Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea
The paper presents the results of a study of the size-at-age and sex variability of plastic characters in the late ontogenesis of Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) from the northern part of the Black Sea. It is established, that males and females can be differentiated into small...
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nasplib_isofts_kiev_ua-123456789-1613212025-02-10T00:02:39Z Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea Размерно-возрастная и половая изменчивость пластических признаков зеленушки глазчатой, Symphodus Ocellatus (Linnaeus, 1758) (Labridae, Perciformes) в северной части Черного моря Розмірно-вікова і статева мінливість пластичних ознак зеленушки плямистої, Symphоdus ocellatus (Linnaeus, 1758)(Labridae, Perciformes), у північній частині Чорного моря Peskov, V.N. Manilo, L.G. The paper presents the results of a study of the size-at-age and sex variability of plastic characters in the late ontogenesis of Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) from the northern part of the Black Sea. It is established, that males and females can be differentiated into small and large individuals based on absolute values of 26 plastic characters. Differences between these groups are highly reliable in all characters. According to the relative values, the differences are significant for 14 characters in males , and 6 characters in females. The expression of sex differences in S. ocellatus increases with age both in absolute and relative values of plastic characters. Among large and small individuals of the Ocellated wrasse, males are significantly larger than females. The structure of the expression and the directionality of sex differences in the Ocellated wrasse also are changed. Приведены результаты изучения размерно-возрастной и половой изменчивости пластических признаков в позднем онтогенезе зеленушки глазчатой, Symphodus ocellatus (Linnaeus 1758) в северной части Черного моря. Установлено, что самцы и самки по абсолютным значениям 26 пластических признаков дифференцируются на мелких и крупных особей, различия между которыми высоко достоверны по всем признакам. По относительным значениям признаков различия достоверны у самцов по 14 признакам, у самок — по 6. Показано, что с возрастом у S. ocellatus увеличивается выраженность половых различий как по абсолютным, так и по относительным значениям пластических признаков. Среди крупных и мелких особей зеленушки самцы достоверно крупнее самок. С возрастом у зеленушки глазчатой изменяются структура выраженности и направленности половых различий. Наведено результати вивчення розмірно-вікової і статевої мінливості пластичних ознак у пізньому онтогенезі зеленушки плямистої, Symphodus ocellatus (Linnaeus, 1758) в північній частині Чорного моря. Встановлено, що самці і самки за абсолютними значеннями 26 пластичних ознак диференціюються на дрібних і великих особин, відмінності між якими високо достовірні за всіма ознаками. За відносними значеннями ознак відмінності достовірні у самців за 14 ознаками, у самок — за 6. Показано, що з віком у S. ocellatus збільшується вираженість статевих відмінностей як за абсолютними, так і за відносними значеннями пластичних ознак. Серед великих і дрібних особин зеленушки самці достовірно більші за самок. З віком у зеленушки плямистої змінюються структура вираженості і спрямованості статевих відмінностей. 2018 Article Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea / V.N. Peskov, L.G. Manilo // Збірник праць Зоологічного музею. — 2018. — Вип. 49 — С. 3-15. — Бібліогр.: 30 назв. — англ. 0132-1102 https://nasplib.isofts.kiev.ua/handle/123456789/161321 597.556.331.5(477) en Збірник праць Зоологічного музею application/pdf Національний Науково-природничий музей НАН України |
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| description |
The paper presents the results of a study of the size-at-age and sex variability of plastic characters in the late ontogenesis of Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) from the northern part of the Black Sea. It is established, that males and females can be differentiated into small and large individuals based on absolute values of 26 plastic characters. Differences between these groups are highly reliable in all characters. According to the relative values, the differences are significant for 14 characters in males , and 6 characters in females. The expression of sex differences in S. ocellatus increases with age both in absolute and relative values of plastic characters. Among large and small individuals of the Ocellated wrasse, males are significantly larger than females. The structure of the expression and the directionality of sex differences in the Ocellated wrasse also are changed. |
| format |
Article |
| author |
Peskov, V.N. Manilo, L.G. |
| spellingShingle |
Peskov, V.N. Manilo, L.G. Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea Збірник праць Зоологічного музею |
| author_facet |
Peskov, V.N. Manilo, L.G. |
| author_sort |
Peskov, V.N. |
| title |
Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea |
| title_short |
Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea |
| title_full |
Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea |
| title_fullStr |
Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea |
| title_full_unstemmed |
Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea |
| title_sort |
size-at-age and sex variability of plastic values in ocellated wrasse, symphodus ocellatus (linnaeus, 1758) (labridae, perciformes), in northern part of the black sea |
| publisher |
Національний Науково-природничий музей НАН України |
| publishDate |
2018 |
| url |
https://nasplib.isofts.kiev.ua/handle/123456789/161321 |
| citation_txt |
Size-at-age and sex variability of plastic values in Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758) (Labridae, Perciformes), in northern part of the Black Sea / V.N. Peskov, L.G. Manilo // Збірник праць Зоологічного музею. — 2018. — Вип. 49 — С. 3-15. — Бібліогр.: 30 назв. — англ. |
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Збірник праць Зоологічного музею |
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2025-12-01T23:51:56Z |
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Zb. prac’ Zool. muz. (Kiïv), 49, 2018 3
Size-at-age and sex variability of plastic values in Ocellated wrasse ...
