https://doi.org/10.52973/rcfcv-e34366
Received: 15/12/2023 Accepted: 30/01/2024 Published: 11/04/2024
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Revista Científica, FCV-LUZ / Vol. XXXIV, rcfcv-e34366
ABSTRACT
In this study, the Tigris bream Acanthobrama marmid individuals (44
females and 31 males) were captured from the Tigris River. The scale
size (as centroid size) and shape were analyzed separately using 2–
dimensional geometric morphometric methods. Procrustes ANOVA
revealed signicant differences in scales size between sexes, while
no difference in shape was observed. Groups based on season and
age showed signicant differences in both size and shape. Female
individuals had larger scale sizes than males, with the scales of the
Autumn group being larger than those of the Spring and Summer
groups. Scale size also increased with age groups. PCA analysis
showed variation in the rst ve components when examined by
age, season, and gender. CVA and DFA results indicated signicant
differences in shape between different age groups and seasonal
groups, but no signicant differences between sexes were observed.
Key words: Leuciscidae; geometric; landmark; morphometric;
scale; shape; Türkiye
RESUMEN
En este estudio, se capturaron individuos de la brema del Tigris
Acanthobrama marmid (44 hembras y 31 machos) del río Tigris.
El tamaño y la forma de las escamas se analizaron por separado
utilizando métodos morfométricos geométricos bidimensionales. El
análisis de la ANOVA de Procrustes reveló diferencias signicativas
en el tamaño de las escamas entre los géneros, mientras que no
se observaron diferencias en la forma. Los grupos basados en la
temporada y la edad mostraron diferencias signicativas tanto en
tamaño como en forma. Los individuos hembra tenían tamaños de
escamas más grandes que los machos, siendo las escamas del grupo
de otoño más grandes que las de los grupos de primavera y verano. El
tamaño de las escamas también aumentó con los grupos de edad. El
análisis de PCA mostró variación en los primeros cinco componentes
al examinar por edad, temporada y género. Los resultados de CVA y
DFA indicaron diferencias signicativas en forma entre diferentes
grupos de edad y grupos estacionales, pero no se observaron
diferencias signicativas entre géneros.
Palabras clave: Leuciscidae; geométrico; punto de referencia;
morfométrico; escama; Turquía
Variation in the shape and size of the scale of the Tigris bream
(Acanthobrama marmid, Heckel, 1843) from the Tigris River, Türkiye
attributed to Seasonality, Age and Sex: A geometric morphometric study
Variación Estacional, por Sexo y Edad de las características de las escamas de la brema del Tigris
(Acanthobrama marmid, Heckel, 1843) del rio Tigris, Turquia: un estudio morfométrico geométrico
Serbest Bilici
1
* , Alaettin Kaya
2
, Muhammed Yaşar Dörtbudak
3
, Tarık Çiçek
4
, Erhan Ünlü
4
1
Şirnak University, Faculty of Agriculture, Department of Animal Science. Sirnak, Türkiye.
2
Dicle University, Faculty of Veterinary Medicine Faculty, Department of Basic Science. Diyarbakır, Türkiye.
3
Harran University, Faculty of Veterinary, Department of Fisheries And Diseases. Şanlıurfa, Türkiye.
4
Dicle University, Faculty of Science, Department of Biology. Diyarbakır, Türkiye.
*Corresponding Author: serbestbilici@hotmail.com.tr
FIGURE 1. The overall body appearance of Acanthobrama marmid (Tigris bream),
Dicle River (Photo by E. Ünlü)
FIGURE 2. Map of the study area which samples obtained. Sample localities
(1. Tigris River (Güçlükonak–1), 2. Tigris River (Güçlükonak–2), 3. Tigris River
(Akdizgin), 4. Tigris River (Damlarca)
Variation in scale characteristics of Acanthobrama marmid / Bilici et al. ____________________________________________________________
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INTRODUCTION
Acanthobrama marmid Heckel, 1843 is a member of the Leuciscidae
family and is found in the Tigris–Euphrates River system, Kuveyk
and Asi Rivers, and likely in Amik Lake and Bardan Stream near
Tarsus [1, 2, 3]. According to Küçük et al. [4], Acanthobrama
marmid is only distributed in the Tigris–Euphrates system, while the
populations in Asi, Seyhan, and Berdan River (Tarsus) are identied
as Achantobrama orontis.
