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Conchological Variability of Anadara Inaequivalvis (Bivalvia, Arcidae) In the Black–Azov Sea Basin

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Conchological Variability of Anadara inaequivalvis (Bivalvia, Arcidae) in the Black-Azov Sea Basin. Anistratenko, V. V., Anistratenko, O. Yu., Khaliman, I. A. - An alien species in the Black Sea and the Sea of Azov - Anadara inaequivalvis (Bruguiere, 1789) - was recently discovered to have a wide range of shell variability. From the investigated samples (over 900 valves) six basic types of the shell commissural opening were defined; they are not, however, discrete conditions. Th e presence of intermediate variants reveals a gradual (continuous) character of the shell variability and indicates that all the samples investigated belong to the same species. Th e variation of some other Anadara characteristics is also discussed, including: quantity and ornamentation of the ribs on the surface of the valves, quantity of chevrons on the ligament area, shape of the hinge plate and quantity of hinge teeth. A comparison of conchological characteristics of Anadara from the Black-Azov Sea Basin with A. inaequivalvis from southeast India shows that ranges and patterns of shell variability of Azov-Black Sea Anadara correspond to variability of A. inaequivalvis from this species type locality - Coromandel Coast of India.
Localities in the Black Sea and the Sea of Azov where the samples were collected: 1 — Cordon “Morskoj”, the Black Sea Biosphere Reserve (BSBR), Gola Prystan District, Kherson Region, Ukraine (46°07 ́54 ̋ N, 32°13 ́39 ̋ E, 2011 and 2012, coll. V. V. Anistratenko and O. Yu. Anistratenko); 2 — Fedotova Spit in the vicinity of Kirillovka settlement, Akimovka District, Zaporizhzhya Region, Ukraine (46°19 ́40 ̋ N, 35°20 ́03 ̋ E, 2005, 2006, 2012, coll. V. V. Anistratenko, O. Yu. Anistratenko and I. A. Khaliman); 3 — Yurkino settlement, Lenino District, Crimea, Ukraine (45°25 ́24 ̋ N, 36°33 ́15 ̋ E, October 12, 2003, coll. V. V. Anistratenko and O. Yu. Anistratenko); 4 — Pichory village, Galy District, Abkhazia (42°26 ́57 ̋ N, 41°32 ́30 ̋ E, June 22, 1989, coll. V. V. Anistratenko); 5 — Chakvy settlement, Adzharia (41°44 ́05 ̋ N, 41°43 ́57 ̋ E, June 21, 1989, coll. V. V. Anistratenko). Рис. 1. Места сбора материала в Чёрном и Азовском морях: 1 — кордон «Морской», Черноморский биосферный заповедник, Голопристанский р-н, Херсонская обл., Украина (46°07 ́54 ̋ N, 32°13 ́39 ̋ E, 2011 и 2012 гг., сбор В. В. Анистратенко, О. Ю. Анистратенко); 2 — Федотова коса возле пос. Кирил- ловка, Акимовский р-н, Запорожская обл., Украина (46°19 ́40 ̋ N, 35°20 ́03 ̋ E, 2005, 2006, 2012 гг., сбор В. В. Анистратенко, О. Ю. Анистратенко, И. А. Халиман); 3 — пос. Юркино, Ленинский р-н, АР Крым, Украина (45°25 ́24 ̋ N, 36°33 ́15 ̋ E, 12 октября 2003 г., сбор В. В. Анистратенко, О. Ю. Анистратенко); 4 — с. Пичори, Галийский р-н, Абхазия (42°26 ́57 ̋ N, 41°32 ́30 ̋ E, 22 июня 1989 г., сбор В. В. Анистратенко); 5 — пос. Чакви, Аджария (41°44 ́05 ̋ N, 41°43 ́57 ̋ E, 21 июня 1989 г., сбор В. В. Анистратенко).
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UDC 594.1:591.157(262.54+560.6)
CONCHOLOGICAL VARIABILITY
OF ANADARA INAEQUIVALVIS BIVALVIA, ARCIDAE
IN THE BLACKAZOV SEA BASIN
V. V. Anistratenko1, O. Yu. Anistratenko1, 2, I. A. Khaliman3
1Schmalhausen Institute of Zoology, NAS of Ukraine,
vul. B. Khmelnitskogo, 15, Kyiv, 01601 Ukraine
E-mail: anistrat@izan.kiev.ua
2Institute of Geological Sciences of NAS Ukraine,
O. Gontchar str., 55-b, Kyiv, 01601 Ukraine
E-mail: anistrat@rambler.ru
3Tavrichesky State Agrotechnological University,
B. Khmelnitsky str., 18, Melitopol, 72319 Ukraine
E-mail: khali@ukr.net
Conchological Variability of Anadara inaequivalvis (Bivalvia, Arcidae) in the Black-Azov Sea Basin.
Anistratenko, V. V., Anistratenko,O. Yu., Khaliman, I. A. — An alien species in the Black Sea and the
Sea of Azov — Anadara inaequivalvis (Bruguiere, 1789) — was recently discovered to have a wide range
of shell variability. From the investigated samples (over 900 valves) six basic types of the shell commissural
opening were de ned; they are not, however, discrete conditions.  e presence of intermediate variants
reveals a gradual (continuous) character of the shell variability and indicates that all the samples investi-
gated belong to the same species.  e variation of some other Anadara characteristics is also discussed,
including: quantity and ornamentation of the ribs on the surface of the valves, quantity of chevrons on
the ligament area, shape of the hinge plate and quantity of hinge teeth. A comparison of conchological
characteristics of Anadara from the Black–Azov Sea Basin with A. inaequivalvis from southeast India
shows that ranges and patterns of shell variability of Azov-Black Sea Anadara correspond to variability of
A. inaequivalvis from this species type locality — Coromandel Coast of India.
Key words: Bivalvia, Anadara, shell morphology, the Black Sea, the Sea of Azov.
  Anadara inaequivalvis (Bivalvia, Arcidae)  -
.  . .,  . .,  . . — Показан широкий диа-
пазон изменчивости раковины Anadara inaequivalvis (Bruguière, 1789) — вида-вселенца в Азово-
Черноморском бассейне. В изученном материале (более 900 створок) выделены 6 основных форм
комиссурального просвета раковины, которые, однако, не являются дискретными состояниями.