Introduction
Body size in fish is determined by their age, growth rate and puberty. It
is known that fish have a strong positive correlation between age and
body size (Cherepanov, 1986), so researchers often study the size or size
-at-age variation of plastic characters instead of age (Fauna…, 1982;
Schevchenko, 1971, etc.). Sex differences in the linear dimensions and
proportions of the body are characteristic for many fish species. Unlike
sexual dimorphism, when males and females differ in qualitative dis-
crete characters, which allow to consider them as two independent
morphs, sex differences in quantitative characters with continuous
variability are characterized by such indicators as expression, direction-
ality and structure.
© V. N. Peskov, L. G. Manilo, 2018
The paper presents the results of a study of the size-at-age and sex variability of plastic char-
acters in the late ontogenesis of Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758)
from the northern part of the Black Sea. It is established, that males and females can be dif-
ferentiated into small and large individuals based on absolute values of 26 plastic characters.
Differences between these groups are highly reliable in all characters. According to the rela-
tive values, the differences are significant for 14 characters in males , and 6 characters in
females. The expression of sex differences in S. ocellatus increases with age both in abso-
lute and relative values of plastic characters. Among large and small individuals of the Ocel-
lated wrasse, males are significantly larger than females. The structure of the expression and
the directionality of sex differences in the Ocellated wrasse also are changed.
K e y w o r d s: plastic characters, size-at-age and sexual variability, Symphodus ocellatus,
Black Sea.
UDC 597.556.331.5(477)
V. N. Peskov, L. G. Manilo
National Museum of Natural History of the NAS of Ukraine
B. Khmelnitsky 15, Kyiv, 01030, Ukraine
E-mail: vladimir.peskov53@gmail.com
leonid.manilo@gmail.com
SIZE-AT-AGE AND SEX VARIABILITY OF
PLASTIC VALUES IN OCELLATED WRASSE,
SYMPHODUS OCELLATUS (LINNAEUS, 1758)
(LABRIDAE, PERCIFORMES),
IN NORTHERN PART OF THE BLACK SEA
Збірник праць Зоологічного музею, 49: 3–15, 2018
4 Зб. пр. Зоол. муз., 49, 2018
V. N. Peskov, L. G. Manilo,
The expression or magnitude and significance of differences between males and
females are usually assessed by average values of characters (for example, using Stu-
dent’s t-test), by integral indicators (main components, factors, canonical variables,
etc.), by difference metrics (Euclidean distance, Mahalanobis metric) or by the value of
specially calculated indicators (Lovich, Gibbons, 1992; Peskov et al., 2017).
The directionality of sex differences is determined by the sign with a corresponding
difference indicator. For example, if males are larger than females (L♂ > L♀), when
comparing them along the body length “L♂ — L♀”, the value of the difference indicator
will have a “+” sign, if vice versa (L♂ <L♀) the sign “–”. In the majority of clupeid and
carp fishes, females are larger than males. Salmon males, as many other species protect-
ing the eggs, are larger than females (Svetovidov, 1964).
The structure of sex differences is determined by the fact that the magnitude and
expression of differences between males and females differ in various characters, there-
fore, all compared characters can be structured (grouped, ranked) according to these
criteria.
Thus, the aim of this paper is to estimate the size-at-age and sex variability of plastic
characters of Ocellated wrasse, Symphodus ocellatus (Linnaeus, 1758), in late ontogene-
sis.
Material and methods
We used materials sampled in the coastal waters of Ukraine in the Black Sea. The
material is housed in the ichthyological collection of the National Museum of Natural
History, National Academy of Sciences of Ukraine (Movchan et al., 2003) (Table 1).
Table 1. Brief characteristics of the processed material
No. Sampling site Sampling date
Sex, number
of specimens
1139 Crimea, Chernomorskyi Distr., Rybachie vill. 25.05.1958 1 ♂
1161
Kherson Reg., Golopristanskyi Distr., cordon
“Yagorlytsky Kut” of the Black Sea Bio-
sphere Reserve, Yagorlytskyi Bay
6–10.07.1980 2 ♂, 3 ♀
1171 Crimea, Sudakskyi Distr., Karadag Reserve 1–23.06.1980 1 ♂, 1 ♀
1182
Nikolaev Reg., Ochakovskyi Distr.,
Pokrovka vill., Yagorlytskyi Bay
1–5.07.1980 6 ♂, 2 ♀
1188
Odessa Reg., Tatarbunarskyi Distr., Sasyk
Lake
18.08–7.09.1967 4 ♂, 5 ♀
3516 Crimea, Sudakskyi Distr., Karadag Reserve 14–31.05.1979 2 ♂, 1 ♀
5564 Crimea, Sudakskyi Distr., Karadag Reserve 24.06–8.07.1980 8 ♂, 1 ♀
5570 Kherson Reg., Golopristanskyi Distr., Tendra 20.07.1988 1 ♂, 16 ♀
5635,
5636
Crimea, near Sevastopol, Kazachia Bay 5–27.06.1974 5 ♂, 6 ♀
5851 Crimea, Sudakskyi Distr., Karadag Reserve 31.05.1979 1 ♂, 1 ♀
7649 Crimea, Sudakskyi Distr., Karadag Reserve 5.08.2002 5 ♂
7920 Crimea, near Yalta, “Cape Martian” Reserve 9.09.2010 1 ♂
8595
Crimea, Chernomorskyi Distr.,
near Olenevka vill., Bolshoi Kostel Bay
23–27.06.2007 3 ♂, 2 ♀
9546
Crimea, Leninskyi Distr.,
Kerch Peninsula, Opuk cape
29–31.05.2013 2 ♂
Zb. prac’ Zool. muz. (Kiïv), 49, 2018 5
Size-at-age and sex variability of plastic values in Ocellated wrasse ...
