Additional
research is needed to quantify any changes in
the number of individuals and their biometric
variables and to validate if an upward trend in
abundance is happening. The continuation of
the study should provide the evidence
necessary to determine possible changes in the
conservation state of this species of community
interest.
CONCLUSIONS
ACKNOWLEDGEMENTS
Even if we consider the adverse changes of
some
ecological
aspects
related
to
environmental factors such as lower water
level, water temperature, and pollution that
could affect the success of the Danube shad
reproduction. The most crucial cause of the
decreasing of the stocks is the overfishing,
mainly in Danube Delta area, which provides
about 90% of total amount of capture in
Danube River.
There are visible decreases in the Pontic shad
population. The research is carried out only
occasionally on the territory of Romania and
Bulgaria. Changes resulting from human
intervention in river systems affect migratory
fish. Dams and river engineering structures will
affect a species such as Alosa immaculata
because their breeding areas have located in
The principal author of the article thanks to
“Dunărea de Jos” University of Galați which
through the University Degree Program, the
doctoral studies contract has supported the
achievement.
REFERENCES
Antipa, G. (1909). The ichthyological fauna of Romania,
XVI. Bucharest, RO: Vasile Adamachi Fund
Publications, Romanian Academy Publishing House
(In Romanian).
Bănărescu, P. (1964). Pisces-Osteichthyes R.P.R. Fauna,
XVIII. Bucharest, RO: R.P.R. Academy Publishing
House (In Romanian).
Cautiș, I., Iliescu, M., Maximov, A. (1957). Caspialose
fund problem form Black Sea. Bul ICP, XV, 47-56.
520
Ciolac, A. (2004). Migration of fishes in Romanian
Danube River. Applied Ecology and Environmental
Research, 2(1), 143-163.
Ciolac, A., Patriche, N. (2004). Structure of Danube
Shad (Alosa pontica, Eichwald 1838.) spawner flocks
migrating for reproduction in Danube River. Appl
Ecol Env Res., 2(2), 53-58.
Cristea, A., Cristea, E. (1958). Some resulted obtained
from prolificity of the Pontic shad (Alosa pontica).
Bul ICP, XVII (3), 78-85.
FAO (2016). Fishery and Aquaculture Statistics. FAO
yearbook, 125(7), Part 1.
Faria, R., Weiss, S., Alexandrino, P. (2006). A molecular
phylogenetic perspective on the evolutionary history
of Alosa spp. (Clupeidae). Molecular phylogenetic
and evolution, 40, 298-304.
Freyhof, J., Kottelat, M. (2008). Alosa immaculata. The
IUCN Red List of Threatened Species 2008:
e.T907A13093654. Retrieved March 12, 2020, from
https://dx.doi.org/10.2305/IUCN.UK.2008.RLTS.T9
07A13093654.en.
GEO no. 57/2007 on the regime of natural protected
areas, the conservation of natural habitats, wild flora
and fauna.
Habitats Directive 92/43/EEC Council Directive
92/43/EEC on the conservation of natural habitats
and of wild Fauna and flora.
Ibănescu, D., Popescu, A., Nica, A. (2016). Estimation of
growth and mortality parameters of the Pontic shad
(Alosa immaculata Bennett, 1835) in Romanian
section of the Danube river. Lucrări Ştiinţifice-Seria
Zootehnie, 285-289.
Kolarov, P. P. (1985). Biological characteristics and
population dynamic of anadromous fish specie. PhD
thesis, Varna: Institute for Fish Resources.
Lenhardt, M., Višnjić-Jeftić, Ž., Navodaru, I., Jarić, I.,
Vassilev, M., Gačić, Z., Nikčević, M. (2012). Fish
stock management cooperation in the Lower
Danube region: A case study of sturgeons and
Pontic shad. In: Lagutov V. (eds) Environmental
Security in Watersheds: The Sea of Azov. NATO
Science for Peace and Security Series C:
Environmental Security, Dordrecht, NL: Springer
Publishing House.
Lenhardt, M., Navodaru, I., Vassilev, M., Kalauzi, A.,
Regner, S., Višnjić-Jeftić, Ž., Tošić, K., SmederevacLalić, M. (2016). Model of the Pontic shad Alosa
immaculata (Bennet, 1835) and anchovy Engraulis
encrasicolus (Linnaeus, 1758) catch in the Danube
river and Black Sea for the period 1920-2008. Acta
zool. bulg., 68(4), 557-561.
Năstase, A., Năvodaru, I., Cernisencu, I., Țiganov, G.,
Popa, L. (2018). Pontic shad (Alosa immaculata)
migrating upstream the Danube river and larval drift
downstream to the Black Sea in 2016. Scientific
Annals of the Danube Delta Institute, 23, 57-68.
Năvodaru, I. (1996). Exploitation of Alosa Pontica in the
Danube Delta, Romania. - In: Co W x i. G. (Ed.):
Stock Assessment in Inland Fisheries, 448-453.
Oxford, UK: Fishing News Books Publishing House.
Năvodaru, I. (1997). The evolution of the Pontic shad
effectives under conditions of ecological of the river
and their maintenance measures. PhD Thesis,
“Dunărea de Jos” University of Galați.
Năvodaru, I. (1998). Migrația scrumbiei de Dunăre
(Alosa pontica) și factorii de mediu. Analele
Științifice ale Institutului Delta Dunării-1997, VI(1),
149-152.
Năvodaru, I., Năstase, A. (2014). New data on pontic
shad (Alosa immaculata Bennet 1835) migration and
drifting larvae in Danube River, Deltaica, 3: 48 pp.
In: Torok, L. ed., Noi date asupra prezenţei marilor
peşti migratory anadromi în Marea Neagră - Zona
marină a Rezervaţiei Biosferei Delta Dunării (New
data on presence of the great anadroumous migratory
fishes in Black Sea - marine zone of Danube Delta
Biosphere Reserve), ISSN 2286–0789; doi:
10.14590/DDI02.D03.
Pallas, P., S. (1811). Zoographia Rosso-Asiatica, sistens
omnium animalium in extenso Imperio Rossico et
adjancetibus, domicilia, mores et descriptiones,
anatomen atque icones plurimorum, III, Petropoli:
Academiae scientarium impress Publishing House.
Pavlov, P. I. (1953). Biologhiceskaia I promislovaia
harakteristika nerestovo stada dunaiskoi seldi. In:
Dunaiska seldi i biologhiceskie osnovi eio promisla,
118-174. Kiev, USSR: Acad. Nauk. Publishing
House.
Schmutz, S. (2006). Assessment of the potential
transboundary effects of the construction of the 270
Bystre Deep Water Navigation Channel on fish and
fisheries. Report to the ESPOO Inquiry Commission
Vienna, 56.
Țiganov, G., Năvodaru, I., Cernișencu, I., Năstase, A.,
Maximov, V., Oprea, L. (2016). Preliminary data on
the studies Alosa immaculata in Romania marine
waters. Scientific Annals of the Danube Delta
Institute, 22, 141-148.
521