Наличие промежуточных вариантов свидетельствует о плавной (непрерывной) изменчивости и
принадлежности всех изученных популяций к одному виду. Обсуждается варьирование некото-
рых других признаков раковины Anadara: количество и скульптированность рёбер на поверхно-
сти створок, количество шевронов на лигаментной площадке, форма замочного края и количество
зубов замка. Сопоставление морфологических характеристик раковины Anadara из Чёрного и
Азовского морей и A. inaequivalvis из юго-восточной Индии подтверждает, что границы и харак-
тер изменчивости моллюсков рода Anadara, обитающих в азово-черноморском бассейне, вполне
соответствуют изменчивости A. inaequivalvis из типовой местности — Коромандельского побере-
жья Индии.
Ключевые слова: Bivalvia, Anadara, морфология раковины, Чёрное море, Азовское море.
Introduction
Over 40 years ago in the Mediterranean Sea at the Italian coast o Ravenna a new alien bivalve mollusk
of the genus Anadara Gray, 1847 has been registered.  e species got into adjacent areas soon (Ghisotti,
1972; Rinaldi, 1972; Ghisotti, Rinaldi, 1976; Nolf, 2010). Mollusks of this group belong to the family Arcidae
Lamarck, 1809 (subfamily Anadarinae Reinhart, 1935) and are, mainly, inhabitants of the tropical and
subtropical seas.
Vestnik zoologii, 48(5 ): 457–466, 2014
DOI 10.2478/vzoo-2014-0054
458 V. V. Anistratenko, O. Yu. Anistratenko, I. A. Khaliman
It is considered that in the Black Sea this mollusk was found for the  rst time in 1981 in coastal areas of
Bulgaria and on a shelf of Romania (Zolotarev, Zolotarev, 1987). However, according to data published by
M.I.Kiseleva (1992), the  rst  nding of this species in the Black Sea should be dated 1968, i. e. practically the
same time as that in the Adriatic Sea. It is possible that introduction of Anadara could happen independently to
di erent basins and/or di erent sites within the same basin suitable for its habitation.  e colonization history
of the Black-Azov Sea Basin by this mollusk was reviewed previously (Anistratenko, Khaliman, 2006); so far
Anadara inhabits in the Black Sea along the entire coast (Sahin et al., 2009), and in the Sea of Azov it occupied
its southern, western and partially northern sectors (Anistratenko et al., 2011).
In the majority of publications (Rinaldi, 1985; Poppe, Goto, 1993; Anistratenko, Khaliman, 2006; Sahin et al.
2009; Finogenova, 2011) the Mediterranean Anadara as well as those in the Black Sea and the Sea of Azov is assigned
to A. inaequivalvis (Bruguière, 1789). Meanwhile, recent substantial data on conchology of A. inaequivalvis from
the southeast coast of India, a district close to the type locality of the species concerned (Lutaenko, 2006), were
published. Comparison of some conchological characters of Anadara from the Mediterranean and the Black Seas
makes the author to doubt on the belonging of these mollusks to the true A. inaequivalvis.
e solution of the problem of Anadara species identi cation that immigrated to the Mediterranean Basin
depends on the study of the variability of its shell in new to this species conditions, in particular. As far as it can
be concluded from the publications, the majority of Anadarinae are not well studied in this respect (Lutaenko,
1993, 2006). Only few works are devoted to variability of Anadara, for instance, in the northwestern part of the
Sea of Japan (Lutaenko, 2002).
e present study focuses on shell variability of Anadara inhabiting the Black-Azov Sea Basin. For
comparison, the materials by K. A. Lutaenko (2006) on conchology of A. inaequivalvis and some related
nominal species of Anadara from the southeast India are discussed.
Material and methods
Material for the present study was collected from coastal casts of the Black Sea and the Sea of Azov from
1989 to 2012 ( g. 1).  e Black Sea material is presented by 17 specimens from the coast of the Caucasus near
Pichory village (Gali District, Abkhazia), three specimens are from the settlement Chakvi (Adzharia); about
800valves were collected within the borders of cordon “Morskoy” of the Black Sea Biosphere Reserve (BSBR,
Golopristan’ District, Kherson Region, Ukraine).  e main material from the Sea of Azov (about 500speci-
mens) was collected along the sea side of Fedotova Spit (Kirillovka settlement, Akimovka District, Zapori-
Fig. 1. Localities in the Black Sea and the Sea of Azov where the samples were collected: 1— Cordon “Morskoj”,
the Black Sea Biosphere Reserve (BSBR), Gola Prystan District, Kherson Region, Ukraine (46°07´54˝ N, 32°13´39˝
E, 2011 and 2012, coll. V. V. Anistratenko and O. Yu. Anistratenko); 2 — Fedotova Spit in the vicinity of Kirillovka
settlement, Akimovka District, Zaporizhzhya Region, Ukraine (46°19´40˝ N, 35°20´03˝E, 2005, 2006, 2012, coll.
V. V. Anistratenko, O. Yu. Anistratenko and I. A. Khaliman); 3 — Yurkino settlement, Lenino District, Crimea,
Ukraine (45°25´24˝ N, 36°33´15˝ E, October 12, 2003, coll. V. V. Anistratenko and O.Yu. Anistratenko); 4
Pichory village, Galy District, Abkhazia (42°26´57˝ N, 41°32´30˝ E, June 22, 1989, coll. V.V.Anistratenko); 5—
Chakvy settlement, Adzharia (41°44´05˝ N, 41°43´57˝ E, June 21, 1989, coll. V.V.Anistratenko).
Рис. 1. Места сбора материала в Чёрном и Азовском морях: 1 — кордон «Морской», Черноморский
биосферный заповедник, Голопристанский р-н, Херсонская обл., Украина (46°07´54˝ N, 32°13´39˝ E,
2011 и 2012 гг., сбор В.В.Анистратенко, О. Ю. Анистратенко); 2 — Федотова коса возле пос.Кирил-
ловка, Акимовский р-н, Запорожская обл., Украина (46°19´40˝ N, 35°20´03˝ E, 2005, 2006, 2012 гг., сбор
В. В. Анистратенко, О. Ю. Анистратенко, И. А. Халиман); 3 — пос.Юркино, Ленинский р-н, АР Крым,
Украина (45°25´24˝ N, 36°33´15˝ E, 12 октября 2003 г., сбор В. В. Анистратенко, О. Ю. Анистратенко);
4 — с.Пичори, Галийский р-н, Абхазия (42°26´57˝ N, 41°32´30˝ E, 22 июня 1989 г., сбор В.В.Анистра-
тенко); 5 — пос. Чакви, Аджария (41°44´05˝ N, 41°43´57˝ E, 21 июня 1989 г., сбор В. В.Анистратенко).