Fish were caught by gill nets at depth of 10–15 m, and fixed in 4 % formaldehyde
solution for subsequent processing in laboratory.
Measurements of plastic characters are made with an electronic caliper, with 0.1 mm
precision: TL — total body length; SL — standard body length (from the beginning of
the upper lip to the beginning of base of caudal fin rays); H — maximum body height
before ventral fins; h — height of the caudal peduncle; aD1 — the distance from the
beginning of the upper lip to the first spiny ray of the dorsal fin; aD2 — the distance
from the beginning of the upper lip to the first soft ray of the dorsal fin; aP — antepecto-
ral length (from the upper lip to beginning of the pectoral fin base); aV — anteventral
length (from the upper lip to beginning of the ventral fin base), aA — anteanal length
(from the upper lip to beginning of the anal fin); pD — postdorsal length (from the end
of dorsal fin base to the beginning of the base of the middle rays of caudal fin); lcaud —
the length of caudal peduncle (from the end of anal fin base before the base of the mid-
dle rays of caudal fin); lD — length of the dorsal fin base; lD1 — length of the base of
the spiny part of the dorsal fin; lD2 — length of the base of the soft part of the dorsal fin;
hD1 — height of the last spiny ray of the dorsal fin; hD2 — maximum height of the soft
part of the dorsal fin; lA — length of the anal fin base; hA — height of the third spiny
ray of the anal fin; lP — pectoral fin length; lV — ventral fin length; hll — distance
from the base of the 2nd spiny ray in dorsal fin to the lateral line; C — head length; hC
— head height at the vertical of the middle of the eye; r — snout length (from the begin-
ning of the upper lip to the anterior edge of the eye); o — horizontal diameter of the eye;
po — postorbital length; io — interorbital length. The following meristic characters
were also counted: D1 — the number of spiny rays in dorsal fin; d2 — the number of soft
rays in dorsal fin; A — the number of spiny rays in anal fin; a — the number of soft rays
in anal fin; P — the number of soft rays in pectoral fin; l.l. — the number of scales
along the lateral side of the body; sp. br. — the number of gill rakers on the first gill
arch.
The sex of the fish was determined by the shape and color of the urogenital papilla (in
females it is large and black).
Differentiation of individuals according to the absolute values of morphometric char-
acters was studied using hierarchical cluster analysis. The Euclidean distance — DE —
was calculated as a measure of the generalized differences between individuals on
absolute values of 26 plastic characters. Based on the fact that the size and age of fish
strongly correlate with each other (Cherepanov, 1986), the groups that emerged were
considered as size-age classes. For each sample, the standard statistical parameters of
the variational series were calculated: minimum (min), maximum (max), arithmetic
average (M) values of the character, arithmetic average error (m), and standard deviation
(SD). The significance of differences between the compared samples was estimated by
the average values of characters using Student’s t-test. The similarity in the structure of
sex and size-age differences was estimated using the Spearman rank correlation coeffi-
cient (Rs), comparing the consistency in the variation of the values of Student's t-test
for 26 plastic characters (Lakin, 1990). Differences at a 5 % level of significance were
considered as statistically significant. Differences between the size-at-age and sex
groups of Ocellated wrasse were analyzed over the entire set of characters using discri-
minant analysis. As a measure of generalized differences in the complex of absolute and
relative plastic characters, we calculated the Squared Mahalanobis Distance SqMD. The
studied material was processed statistically using the STATISTICA 6.0 and Microsoft
Office Excel software packages.
Results and discussion
Brief biological characteristics . The species distribution covers the waters of the
East Atlantic (Bay of Biscay), the Mediterranean, Adriatic, Aegean, Marmara,
6 Зб. пр. Зоол. муз., 49, 2018
V. N. Peskov, L. G. Manilo,
Black Sea, and the western part of the Azov Sea to the Berdyansk Spit (Svetovidov,
1964; Diripasco et al., 2011; Vasil’eva, Luzhnyak, 2013; Bilecenoglu et al., 2002;
Golani et al., 2006). In the waters of Ukraine this species is found from the Zmeinyi
Island (Snigirev, 2008) and further eastward from the Danube coast to the Kerch Strait in
the waters of Ukraine (Vinogradov, 1960; Pinchuk, Tkachenko, 1996; Boltachev, Kar-
pova, 2012). It was often observed in the Odesa Bay until 2003, however, it was not
registered here later (Chernikova, Zamorov, 2011). The isolated population is known
from the Tiligul estuary (Kovtun, Tarasenko, 2005). Target species inhabits coastal
biotopes with rocky and (less often) sandy soils among the thickets of underwater vegeta-
tion to a depth of 30 m, but is most numerous in shallow waters (up to 5–10 m). The food
spectrum includes invertebrates: small crustaceans, mollusks, sea worms, as well as
bryozoans, hydroid polyps and plants (Vasil’eva, 2007; Quignard, Pras, 1986), and diet
changes throughout the year are a specific adaptive response to transformations in the
environment (Ouannes-Ghorbel et al., 2003). Males grow faster than females (Quignard,
Pras, 1986). In the Black Sea, it reaches sexual maturity at the age of one year at a length
of about 3.5 cm (Salekhova, 1971). Spawning is multiportional. Reproduction in the
Black Sea continues from April to August under water temperature from 12 to 26 °C,
spawning peaks from the end of May to the end of June (Salekhova, 1971). Species
biology has been well studied in the Mediterranean Sea (Taborsky et al., 1987; Alonso,
Wagner, 1999, 2000; Kabasakal, 2001; Alonso, 2008; Lipej et al., 2009; Alonso, Heck-
man, 2010). Standard body length reaches 14 cm, usually 7–8 cm (Vasil’eva, 2007). In
the Black Sea, the age limit for females is 3+, and 5+ years for males (Salekhova, 1971).