This species is characterized by its compressed body structure and
humped back structure on the back of the head, which is especially
evident in large individuals. It does not have whiskers and has small
scales. A eshy keel is located between the base of the pelvic ns
and the ventral n. Additionally, it possesses a thick, spine–like, and
smooth terminal unbranched dorsal n ray, as well as a long anal n
(15–22 branches) [2, 3]. Its ns are orange–red in color (FIG. 1). It is
a benthopelagic species, typically found in shallow, slow–moving
waters with sandy or muddy bottoms. Acanthobrama marmid plays
a vital role in the ecosystem as a prey species for larger predatory
sh and an important component of the food web. In rural areas, the
local population consumes it [5].
Scales are structures embedded in the epidermal layer of the
sh's body, which are also used in species identication. Ctenoid
and cycloid scales are particularly used as identication tools in
systematic studies. Compared to molecular techniques, scales are
cost–effective, non–destructive, convenient to use, and can serve
as suitable bony structures for species identication due to their
resistance to digestion by predators' digestive systems [6, 7]. Scale
morphology is used in taxonomy and classication studies and has
been evaluated for ontogenetic analyses [8] and morphology [9, 10, 11,
12]. The morphological and morphometric characteristics of scales
are one of the methods used in the identication and differentiation
of fish species and populations [13, 14, 15, 16, 17]. Biological
characteristics and age determination studies of Acanthobrama
marmid are available [18, 19, 20, 21, 22, 23].
The study utilized geometric morphometric methods to ascertain
the distinctive structure of scales attributed to the species and to
discern variations among season, age, and male and female individuals.
MATERIAL AND METHODS
In this study, we collected a total of 75 specimens of Acanthobrama
marmid, including 44 females and 31 males, from the Tigris River. The
localities where the samples were collected are shown in FIG. 2, and
the seasonal, sexual, and age distributions of the samples, as well
as some water parameters of the locality where they were collected,
are given in TABLE I.
The sex of each sh was determined by observing their gonads. Scales
from the front and upper sections of the lateral lines of the dorsal ns were
taken to determine their age and morphology. The sh scales tissue was
cleaned with 5% NaOH for 2 hours, then washed with distilled water, and
immersed in 96% ethanol for several minutes to remove any remaining
water. Following this the scales were placed between two slides and
photographed by an stereo microscope (Olympus SZX7, Tokyo, Japan) and
a digital camera (OLYMPUS Camedia C–5060 5.1 MP w/4× Optical Zoom,
Tokyo, Japan) under 20× and 40× magnications. Images were analyzed
TABLE I
Samples distribution and water parameters of the study.
Season Female Male Total
Autumn (November)
12 – 12
Spring (April)
22 16 38
Summer (July)
10 15 25
Age
II III IV V VI VII
Sample number
7 23 23 15 5 2
Date
Water
temperature
(°C)
pH
dissolved
oxygen (O
2
)
O
2
(%)
Electrical
conductivity
(EC) µS·cm
-1
26/04/2021 17.7 8.3 9.11 101 306
01/07/2021 24.6 7.86 7.67 97.2 474
04/11/2021 13.5 8.2 8.62 96.8 365
FIGURE 3. Landmark denitions used in the sh scales
FIGURE 4. Box and Violin plot of CS of scales by sex F: female, M: male
FIGURE 5. Box and Violin plot of CS of scales by season Au: autumn, Sm: summer,
Sp: spring
FIGURE 6. Box and Violin plot of CS of scales by age. Numbers represent ages
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by geometric morphometric procedure [24, 25, 26]. Subsequently, six
landmarks (FIG. 3) were digitized using tpsDig ver. 2.32 [27] software, and
Procrustes analysis was conducted. Following the separation of shape
and size (centroid size=CS) of the samples, Procrustes ANOVA, PCA, CVA/
MANOVA, and DFA analyses were performed using Morpho J1.06d [28],
R Core Team [29] and Jamovi Ver. 2.4 [30] programs.
RESULTS AND DISCUSSION
When the results of Procrustes ANOVA are examined, a signicant
difference in size (CS) between sex is found (P=0.0051), while no
difference in shape is observed. Groups based on season and age
are signicant in both size and shape (P<0001) (TABLE II).