459Conchological Variability of Anadara inaequivalvis...
zhzhya Region, Ukraine), about 50 other specimens came from the Sea of Azov coast of the Kerch Peninsula
(Yurkino settlement, Lenino District, Crimea, Ukraine). Additionally, some specimens from coastal area of
the Kerch Strait were studied, namely, from Tuzla Spit, Ak-Burun Cape and Opuk Cape, kindly provided by
Yu.V.Vernigorova and T. V. Shevchenko (Institute of Geological Sciences of NAS of Ukraine, Kyiv).
Shell shape variability of Anadara inaequivalvis was estimated based on a variety of an outline joining, i.e.
of a shell’s commissural opening. For this purpose we specially selected and studied 508 discrete valves (259le
and 249 right) from the BSBR and 400 valves (200 le and 200 right) from the Fedotova Spit (all collected in
2012). In specimens with two valves, only one of them was considered, thus, each valve considered has to be
considered as an individual specimen. Comparison of the shell shape was carried out as follows. On a transpar-
ent plastic plate an outline of commissural opening of the valve enclosed below was drawn by a marker. Each
of subsequent valves was compared to the received contour leveling them on hinge plate. If contours do not
coincide (excepting the sizes), a new contour from a di erent specimen was drawn. In case of an intermediate
form a valve was sorted to the closest of the basic types.
Beside the shell shape we analyzed variability of some other parameters: a number of radial ribs on the
surface of valves and their ornamentation, a quantity of chevrons on the ligament area, a shape of the hinge
plate and a number of hinge teeth.
Length of shells was determined on graph paper, their thickness was measured by caliper at the middle
part of the ventral edge. Lock studying and calculation of a number of ribs and chevrons was carried out with an
optical stereomicroscope MBS–9. Photos of shells are taken with “Canon Power Shot SX 120” camera.
Results and discussion
According to our observations Anadara inaequivalvis in the Black-Azov Sea Basin
possesses wide variability of a shell ( g. 2). Various degree of variability is characteristic for
its shell shape and thickness, shape of the hinge plate, number, shape and the relative size
of hinge tooth, quantity and surface texture of the radial ribs, prevailing color of the shell
and other features.
e most indicative, in our opinion, is the analysis of variability of a shell shape,
whereas other properties in a context of this research are considered to be minor and are
discussed in fewer details.
Usually the conchological variability is estimated with the aid of a verbal description
and/or quantitative parameters. However, in most cases the patterns and limits of shell vari-
ability can be exhibited with images much more clearly. In the research presented we used a
visually selected set of the main types of outline of Anadara shell valves joining (=locking)
which is called here “forms” ( g. 3). Numbering of these forms, i. e. certain types of a con-
tour of the shell commissural opening is selected arbitrarily and used only to substitute the
subjective verbal equivalents, such as “orbicular”, “ovate-trapezoid”, “rounded-tetragonal”,
etc. However, in the discussion of qualitative and quantitative characteristics of particular
forms we are compelled to use also similar statements (see below).
Within the studied samples of Anadara from the Black-Azov Sea Basin we de ne six
basic types of the shell commissural opening ( g. 3).  e form 4 exhibits the highest rate
of occurrence, whereas extreme forms occur rarely. Comparison of two large samplings of
the Black and the Azov seas (see Material and methods) shows that in both localities the
form 4 predominates, being the  rst dominant; the second dominants are forms 2 and 1
at the Azov Fedotova Spit and forms 2 and 3 at the BSBR. Occurrence frequency of other
forms in both samplings di ers considerably ( g. 4). We explain this as follows.  e domi-
nant form, i. e. the form 4 (non-equilateral trapezoid), represents a basic conchological
morphotype, generated here on the basis of initially introduced individuals whereas other
forms are its morphological (autochthonous) variants.  e relations between all de ned
forms can be expressed schematically ( g. 5) in two morphological ranges.  e rst range
4–2–1–3 re ects a gradual reduction of shell height and its relative lengthening; in the
form 3 it combines with rounding of anterior edge and elongation of posterior one.  e
second line 4–6–5 demonstrates a transition from the basic morphotype 4 to the form 6
by shortening of posterior edge (the contour of commissural opening comes nearer to an
equilateral trapezoid) and further to the form 5, an opening of which is almost roundish
caused by subsequent increasing of relative height of a shell. Herewith a hinge plate in form
5 is considerably shortened that makes it markedly arcuate.
460 V. V. Anistratenko, O. Yu. Anistratenko, I. A. Khaliman
It should be stressed that forms discussed here can not be considered as discrete states,
and no taxonomic status might be assigned to them. Although the intermediate variants make
an insigni cant part of the material studied, their existence evidences a smooth (continuous)
variability. Existence of transitional forms allows to consider Anadara successfully making
them at home in the Black and the Sea of Azov belonging to one species with wide morpho-
logical variability but not to several distinct “narrow” species. It corroborates with extremely
low probability of invading into the Mediterranean basin (one-time occasion or repeatedly in
to the Adriatic, the Black Sea and the Sea of Azov), 2–3 allied but di erent species of Anadara.
Finally, a dispersion of domination order of conchological morphotypes revealed in the com-
pared samplings of A. inaequivalvis means an independent existence of these populations and
only re ects particular characteristics of local conditions of their habitation.
Dimensional characteristics of A. inaequivalvis shell occurring in the Adriatic Sea and
in the Black-Azov Sea Basin di er signi cantly.  e length of this species shell in the Adri-
atic Sea reaches up to 70–75 mm, maximally even 80 mm (Rinaldi, 1985).  e mollusks liv-
ing in the Black Sea and in the Sea of Azov are much smaller (Anistratenko et al., 2011).  e
largest shell from our materials is of 65.0 mm length (table 1), but quantitatively dominate
Fig. 2. Shells of A. inaequivalvis from the Black Sea (G–L) and the Sea of Azov (А–F): А–F — specimens from
the northwestern part of the Sea of Azov (Fedotova Spit); G–L — specimens from the northwestern part of the
Black Sea (Cordon “Morskoj”, the BSBR). A, C, E, G, I, K — inner side of the valves; B, D, F, H, J, L — outer side
of the same specimens.