Brief diagnosis . D XIII–XV+8–11; A III+8–11; P 12; V I 5; l.l. 30
(Golani et al., 2006, Mediterranean Sea). D (XIII) XV+(9) 10; A III+(9) 10; P I+11–12;
l.l. 32–34 (Svetovidov, 1964). D XIII–XV+8–11; A III+8–11; l.l. 30–34; sp.br. 14–18
(Vasil’eva, 2007, Black Sea). D XIV+9–10; A III+9–10; P I+11–12; V I 5; l.l. (31) 32–
34 (Movchan, 2011, Black Sea). D XIII–XV (M=13.89±0.040)+9–11 (M=9.95±0.050);
A III+9–11 (M=9.91±0.051); P 11–13 (M=12.01±0.022); l.l. 30–34 (M=31.86±0.094);
sp.br. 14–16 (M=14.78±0.080) (original data, northern part of the Black Sea).
The distance from the base of the second spiny ray in dorsal fin to the lateral line is less
than half as long as the soft part of the dorsal fin (37.29–50.00 %, mean 47.13 % in
males and 44.19 % in females). Body height is equal or slightly greater than the head
length. The dorsal fin is long, consists of spiny and soft parts. The length of its base
averages 53.73 % of the standard body length (SL) in males and 52.82 % in females. The
pectoral fins are longer than ventral, with an average of 21.63 % SL in males and 20.52
% in females. Ocellated wrasse reaches sexual maturity at the age of one year at a length
of about 3.5 cm in the Black Sea. Therefore, all investigated specimens are mature.
Size and age variabili ty . According to the results of hierarchical cluster analysis
(Fig. 1), both males and females of the Ocellated wrasse are differentiated into small
(SL♂ = 43–70 mm and SL♀ = 42–56 mm) and large (SL♂ = 71–93 mm and SL♀ = 59–72
mm) individuals.
Differences between large and small individuals are highly significant in the mean
absolute values of 26 characters in both males (Table 2) and females (Table 3). At the
same time, in females these differences are noticeably larger in most characters than
those in males, with the exception of three signs (aP, hD2, hA), the size-age differences
for which are greater in males. The generalized differences between small and large
individuals in males (SqMD = 21.99) are slightly larger than in females (SqMD = 16.48).
In large males, the relative values of 10 characters are on average significantly greater
than those of small males, while the relative value of the anteventral length (aV), the
height of the 3rd spiny ray of the anal fin (hA), the length of the snout (r) and the hori-
zontal diameter of the eye (o), on the contrary, are significantly bigger in small males
(Table 4).
Zb. prac’ Zool. muz. (Kiïv), 49, 2018 7
Size-at-age and sex variability of plastic values in Ocellated wrasse ...
Fig. 1. Differentiation of males and females of Symphodus ocellatus in absolute values of 26
plastic characters.
Ward`s method
Squared Euclidean distances
0
20
40
60
80
100
120
(D
li
n
k
/D
m
a
x
)*
1
0
0
Males
0
20
40
60
80
100
120
(D
lin
k
/D
m
a
x
)*
1
0
0
Females
8 Зб. пр. Зоол. муз., 49, 2018
V. N. Peskov, L. G. Manilo,
Table 2. The results of the comparison of absolute values (mm) of plastic characters in two
size-age groups of Symphodus ocellatus (males)
Note. min-max — minimum and maximum values of the character; M — mean value of the
character; SD — standard deviation; tst — Student’s criterion, *** P < 0.001.
Charac-
ter
Small individuals (n = 20) Large individuals (n = 22)
tst min max M SD min max M SD
SL 43.0 70.0 59.65 7.11 71.0 93.0 78.68 6.61 8.96***
H 13.1 25.9 20.81 3.59 22.9 35.1 28.94 2.76 8.16***
h 5.7 10.3 8.40 1.39 10.2 13.3 11.54 0.93 8.54***
aD1 14.4 23.5 20.29 2.40 23.6 32.8 26.74 2.49 8.55***
aD2 28.9 47.3 41.09 4.70 47.5 66.2 54.46 4.83 9.09***
aP 16.3 24.2 20.53 1.92 22.7 33.5 26.50 2.88 7.97***
aV 18.7 31.3 25.53 2.82 28.6 41.8 32.54 3.73 6.91***
aA 26.5 43.2 37.14 4.22 43.3 60.4 48.46 4.95 8.00***
pD 6.6 11.5 8.88 1.33 9.2 13.7 11.25 1.11 6.24***
lcaud 7.1 10.9 8.83 1.16 10.0 12.7 11.33 0.86 7.86***
lD 21.4 36.8 31.35 4.49 38.0 54.6 43.21 4.06 8.95***
lD1 15.4 26.8 21.27 2.95 24.5 36.3 29.14 2.57 9.17***
lD2 6.0 13.1 10.08 1.85 11.2 18.3 14.08 1.81 7.08***
hD1 6.6 9.7 7.88 0.87 8.2 12.4 9.85 1.09 6.54***
hD2 6.0 11.7 9.05 1.50 9.3 16.7 13.46 1.99 8.16***
lA 10.6 17.7 14.75 2.18 17.8 26.1 20.54 2.20 8.56***
hA 5.4 8.0 6.73 0.81 6.3 9.5 8.13 0.89 5.36***
lP 9.1 14.9 12.75 1.58 13.7 21.6 17.21 1.73 8.72***
lV 7.3 12.6 10.36 1.50 11.3 19.8 14.29 1.96 7.32***
hll 2.8 6.1 4.72 0.85 5.5 8.9 6.65 0.87 7.28***
C 15.6 23.8 20.85 2.35 24.6 33.7 27.66 2.70 8.75***
hC 8.0 14.1 11.43 1.73 12.7 19.4 15.73 1.74 8.03***
r 5.2 8.1 6.76 0.72 7.3 11.8 8.60 1.20 6.08***
o 3.2 4.9 4.23 0.39 4.5 5.8 5.20 0.30 9.11***
po 6.9 11.7 9.61 1.54 11.2 16.2 13.36 1.28 8.53***
io 2.9 5.5 4.33 0.87 4.7 7.2 5.94 0.63 6.84***
Zb. prac’ Zool. muz. (Kiïv), 49, 2018 9
Size-at-age and sex variability of plastic values in Ocellated wrasse ...