Рис. 2. Раковины A. inaequivalvis из Чёрного и Азовского морей: А–F — экземпляры с северо-западного
побережья Азовского моря (Федотова коса); G–L — экземпляры с северного побережья Чёрного моря
(кордон «Морской», ЧБЗ). A, C, E, G, I, K — внутренняя сторона створок; B, D, F, H, J, L — наружная
сторона тех же экземпляров.
A
G
C
I
E
K
B
H
D
J
F
L
461
Conchological Variability of Anadara inaequivalvis...
Fig. 3. Basic forms of the shell shape i. e. commissural opening in A. inaequivalvis from the Black Sea and
the Sea of Azov. A, B — form 1; C, D — form 2; E, F — form 3; G, H — form 4; I, J — form 5; K, L — form 6.
Allspecimens — from the BSBR. Numeration is selected arbitrarily, explanations see in the text.
Рис. 3. Основные формы комиссурального просвета раковины A. inaequivalvis из Чёрного и Азовского
морей на примере правой створки. A, B — форма 1; C, D — форма 2; E, F — форма 3; G, H — форма 4; I, J—
форма 5; K, L — форма 6. Все экземпляры из ЧБЗ. Нумерация форм произвольная, пояснения в тексте.
Fig. 4. A ratio of basic types of the shell shape in A. inaequivalvis from the Black Sea and the Sea of Azov. Enu-
merated forms correspond to those in the text discussed; the  rst symbol speci es an absolute number of valves
in the sample, the second symbol means a percentage share of this form in the sample.
Рис. 4. Соотношение основных вариантов изменчивости раковины A. inaequivalvis в пробах из Чёрного и
Азовского морей. Нумерованные формы соответствуют таковым, обсуждаемым в тексте; первая цифра обо-
значает абсолютное количество створок данной формы в пробе, вторая — доля этой формы в процентах.
A
G
1
4
2
5
3
6
C
I
E
K
B
H
D
J
F
L
462 V. V. Anistratenko, O. Yu. Anistratenko, I. A. Khaliman
valves reaching about 40 mm in length. Meanwhile, according to K. A. Lutaenko (2006)
maximum length of A. inaequivalvis shell in the southern India is 66 mm which is compa-
rable to that of the Black Sea Anadara.
Among the other features of Anadara shell, number of radial ribs on surface of valves,
number of chevrons on a ligament area, hinge plate shape, number and peculiarities of
hinge teeth location have essential value for the correct speci c identi cation (table 1).
Also, one of the determinative features of anadarins is valve-inequality; at least, in the  g-
ures of Anadara inaequivalvis group presented in works of J.Chemnitz and L.Reeve, this
character is expressed distinctly (Lutaenko, 2006: 113,  g. 5). It is signi cant that inequival-
vity in Anadara individuals from the Black and the Azov Seas is barely visible — a ventral
spacing between the valves of adult specimens does not exceed 2 mm ( g. 6, C–F).
e number of radial ribs on valve surface of the specimens studied varied from 30 to
36. Some distinctions of this characteristic were revealed in shells from di erent localities
(table 1). Upon the average the shells from the Black Sea bear 1–2 ribs more than those from
the Sea of Azov and possess somewhat wider limits of variation of this character ( g. 7).
As our material shows, the sculpture of ribs — their granularity in the middle and
the anterior parts of the shell is more developed on the le valve ( g. 6, A, B). However,
the right valves also have the expressed rugosity of ribs formed by thin transversal ribs.
Perfectly smooth look only ribs of the valves worn enough. On a photo of A.inaequivalvis
shells from the southern India provided by K. A. Lutaenko (2006: table 6, A, B, E, F) the
surface of ribs is absolutely lacking rugosity; the author of the article draws attention to this
fact also in the text (Lutaenko, 2006: 114).
Table 1. Morphological characters of the shells A. inaequivalvis from the Black Sea and the Sea of Azov
(dimensions are given in millimeters)
 1.    A. inaequivalvis    
 (    )
N
Le /Right
valve
(L/R)
Morpho-
type of
the shell
Length
oftheshell
mean
(min–max)
Number
ofradial ribs
mean
(min–max)
Number
ofchevrons
mean
(min–max)
ickness
oftheshell wall
mean
(min–max)
Number
ofteeth
mean
(min–max)
Cordon Morskoj, the Black Sea Biosphere Reserve, Gola Prystan District, Kherson Region, Ukraine
13 R 1–6 38.5 (20–53) 34.1 (32–36) 1.9 (1–3) 1.9 (1.0–2.6) 44.2 (39–50)
7 L 1, 3, 4, 5 37.6 (10–57) 33.4 (30–35) 1.9 (0–3) 1.9 (0.4–2.5) 43.3 (27–55)
Coast of the Caucasus near Pichory Village, the Black Sea (Gali district, Abkhazia)
6 R 2, 4, 6 35.0 (22–48) 34.6 (33–36) 1.5 (1–2) 1.6 (1.0–2.1) 46 (35–53)
7 L 2, 4, 6 34.7 (22–52) 34.7 (32–36) 1.6 (1–3) 1.7 (1.2–2.3) 45.9 (40–53)
Fedotova Spit near Kirillovka settlement, the Sea of Azov (Akimovka District, Zaporozh’e Region, Ukraine)
13 R 1–4, 6 34.1 (23–43) 32.2 (31–35) 1.4 (1–2) 1.5 (1.1–2.1) 40.5 (32–51)
18 L 1–4, 6 35.5 (26–45) 32.5 (31–34) 1.4 (1–2) 1.9 (1.3–2.9) 40.4 (32–48)
Coast of the Kerch Peninsula, the Sea of Azov (Yurkino settlement, Lenino District, Crimea, Ukraine)
1 L 2 65 34 4 3.1 56
Kerch Strait, Tuzla Spit (Crimea, Ukraine)
1 R 2 48 34 2 1.6 46
2 L 2, 4 54.5 (53–56) 33.5 (33–34) 2 2.25 (2.1–2.4) 48 (46–50)
Kerch Strait, Ak–Burun Cape (Crimea, Ukraine)
2 R 2, 3 39.5 (38–41) 34 2 1.5 (1.4–1.6) 44 (38–50)
2 L 4 49.5 (49–50) 34 (33–35) 2 (1–3) 2.45 (2.3–2.6) 48.5 (45–52)
Kerch Peninsula, Opuk Cape, the Black Sea (Crimea, Ukraine)
5 R 1–4 52.4 (47–56) 35 (33–37) 2.4 (1–3) 2.3 (2.0–2.7) 49.4 (43–56)
7 L 1, 2, 4 50 (33–61) 33.6 (33–35) 2.3 (1–3) 2.3 (1.3–3.0) 50.1 (46–54)
463Conchological Variability of Anadara inaequivalvis...
ere are 1 to 4 chevrons on a ligament area of Anadara from the Black-Azov Sea Ba-
sin; usually two of them are available.  e number of chevrons partly depends on the shell
size though this dependence apparently is not a direct one.  us, a specimen of 65 mm
length has four chevrons while the majority of valves of di erent size classes have either 1
or 2 chevrons (table 1). No any correlation between the number of chevrons and the mor-
photype of the commissural opening was registered.
Hinge plate of Anadara studied is almost straight, only specimens with relatively short-
er shell (forms 5 and6) possessed the plate in a shape of a slightly curved arch (see above).
e number of hinge teeth of equally-sized shells (i. e. having the similar age) is not con-
Fig. 5. Morphological ranges of basic types of the shell shape in A. inaequivalvis from the Black-Azov Sea Basin.
Outlines of the shell commissural opening are given in the same scale.  e form 4 is de ned as initial for all
other modi cations presented.
Рис. 5. Морфологические ряды основных форм A. inaequivalvis из Азово-Черноморского бассейна. Кон-
туры комиссурального просвета раковины приведены к одному масштабу. Форма 4 определена в каче-
стве исходной для всех остальных вариантов изменчивости.
Fig. 6. Morphology of the shell A. inaequivalvis from the Black-Azov Sea Basin: A, B — le valve with well-
sculptured radial ribs (Caucasus coast near Pichory village, Abkhazia); C–F — inequivalvity in adult shells:
C,D — specimen from the Sea of Azov, Fedotova Spit, E, F — specimen from the Black Sea, cape Opuk. Arrows
indicate the line of valves occlusion.
Рис. 6. Морфология раковины A. inaequivalvis из Азово-Черноморского бассейна: A, B — левая створка
с хорошо выраженной скульптурой на поверхности радиальных рёбер (побережье Кавказа у с. Пичори,
Абхазия); C–F — неравностворчатость взрослых раковин: C, D — экземпляр из Азовского моря, Федо-
това коса, E, F — экземпляр из Чёрного моря, м. Опук. Стрелки показывают линию смыкания створок.
ACE
BDF
464 V. V. Anistratenko, O. Yu. Anistratenko, I. A. Khaliman
stant; the increase of the shell size usually led to the increase of the number of teeth.  us,
the shell 10 mm long was supplied with 27 teeth, lock of the shell 35–40 mm long bore from
37 to 49 teeth, and in specimens with length over 45 mm we registered from 44 to 56teeth
(table 1). On hinges of the Sea of Azov specimens (and more rarely those of the Black Sea)
some anomalies of teeth were revealed: on posterior edge, and sometimes on anterior edge
of the teeth the series up to 2–3 teeth merged in one tooth ( g. 2, G); also teeth of irregular
shape as calluses, instead of normal plates were sometimes observed.
In addition, judging by the illustrations presented by K.A. Lutaenko (2002: 38,  g. 5),
variability of the shell shape of A. inaequivalvis from the Holocene deposits of Peter the
Great Bay (northwestern Sea of Japan) demonstrated similar patterns with those revealed
in the Black-Azov Sea Basin (data presented here).
Comparison of morphological types of Anadara shells from the Black Sea and the Sea
of Azov on the one side, and A. inaequivalvis inhabiting the southeast India on the other
side, shows the following : the most “popular” morphotype among the Anadara from the
Black–Azov Sea Basin (form 4) demonstrates a strong resemblance to A.inaequivalvis of
Coromandel Coast of India. All images of this species shells from the type locality given by
K.A.Lutaenko (2006, phototable 3, E–I; phototable 6, A–H,)  t well to variability limits of
Anadara from the Black Sea and the Sea of Azov. Shells of A. inaequivalvis from Romania
(Lutaenko, 2006, phototable 11, E–H) by a contour of commissural opening also correspond
well to the form 4; and shells from the Adriatic Sea (Lutaenko, 2006, phototable 11, A–D)
correspond to the form 1. Concerning the number of a shell radial ribs A. inaequivalvis
from southern India, bearing from 31 to 36 and usually 33–35 (Lutaenko, 2006), does not
exceed variability limits of this character in mollusks from the Black and Azov seas (table1;
g. 7).
ereby, a conspeci ty of at least one of alien Anadara nominal species known in the
Mediterranean Basin (Morello, Solustri, 2001; Morello et al., 2004 for records of Anadara
demiri in the Adriatic Sea) to A. inaequivalvis can be considered as properly grounded. To
verify this decision more data on variability of this species within its native area are to be
obtained as well as the molecular and genetic analyses are to be completed.
Conclusion
e present study shows a wide morphological variability of A. inaequivalvis shell
characters in the Black-Azov Sea Basin, i. e. far beyond of this species native area.  ereby,
the invader species realizes its adaptive abilities and apparently grow roots here in the newly
occupied region of the World Ocean.
Fig. 7.  e number of radial ribs in A. inaequivalvis from the Black Sea and the Sea of Azov.
Рис. 7. Количество радиальных рёбер A. inaequivalvis из Чёрного и Азовского морей.
465Conchological Variability of Anadara inaequivalvis...
e existence of intermediate forms proves a smooth (continuous) variability of the
shell and belonging of all the studied populations to the same species. Revealed distinctions
in a quantitative ratio of morphotypes in compared samples of A. inaequivalvis mean
independent existence of these populations and re ect particularities of local living
conditions.