Table 3. The results of the comparison of absolute values (mm) of plastic characters in two
size-age groups of Symphodus ocellatus (females)
Note. min-max — minimum and maximum values of the character; M — mean value of the
character; SD — standard deviation; tst — Student’s criterion, *** P < 0.001.
Char-
acter
Small individuals (n = 21) Large individuals (n = 17)
tst
min max M SD min max M SD
SL 42.0 56.0 48.00 3.94 59.0 72.0 64.24 3.65 13.16***
H 14.0 20.5 16.73 1.82 19.9 26.1 22.84 1.89 10.07***
h 4.9 7.3 6.18 0.65 7.7 9.5 8.46 0.61 11.16***
aD1 13.3 20.2 16.45 1.74 19.0 25.5 22.16 1.87 9.65***
aD2 28.6 39.2 33.33 2.79 39.8 50.7 44.00 3.27 10.68***
aP 13.8 20.6 16.44 1.87 18.1 25.9 21.55 2.11 7.81***
aV 16.6 26.4 19.99 2.47 24.0 33.7 27.34 2.53 8.99***
aA 26.3 38.4 30.86 3.28 37.2 50.8 42.78 3.65 10.47***
pD 5.7 8.4 6.79 0.69 8.2 10.2 9.32 0.70 11.16***
lcaud 5.8 7.7 6.58 0.58 7.9 10.5 8.75 0.65 10.79***
lD 21.5 29.6 25.44 2.08 30.8 38.1 33.78 2.35 11.44***
lD1 15.4 20.2 17.46 1.28 20.1 26.3 22.84 1.67 10.92***
lD2 6.1 10.8 7.98 1.09 9.1 13.7 10.94 1.33 7.38***
hD1 4.5 7.0 6.29 0.61 6.8 9.0 7.72 0.64 6.98***
hD2 5.2 8.2 6.77 0.89 7.9 10.5 8.96 0.86 7.72***
lA 9.0 11.8 10.59 0.80 12.6 16.4 14.66 1.21 11.97***
hA 4.2 6.6 5.54 0.60 5.6 7.4 6.49 0.56 5.06***
lP 8.1 11.7 9.80 1.10 12.0 15.1 13.28 0.85 10.97***
lV 6.5 9.8 8.17 0.93 9.2 12.2 10.68 0.79 8.97***
hll 2.8 4.6 3.51 0.46 4.1 6.0 4.82 0.50 8.38***
C 14.3 20.7 16.82 1.90 19.8 26.0 22.29 1.90 8.83***
hC 6.0 10.6 8.62 1.10 10.7 14.2 12.05 1.13 9.42***
r 3.8 6.9 4.92 0.79 5.4 8.1 6.72 0.82 6.84***
o 3.4 4.3 3.84 0.24 4.1 5.0 4.68 0.25 10.45***
po 6.4 9.3 7.54 0.98 8.9 12.3 10.46 1.09 8.60***
io 2.7 3.8 3.33 0.29 3.9 5.1 4.57 0.35 11.68***
10 Зб. пр. Зоол. муз., 49, 2018
V. N. Peskov, L. G. Manilo,
Large females have significantly higher relative values of aA, hC and po than small
females. The relative values of hD1, hA, and o are significantly higher in small females
(Table 5). The generalized differences in the relative values of plastic characters are
much bigger in females (SqMD = 8.03) in contrast to males (SqMD = 5.44).
Table 4. Comparison results of relative values (%) of plastic characters in two size-age
groups of Symphodus ocellatus (males)
Note. min-max — minimum and maximum values of the character; M — mean value of the
character; SD — standard deviation; tst — Student’s criterion, * — at P < 0/05; ** — at P < 0/01;