From the taxonomic point of view the data obtained probably allow to identify
Anadara inhabiting the Black-Azov Sea Basin as A. inaequivalvis, since they correspond
to A. inaequivalvis from the type locality. For more tenable decision the molecular and
genetic analyses of as more as possible number of Anadarinae species should have a
crucial importance.
e authors are grateful to sta of the Black Sea Biosphere Reserve of the National Academy of Sciences
of Ukraine for their hospitality and kind assistance during  eld works in 2011 and 2012. Also we thank
Yu.V. Vernigorova and T.V. Shevchenko (Institute of Geological Sciences of NAS of Ukraine, Kyiv) for the
opportunity to examine their samples from the area of the Kerch Strait. We are much obliged to two anonymous
reviewers for their constructive criticism, helpful comments and suggestions.
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Received 3 March 2014
Accepted 24 September 2014
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... Ширина ареи под макушкой (у азовоморских форм) -0,15-0,20 от выпуклости (Чихачев и др., 1994). Число шевронов на арее у черноморских форм -0-4 (Anistratenko et al., 2014), у форм из Адриатического моря -1-3 (Лутаенко, 2006). Макушки створок выступающие, слегка смещены к переднему краю. ...
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Lessepsian invader Anadara kagoshimensis is one of the important elements in the adaptive transformation of the Black sea shelf ecosystem in the second half of the XX century. The aim of the work is to analyse the Black Sea spatial and temporal colonization by this species according to the benthic database of Benthos Ecology Department of IMBR RAS (for the Crimean coasts) and literary sources for the whole Black Sea basin. The phases of this process are considered. A 10-year latency period followed after the first detection anadara in 1968 at Caucasian shelf. The main wave of anadara colonization in the Black Sea covered the Western and Eastern parts of the sea and fall within the period from second half of 1970s to 1980s. During that time period the extended development of A. kagoshimensis lead to forming own reproductive potential for next steps of expansion of this species to Anatolian and Crimean coasts in 1990s. About 20–25 years delay in detection of introduced species at the Northern and Southern parts of the Black Sea shelf, in presence at the Western and Eastern parts, could be an evidence of existing hydrological barrier between far distant nearshore areas of the Black Sea shelf. The reasons of the fluctuations in development of this species settlements are discussed. The phenomenon of introduction and mass development of A. kagoshimensis in the Black Sea during its the “most demand” period as a consumer of excess amount of organic matter, in conditions of eutrophication’s peak in the Black Sea basin at the end of 1980s — the beginning of 1990s is marked. Estuarine areas of the Black Sea shelf enriched by organic matter are the most favourable conditions for the mass development of this species. This quantitative rising is due to wide ecological plasticity and stress tolerance of A. kagoshimensis, which are determined by its physiological and biochemical adaptations to a hypoxic conditions often appearing during eutrophication. Based on parameters of abundance and biomass the actual implementation of A. kagoshimensis biotic potential in the Black Sea is revealed at least within Eastern and Western areas of the shelf, where peaks of its quantitative development are already passed in previous years. This conclusion for the Crimean sector is not obvious because of more recent invasion by anadara of this region. A decrease and stabilization of A. kagoshimensis development, with the exception of local zones near organic ”pollution” sources should be expected under maintaining the tendency of the basin deeutrophication.
... These marine communities are heavily affected by three invasive mollusk species, especially in the NW Black Sea: Mya arenaria, Rapana venosa, and Anadara sp. (see for taxonomy discussion of the latter Anistratenko et al., 2014;Anistratenko & Khaliman, 2006;Krapal et al., 2015). In areas with strong freshening, such as the Razim-Sinoe system, freshwater mollusk species, including non-native bivalves (i.e., S. woodiana, C. fluminea) and viviparids, expanded at the cost of PC species Velde et al., 2019). ...
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The unique aquatic Pontocaspian (PC) biota of the Black Sea Basin (BSB) is in decline. The lack of detailed knowledge on the status and trends of species, populations, and communities hampers a thorough risk assessment and precludes effective conservation. This paper reviews PC biodiversity trends in the BSB (Bulgaria, Romania, Moldova, Ukraine, and Russia) using endemic mollusks as a model group. We aim to assess changes in PC habitats, community structure, and species distribution over the past century and to identify direct anthropogenic threats. The presence/absence data of target mollusk species were assembled from literature, reports, and personal observations. Pontocaspian biodiversity trends in the northwestern BSB coastal regions were established by comparing 20th- and 21st- century occurrences. The direct drivers of habitat and biodiversity change were identified and documented. We found that a pronounced decline of PC species and communities is driven by (a) damming of rivers, (b) habitat modifications that disturbed previous natural sa-linity gradients and settings in the studied area, (c) pollution and eutrophication, (d) invasive alien species, and (e) climate change. Four out of the 10 studied regions, namely, the Danube Delta– Razim Lake system, Dniester Liman, Dnieper– Bug estuary, and Taganrog Bay– Don Delta, contain favorable ecological conditions for PC com-munities and still host threatened endemic PC mollusk species. Distribution data are incomplete, but the scale of deterioration of PC species and communities is evident from the assembled data, as are major direct threats. Pontocaspian biodiversity in the BSB is profoundly affected by human activities. Standardized observation and collection data as well as precise definition of PC biota and habitats are necessary for targeted conservation actions. This study will help to set the research and policy agenda required to improve data collection to accommodate effective conservation of the unique PC biota.
... These marine communities are heavily affected by three invasive mollusk species, especially in the NW Black Sea: Mya arenaria, Rapana venosa, and Anadara sp. (see for taxonomy discussion of the latter Anistratenko et al., 2014;Anistratenko & Khaliman, 2006;Krapal et al., 2015). In areas with strong freshening, such as the Razim-Sinoe system, freshwater mollusk species, including non-native bivalves (i.e., S. woodiana, C. fluminea) and viviparids, expanded at the cost of PC species Velde et al., 2019). ...
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Aim The unique aquatic Pontocaspian (PC) biota of the Black Sea Basin (BSB) is in decline. Lack of detailed knowledge on the status and trends of species, populations and communities hampers a thorough risk assessment and precludes effective conservation. This paper aims to review PC biodiversity trends using endemic molluscs as a model group. We aim to assess changes in PC habitats, community structure and species distribution over the past century and to identify direct anthropogenic threats. Location Black Sea Basin (Bulgaria, Romania, Moldova, Ukraine and Russia). Methods Presence/absence data of target mollusc species was assembled from literature, reports and personal observations. PC biodiversity trends in the NW BSB coastal regions were established by comparing 20th and 21st century occurrences. Direct drivers of habitat and biodiversity change were identified and documented. Results A very strong decline of PC species and communities during the past century is driven by a) damming of rivers, b) habitat modifications negatively affecting salinity gradients, c) pollution and eutrophication, d) invasive alien species and e) climate change. Four out of 10 studied regions, namely, the Danube Delta – Razim Lake system, Dniester Liman, Dnieper-South Bug Estuary and Taganrog Bay-Don Delta contain the entire spectrum of ecological conditions to support PC communities and still host threatened endemic PC mollusc species. Distribution data is incomplete, but the scale of deterioration of PC species and communities is evident from the assembled data, as are major direct threats. Main conclusions PC biodiversity in the BSB is profoundly affected by human activities. Standardised observation and collection data as well as precise definition of PC biota and habitats are necessary for targeted conservation actions. This study will help to set the research and policy agenda required to improve data collection to accommodate effective conservation of the unique PC biota.
... A complex of bivalve filter-feeder mollusks is an important component of the Black Sea ecosystems (Zenkevich, 1986). Their potential competitor is a representative of genus Anadara Gray, 1847, which almost simultaneously invaded the Mediterranean and Black seas about 50 years ago and which identification to species was a subject of debate (Lutaenko, 2006(Lutaenko, , 2015Revkov, 2016;Anistratenko et al., 2014). Anadara that colonized the Black Sea was confirmed by modern genetic studies using barcoding to exhibit 99.8-100% similarity to Anadara kagoshimensis (Tokunaga, 1906) from Japan (Krapal et al., 2014). ...
Article
Bivalve mollusk Anadara kagoshimensis is classified among the top 100 dangerous invasive species of the Mediterranean and Black seas. Using as an example the colonization by Anadara of the Kazachya (Cossack) Bay (Crimea, the Black Sea), the article identifies and discusses biocenotic relations of the mollusk with sea bottom substrates, epibionts, potential environmental competitors, and predator Rapana venosa. Relations of A. kagoshimensis to the complex of phytoepibionts and zooepibionts are presented for the first time. It is shown that Anadara can be an edificator and form a specific complex of algoconsorts and zoocon-sorts, while playing a positive role in increasing and preserving the species diversity in certain areas of the Black Sea. The increase in biomass and diversity of filter feeders following the occurrence of Anadara in the study region argues in favor of its positive effect on the biocenosis.
... This observation demonstrates the ongoing of pontization process in the Sea of Azov, that is, the enrichment of the sea fauna by certain Black Sea species, and gives a reason to assume that the number of Black Sea species found in the Sea of Azov will increase in the future . Moreover, the invasion of species from distant sea basins into the Sea of Azov is continuing, and therefore, regular monitoring of these species is also necessary Anistratenko et al., 2014). ...
Article
Summarized data on the fauna composition, distribution, and ecology of gastropod and bivalve mollusks of the Utlyuk Liman in the northwestern part of the Sea of Azov is presented. The total number of mollusk species identified was 63; 43 species belonged to the class Gastropoda, and 20 species, to the class Bivalvia. The distribution of mollusks in the liman has extremely irregular character, whereas the distribution of species along the marine shore of Biryuchii Ostrov spit is more homogeneous. Euryhaline Mediterranean species represent the core of liman malacofauna; some taxa of the Ponto-Caspian zoogeographical complex (Dreissena polymorpha and species of the genus Theodoxus) and invader species from distant sea basins (Mya arenaria and Anadara inaequivalvis) were also identified.
... A. kagoshimensis is a successful invasive species to the Mediterranean, Black, and Azov seas but was misidentified for a long time as Anadara (Scapharca) cornea (reeve 1844) (ghisotti 1973;zoLotarev & zoLotarev 1987) or Anadara (Scapharca) inaequivalvis (Bruguière 1789) (ghisotti & rinaLDi 1976;rinaLDi 1978rinaLDi , 1994PoPPe & goto 1993;hrs-BrenKo & LegaC 1996;zenetos & al. 2003;anistratenKo & al. 2014). Although I clearly noted differences between the Indian (type locality) and Mediterranean basin anadarines belonging to this complex (LutaenKo 2006), it was huBer (2010) who suggested for the first time that the Mediterranean species is instead A. kagoshimensis. ...
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Carl Emil Lischke (1813–1886) contributed much to Japanese marine malacology and published the celebrated “Japanische Meeres-Conchylien”, in three volumes (1869–1874). Part of his collection, including samples brought from Japan by the Russian Navy officer N.A. Birilev, is stored in the Zoological Institute, Russian Academy of Sciences (ZIN, St. Petersburg). In total, Lischke enumerated ten species of Arcidae from Japan, and we have located seven of Lischke’s original arcid species in the ZIN collection. In modern nomenclature there are six species: Arca avellana Lamarck 1819, Acar plicata (Dillwyn 1817), Barbatia (Savignyarca) virescens (Reeve 1844), Barbatia (Abarbatia) decussata (G.B. Sowerby I 1833), Anadara (Scapharca) kagoshimensis (Tokunaga 1906), and Anadara (Tegillarca) granosa (L. 1758). Also, among them are syntypes of Arca subcrenata Lischke 1869 (= A. kagoshimensis) illustrated here for the first time. All species are reviewed with regard to their present-day taxonomic position, and illustrated. The history of the C.E. Lischke and N.A. Birilev collections is briefly described.
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The hypothesis on non-random correlation between abnormalities in the structure of hinge plate and infection of mollusks Macoma balthica with trematodes of the family Gymnophallidae has been tested on the basis of material from the Barents Sea. Significant correlation between the presence of warts and infection was established upon intraand interpopulation comparison. The hypothesis states that parasitizing of trematodes in the extrapallial cavity of mollusks influences the mantle functioning and provokes abnormalities in the hinge plate structure.