*** — P < 0/001.
Charac-
ter
Small individuals (n = 20) Large individuals (n = 22)
tst
min max M SD min max M SD
SL 53.0 85.0 74.40 8.93 89.0 120.0 87.10 14.99 8.96***
H 43.0 70.0 59.65 7.11 71.0 93.0 69.62 11.76 2.89***
h 30.5 39.2 34.69 2.59 30.1 40.8 35.80 2.54 2.64**
aD1 12.6 16.1 14.01 0.93 13.6 16.2 14.36 0.88 0.17
aD2 32.4 36.0 34.02 1.05 32.6 35.8 33.99 1.00 0.42
aP 65.1 74.5 68.95 2.25 64.2 72.5 69.09 2.00 1.72
aV 32.0 37.9 34.55 1.50 29.7 38.1 34.08 1.73 2.50**
aA 39.8 47.0 42.90 2.01 38.7 46.3 42.06 2.20 1.50
pD 59.2 67.2 62.33 1.88 59.3 65.7 61.91 1.77 1.76
lcaud 13.2 16.9 14.87 1.15 11.7 15.5 14.58 1.04 1.49
lD 13.4 17.0 14.82 0.95 12.8 15.8 14.61 0.87 3.82***
lD1 48.9 55.8 52.42 2.28 50.3 58.7 53.73 2.42 2.39**
lD2 32.0 40.6 35.63 2.01 32.8 39.9 36.38 2.05 2.39**
hD1 14.0 19.8 16.80 1.65 15.6 20.5 17.35 1.48 1.88
hD2 10.6 15.5 13.32 1.68 11.1 14.2 12.91 1.38 3.96***
lA 11.7 17.7 15.14 1.41 12.7 19.6 16.15 1.86 3.56***
hA 21.6 27.3 24.67 1.36 23.6 28.3 25.40 1.44 2.45**
lP 8.8 14.0 11.39 1.63 8.5 12.5 10.85 1.43 1.67
lV 20.2 22.7 21.36 0.68 18.8 23.4 21.63 1.03 2.01
hll 15.7 19.8 17.33 1.07 15.5 21.3 17.74 1.33 0.48
C 38.4 50.0 46.90 3.09 40.7 50.0 47.13 2.87 0.43
hC 33.5 36.3 34.98 0.84 32.2 37.5 35.07 1.22 2.09*
r 49.5 60.3 54.65 3.22 49.9 66.9 55.84 3.68 2.49**
o 28.8 34.9 32.49 1.89 27.5 35.0 31.71 2.06 3.87***
po 18.5 22.9 20.35 1.23 16.5 21.4 19.59 1.40 2.73***
io 40.7 51.6 45.89 3.33 43.5 51.5 47.18 3.10 1.65
Zb. prac’ Zool. muz. (Kiïv), 49, 2018 11
Size-at-age and sex variability of plastic values in Ocellated wrasse ...
Sex differences . In both size-age groups, males are significantly larger than fe-
males (Tables 3, 4, 6). At the same time, the differences between large males and fe-
males on most characters are significantly bigger than those between small ones. Sex
differences only on 4 characters (aP, aV, aA and r) are bigger in group of small Ocellat-
ed wrasse.
Table 5. Comparison results of relative values (%) of plastic characters in two size-age
groups of Symphodus ocellatus (females)
Note. min-max — minimum and maximum values of the character; M — mean value of the
character; SD — standard deviation; tst — Student’s criterion, *** P < 0.001.
Charac-
ter
Small individuals (n = 21) Large individuals (n = 17)
tst min max M Sx min max M SD
SL 53.0 70.0 60.67 5.10 73.0 89.0 80.06 4.64 13.16***
H 42.0 56.0 48.00 3.94 59.0 72.0 64.24 3.65 1.22
h 31.4 37.9 34.82 1.90 32.1 38.4 35.54 1.70 1.87
aD1 11.7 13.6 12.85 0.52 12.2 14.2 13.17 0.51 0.42
aD2 30.0 36.7 34.24 1.66 31.7 37.7 34.47 1.68 1.61
aP 66.9 71.6 69.45 1.20 65.6 72.8 68.46 2.27 1.07
aV 31.1 38.0 34.19 1.76 30.2 36.8 33.52 2.07 1.29
aA 38.3 48.0 41.54 2.33 39.7 47.3 42.51 2.31 2.77***
pD 60.2 69.8 64.21 2.52 62.0 71.1 66.52 2.58 1.15
lcaud 12.8 15.9 14.14 0.98 12.1 16.3 14.53 1.04 0.28
lD 12.2 15.7 13.73 0.81 12.4 15.7 13.65 0.98 0.83
lD1 48.5 55.7 53.02 1.81 49.7 55.9 52.57 1.60 1.64
lD2 33.6 38.9 36.43 1.69 32.4 38.7 35.55 1.62 0.82
hD1 13.8 19.3 16.60 1.40 14.0 20.5 17.01 1.65 2.87***
hD2 9.0 15.2 13.15 1.44 10.4 14.4 12.03 0.96 0.37
lA 11.3 16.0 14.09 1.35 12.7 15.6 13.95 0.95 1.73
hA 20.0 24.9 22.10 1.22 21.2 25.5 22.83 1.32 3.97***
lP 8.4 13.6 11.58 1.31 8.6 11.8 10.12 0.96 0.88
lV 18.5 22.3 20.39 1.14 19.4 22.3 20.68 0.87 1.18
hll 14.1 18.8 17.01 1.13 14.8 17.5 16.63 0.82 0.19
C 37.8 49.2 44.11 3.13 37.3 49.5 44.29 3.01 0.59
hC 31.1 37.6 34.99 1.75 32.4 37.7 34.67 1.63 3.81***
r 41.1 55.1 51.20 2.78 51.0 58.3 54.07 1.85 1.18
o 26.1 34.2 29.17 2.49 25.9 34.0 30.12 2.45 3.81***
po 20.3 27.2 23.00 1.86 18.8 22.7 21.08 1.24 3.47***
io 42.4 47.7 44.74 1.37 43.5 50.5 46.92 2.28 1.57
12 Зб. пр. Зоол. муз., 49, 2018
V. N. Peskov, L. G. Manilo,
Sex differences by the relative size of plastic characters are expressed in both size-
age groups of males and females of Ocellated wrasse. The mean values of 7 relative
characters (h, lcaud, hD2, lA, lP, hll and hC) are significantly higher in males of both size-
age groups and only 2 characters (aA and o) are greater in females in contrast to males.