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The Black Sea is in transition from a freshwater to a marine environment and has a very low biodiversity. Therefore, it appears to be particularly vulnerable and easy target for many exotic species. Anthropogenic introductions of exotic species into the Black Sea began in the 19th century, but accelerated during the second half of the century. There are about 36 kinds of introduced species in Black and Azov seas now and some of them had already severe damage to part or whole ecosystem, while some may have limited or neutral impacts. Current status of two exotic benthic species presented here: Rapa Whelk, Rapana thomasiana and blood-cockle, Anadara inaeguivalvis. Most dramatic changes to benthic ecosystem of the Black Sea has taken place after introduction of predatory gastropod Rapa whelk from Far East (Sea of Japan) to the Black Sea in 1940s and has since spread to the Aegean and Adriatic Seas. The second species blood-cockle is a filtering feeding bivalve introduced 20 years ago, but not so well-known since its invasion ability and impact on ecosystem seems to be not very devastating. Particularly rapid distribution and increased biomass of Rapa Whelk caused severe damage to narrow benthic ecosystem. It has direct (predation on bivalves) and indirect (fishing with dredges) negative impacts on the ecosystem. Annual Rapa. whelk catches from Turkey and Bulgaria reached totally 13,000 ton year-1. This study presents updated information on spatial distribution, habitats preferences, population structures, ecological and economical impacts of two exotic species; Rapa whelk and blood-cockle.
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The bivalve mollusk Anadara inaequivalvis (Bruguie`re, 1789) is found for the first time in the Northwest part of the Sea of Azov. The find registers a final stage of settling by this species in the Azov-Black Sea basin. Distribution, ecological peculiarities of this mollusk and significance of its penetration for the local fauna are discussed. Key words: Bivalvia, Arcidae, Anadara, malacofauna, the Black Sea, the Sea of Azov.
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Anistratenko V.V., Khaliman I.A & Anistratenko O.Yu. 2011. The Molluscs of the Sea of Azov. Kiev: Naukova dumka. 173 pp. [in Russian with English Abstract]. The present book is the monographic description of molluscs inhabiting the Sea of Azov, adjacent lagoons (Molochnyj and Utlyuk) and Taganrog Bay. In foreword chapter the general characteristics of the Sea of Azov and short essay on the history of its molluscs study are given. General part consists of Material and Methods of the present study description, morphologic sketch of Phylum Mollusca as well as Gastropod and Bivalve classes; main characteristics of soft body and shell used for species identification are also provided here. Systematic part includes the illustrative description of all species registered within the region studied: 70 species of Gastropoda and 26 species of Bivalvia (except for freshwater molluscs). The final chapter considers some problems of the Azov Sea molluscs’ geographic distribution and ecology depending on water salinity. At the end of the monograph the reference list and alphabetic index for scientific names of taxa are also provided. The publication is intended for professionals in the field of invertebrate zoology, hydrobiology, it is also addressed to students in biology and to wide audience of nature lovers.
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This paper reports the observations recorded on the arcid clam, Anadara demiri, from its first reporting in the Adriatic Sea three years ago, when its presence only was noted, to the present day when its invasive potential is confirmed. Repeated recruitment events were observed and confirmed by the co-existence of two or more cohorts within the population. Anadara demiri appears to be a year-round spawner with two spawning peaks (late winter and summer). Together with the congeneric invader, Anadara inaequivalvis, it has colonized the external belt of the fine well-sorted sand biocoenosis where they may numerically dominate the benthic community. Anadara demiri appears to have gained full status as a component of the Adriatic Sea coastal benthic community despite its very recent first entry. Numerous features make it a strong competitor among which is the ability, retained into adult life, of attaching, by means of byssus threads to all kinds of hard substrata, particularly to other live molluscs such as the gastropod Aporrhais pespelecani and the bivalves Anadara inaequivalvis and Chamelea gallina. The presence of species actively interfering with A. demiri in various ways (the polychaetes, Polydora sp. and Sabellaria spinulosa, that bore into and foul the shell, respectively, and the predatory asteroid Astropecten irregularis) is discussed.
First record of Anadara demiri (Piani, 1981) (Bivalvia: Arcidae) in Italian waters // Bollettino Malacologico
  • E Morello
  • C Solustri
Morello, E., Solustri, C. First record of Anadara demiri (Piani, 1981) (Bivalvia: Arcidae) in Italian waters // Bollettino Malacologico. -2001. -37, N 9-12. -P. 231-234.
Population characteristics of settlements Anadara inaequivalvis (Bivalvia, Arcidae) of Odessa Black Sea region // Ecological security of coastal and shelf zones and integrated utilization of a shelf resources. Collection of scientifi c works
  • N L Finogenova
Finogenova, N. L. Population characteristics of settlements Anadara inaequivalvis (Bivalvia, Arcidae) of Odessa Black Sea region // Ecological security of coastal and shelf zones and integrated utilization of a shelf resources. Collection of scientifi c works. -2011. -25. -P. 392-399. -Russian : Финогенова Н. Л. Популяционные характеристики поселений Anadara inaequivalvis (Bivalvia, Arcidae) Одесского региона Черного моря.
Osservazioni relative a molluschi appartenenti al genere Anadara viventi in Adriatico // Conchiglie. -1972
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Rinaldi, E. Osservazioni relative a molluschi appartenenti al genere Anadara viventi in Adriatico // Conchiglie. -1972. -8. -P. 121-124.
Черноморский биосферный заповедник Анистратенко); 2 — Федотова коса возле пос. Кирилловка , Акимовский р-н, Запорожская обл); 3 — пос
  • В В Сбор
  • О Анистратенко
Рис. 1. Места сбора материала в Чёрном и Азовском морях: 1 — кордон «Морской», Черноморский биосферный заповедник, Голопристанский р-н, Херсонская обл., Украина (46°07´54˝ N, 32°13´39˝ E, 2011 и 2012 гг., сбор В. В. Анистратенко, О. Ю. Анистратенко); 2 — Федотова коса возле пос. Кирилловка, Акимовский р-н, Запорожская обл., Украина (46°19´40˝ N, 35°20´03˝ E, 2005, 2006, 2012 гг., сбор В. В. Анистратенко, О. Ю. Анистратенко, И. А. Халиман); 3 — пос. Юркино, Ленинский р-н, АР Крым, Украина (45°25´24˝ N, 36°33´15˝ E, 12 октября 2003 г., сбор В. В. Анистратенко, О. Ю. Анистратенко);