In addition, sex differences are significant in two characters (pD and r) in the group of
small individuals only and in four characters (lD, lD1, lV and io) in the group of large
individuals (in both cases, males have average values of relative characters). The gener-
alized differences between males and females in the group of small individuals are
somewhat larger in the body proportions (SqMD = 23.21) than in its linear dimensions
(SqMD = 19.53). On the contrary, the generalized sex differences in the body linear
dimensions (SqMD = 45.50) are noticeably bigger than those in its proportions
(SqMD = 32.60) in group of large individuals.
Data given above showed that sex differences in Ocellated wrasse increase with age
both in absolute and relative values of plastic characters; it is accompanied by increased
morphological differentiation between males and females in overall size and body
proportions.
The expression and directionality of sex differences in Ocellated wrasse are mani-
fested in the fact that the mean values of all 26 plastic characters in males are signifi-
cantly higher compared with females (Tables 3, 4). At the same time, in group of large
mature fishes, the expression of sex differences on 22 characters is greater than in small
ones, and only in 4 is the opposite (Table 6).
Table 6. The magnitude of sex differences (tst) in absolute and relative values of plastic
characters in two size-age groups of Symphodus ocellatus
Charac-
ter
Absolute values Relative values
Large Small Large Small
SL 8.68*** 6.45*** 8.68*** 6.45***
H 8.17*** 4.55*** 2.09* 0.18
H 12.47*** 6.50*** 7.76*** 4.90***
aD1 6.56*** 5.83*** 1.10 0.51
aD2 8.06*** 6.39*** 1.13 0.88
aP 6.19*** 6.92*** 0.23 0.70
aV 5.18*** 6.68*** 1.68 2.00
aA 4.13*** 5.30*** 7.01*** 2.71*
pD 6.65*** 6.24*** 0.68 2.17*
lcaud 10.65*** 7.80*** 2.71* 3.96***
lD 9.10*** 5.37*** 4.19*** 0.93
lD1 9.24*** 5.32*** 2.71* 1.38
lD2 6.26*** 4.41*** 1.80 0.42
HD1 7.66*** 6.71*** 1.62 0.35
hD2 9.52*** 5.89*** 7.09*** 2.44*
lA 10.63*** 8.03*** 7.96*** 6.34***
hA 7.05*** 5.33*** 0.71 0.42
lP 9.30*** 6.88*** 3.54** 3.34**
lV 7.83*** 5.55*** 4.03*** 0.94
hll 8.24*** 5.61*** 3.27** 2.88**
Zb. prac’ Zool. muz. (Kiïv), 49, 2018 13
Size-at-age and sex variability of plastic values in Ocellated wrasse ...
An increase of the body length in large individuals in contrast to small ones, as well as
in males in contrast to females, are accompanied by an increase in the size of all other
25 characters. It is important to find out if the structure of the variability of all 26 plastic
characters is the same in all four cases. By comparing the difference vectors (Student’s t
-values) using the Spearman rank correlation coefficient (Rs), an average level of simi-
larity was established (Rs = 0.54; p < 0.01) in the structure of differences between small
and large individuals in males and females. The average level of similarity (Rs = 0.52;
p < 0.01) was also noted between the structure of sex differences in group of large
individuals and size-age differences in females. In all other cases, the level of similarity
of the vectors of differences was below the average level (0.09–0.39).
The results of the study confirm the data of Shevchenko (1971) that males have larger
relative sizes of the anal fin base, while females have larger eye diameter than males. At
the same time, our data contradict the conclusion of Shevchenko (1971) that males are
more tall and have larger relative sizes of the head and dorsal fin. According to our data,
the males are characterized by significantly higher relative values of the height and
length of the caudal peduncle, the height of the soft part of the dorsal fin, pectoral fin
length, head height and the snout length, as well as the females have larger anteanal
distance (Tables 4, 5). In all cases, the value of the Student’s t-test for the aforemen-
tioned features significantly exceeds the critical table values.
Female mate choice copying affects sexual selection in wild
populations of the ocellated wrasse. Animal behaviour., 75, 1715–1723.
doi:10.1016/j.anbehav.2007.09.031.
The unexpected but understandable
dynamics of mating, paternity and paternal care in the Ocellated wrasse. Proc. R. Soc. B.,
277, 115–122. doi:10.1098/rspb.2009.1425.
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Mediterranean wrasse males refuse present mates to increase future success. Behavioral
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. Dynamic games and field experiments
examining intra- and intersexual conflict: explaining counterintuitive mating behavior in a
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the marine fishes of Turkey. Zootaxa, 113, 1–194.
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la. Simferopol: Biznes-Inform, 1–224 [In Russian].
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zhivotnyh. Novosibirsk: Nauka. 1–240.
Note. * — at P < 0/05; ** — at P < 0/01; *** — P < 0/001.
Charac-
ter
Absolute values Relative values
Large Small Large Small
C 7.29*** 6.02*** 0.93 0.02
hC 7.97*** 6.19*** 3.07** 3.67***
R 5.80*** 7.77*** 1.21 4.82***
o 5.95*** 3.76*** 5.56*** 5.42***
po 7.62*** 5.09*** 1.91 1.44
io 8.62*** 4.87*** 2.29* 1.20
14 Зб. пр. Зоол. муз., 49, 2018
V. N. Peskov, L. G. Manilo,
Fish fauna of the Odessa Bay (the
Black Sea) in the first decades of the XXI century. Morskyi ecologichnyi zhurnal. 10, 3, 76–
85 [In Russian].
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the Azov Sea. Berdyansk: Izd-vo OOO “NPK Inter-M”. 1–288 [In Russian].
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1–384.
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sympatric labrids (Perciformes, Labridae) from south-eastern Aegean Sea, Turkey. Nether-
lands Journal of Zoology, 51 (4), 439–455.
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ischezajuschih vidov gidrobiontov severnoi chasti Chernogo morya. Ekologiya ta suspilstvo.
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M. Gor’kogo, vyp. 2, Odesa, 112–124.
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revised and enlarged. Moscow, Vysschaja shkola. 1–352 [In Russian].
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characteristics of three labrid species in the Gulf of Trieste (northern Adriatic Sea). Acta
Adriatica, 50 (2), 139–150.
A review of techniques for quantifying
sexual size dimorphism. Growth, Development and Aging, 56, 269–281.
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444 [In Ukrainian].
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Natural History, Ukrainian Academy of Sciences. Cyclostomata and Fishes. Kiev: Zoological
Museum NMNH of Ukraine, 1–342 [In Russian].
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Inst. Natn. Scien. Tech. Mer de Salammbô, 30, 33–37.
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direction and structure of sexual differences in amphibians and reptiles: a case study on Rana
temporaria Linnaeus, 1758 and Lacerta viridis Laurenti, 1768. Zbirnik prac Zoologicnogo
muzeu, 48, 54–69.
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skogo biosfernogo zapovednika (annotirovannye spiski vidov). Ryby morskih akvatoriy.
Vestnik Zoologii. Otdeln. vyp., 1, 5–14.
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Embrionalnyi i ranniy postembrionalnyi periody razvitiya
chernomorskih zelenushek roda Crenilabrus. Biologiya morya. Vyp. 23. Voprosy ekologii ryb
yuznyh morey. (Respubl. mezhvedomstvennyi sborn.). Kiev: Naukova dumka, 36–77 [In
Russian].
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zelenushek roda Crenilabrus v Chernom more. Biologiya morya. Vyp. 23. Voprosy ekologii
ryb yuznyh morey. (Respubl. mezhvedomstvennyi sborn.). Kiev: Naukova dumka, 78–96 [In
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line and Anadromous Species with Color Illustrations Collected by S.V. Bogorodsky. Mos-
cow, VNIRO Publishing: 1–238 [In Russian].
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В. М. Пєсков, Л. Г. Маніло
РОЗМІРНО-ВІКОВА І СТАТЕВА МІНЛИВІСТЬ ПЛАСТИЧНИХ ОЗНАК ЗЕЛЕНУШКИ
ПЛЯМИСТОЇ, SYMPHОDUS OCELLATUS (LINNAEUS, 1758) (LABRIDAE, PERCI-
FORMES), У ПІВНІЧНІЇ ЧАСТИНІ ЧОРНОГО МОРЯ
Наведено результати вивчення розмірно-вікової і статевої мінливості пластичних ознак у
пізньому онтогенезі зеленушки плямистої, Symphodus ocellatus (Linnaeus, 1758) в північній
частині Чорного моря. Встановлено, що самці і самки за абсолютними значеннями 26
пластичних ознак диференціюються на дрібних і великих особин, відмінності між якими
високо достовірні за всіма ознаками. За відносними значеннями ознак відмінності достові-
рні у самців за 14 ознаками, у самок — за 6. Показано, що з віком у S. ocellatus збільшуєть-
ся вираженість статевих відмінностей як за абсолютними, так і за відносними значеннями
пластичних ознак. Серед великих і дрібних особин зеленушки самці достовірно більші за
самок. З віком у зеленушки плямистої змінюються структура вираженості і спрямованості
статевих відмінностей.
К л ю ч о в і с л о в а: пластичні ознаки, розмірно-вікова і статева мінливість,
Symphodus ocellatus, Чорне море.
В. Н. Песков, Л. Г. Манило
РАЗМЕРНО-ВОЗРАСТНАЯ И ПОЛОВАЯ ИЗМЕНЧИВОСТЬ ПЛАСТИЧЕСКИХ ПРИ-
ЗНАКОВ ЗЕЛЕНУШКИ ГЛАЗЧАТОЙ, SYMPHODUS OCELLATUS (LINNAEUS, 1758)
(LABRIDAE, PERCIFORMES) В СЕВЕРНОЙ ЧАСТИ ЧЕРНОГО МОРЯ
Приведены результаты изучения размерно-возрастной и половой изменчивости пластиче-
ских признаков в позднем онтогенезе зеленушки глазчатой, Symphodus ocella-
tus (Linnaeus 1758) в северной части Черного моря. Установлено, что самцы и самки по
абсолютным значениям 26 пластических признаков дифференцируются на мелких и
крупных особей, различия между которыми высоко достоверны по всем признакам. По
относительным значениям признаков различия достоверны у самцов по 14 признакам, у
самок — по 6. Показано, что с возрастом у S. ocellatus увеличивается выраженность поло-
вых различий как по абсолютным, так и по относительным значениям пластических
признаков. Среди крупных и мелких особей зеленушки самцы достоверно крупнее самок.
С возрастом у зеленушки глазчатой изменяются структура выраженности и направленно-
сти половых различий.
К л ю ч е в ы е с л о в а: пластические признаки, размерно-возрастная и половая изменчи-
вость, Symphodus ocellatus, Черное море.
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