TABLE II
Procrustes ANOVA results (F: Goodal’s F, CS: Centroid Size)
F P–value Pillai tr. P–value
Sex
CS 8.33
0.0051
Shape
0.91 0.5110 0.13 0.2719
Season
CS
13.97
< 0.0001
Shape
4.51
< 0.0001 0.51 0.0006
Age
CS
11.12
< 0.0001
Shape 2.44 < 0.0001 0.83 0.0101
In female individuals, scale size is larger than in males, and the
scales of the Autumn group are larger than those of the Spring and
Summer groups. Scale size increases with age groups (FIGS.4, 5, 6).
In PCA analysis, when examined by age, PC1 accounts for 29.5%,
PC2 for 22.7%, and the rst ve components explain 85.4% of the
total variation. When examined by season, PC1 accounts for 32.9%,
PC2 for 19.9%, and the rst ve components explain 85.3% of the
total variation. When examined by sex, PC1 accounts for 36%, PC2
for 20.5%, and the rst ve components explain 86.2% of the total
variation (FIGS. 7, 8, 9).
When looking at the CVA results, the 6–year age group differs
signicantly from all other groups except the 7–year–old group,
while there is no signicant difference between the 3–4 and 4–5 age
groups and among other groups (TABLE III; FIG. 10). When examining
the seasonal groups, there is no signicant difference between the
Summer and Spring groups, while the difference between Autumn–
Summer and Autumn–Spring is signicant (TABLE IV; FIG. 11). There
is no signicant difference between sex (F–M) (TABLE V; FIG.12).
FIGURE 7. Scatter plot of principal component analysis (PCA) showing the
distribution of scales by age. Number represent ages
FIGURE 10. CVA plot of scales by age. Number represent ages
FIGURE 11. CVA plot of scales by Season. Au: autumn, Sm: summer, Sp: spring
FIGURE 12. CVA plot of scales by sex. F: female, M: male
FIGURE 8. Scatter plot of principal component analysis (PCA) showing the
distribution of scales by season. Au: autumn, Sm: summer, Sp: spring
FIGURE 9. Scatter plot of principal component analysis (PCA) showing the
distribution of scales by sex. F: female, M: male
Variation in scale characteristics of Acanthobrama marmid / Bilici et al. ____________________________________________________________
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TABLE III
CVA result of scales by age
Age
2 3 4 5 6
M.Dist /
P–value
P.Dist /
P–value
M.Dist /
P–value
P.Dist /
P–value
M.Dist /
P–value
P.Dist /
P–value
M.Dist /
P–value
P.Dist /
P–value
M.Dist /
P–value
P.Dist/
P–value
3
1,5243 /
0,0497
0,0471 /
0,1212
4
1,8217 /
0,0344
0,0728 /
0,0133
0,9992 /
0,1436
0,0381 /
0,0511
5
2,1384 /
0,0028
0,0752 /
0,0138
1,3902 /
0,0084
0,0408 /
0,0633
1,0575 /
0,3308
0,0213 /
0,8375
6
3,2478 /
0,0056
0,1391 /
0,0080
2,8103 /
0,0001
0,1132 /
0,0001
2,1309 /
0,0204
0,0813 /
0,0067
2,1597 /
0,0358
0,0810 /
0,0125
7
2,5964 /
0,3006
0,0818 /
0,3589
2,5553 /
0,1412
0,0687 /
0,3034
2,1714 /
0,5779
0,0526 /
0,8124
1,9358 /
0,7225
0,0559 /
0,7136
2,7091 /
0,5497
0,0866 /
0,3733
M.Dist: Mahalanobis distance, P.Dist: Procrustes distance,
P–value: value of permutation test
TABLE IV
CVA result of scales by season
Autumn Summer
M.Dist / P–value P.Dist / P–value M.Dist / P–value P.Dist / P–value
Summer 2.4640 / <
0.0001 0.0877 / 0.0001
Spring 2.2869 / < 0.0001 0.0768 / 0.0001 0.6419 / 0.6112 0.0213 / 0.4966
M.Dist: Mahalanobis distance, P.Dist: Procrustes distance,
P–value: value of permutation test
TABLE V
CVA Result of scales by gender
Female
M.Dist / P–value P.Dist / P–value
Male 0,7875 / 0,1833 0,0209 / 0,4603
M.Dist: Mahalanobis distance, P.Dist: Procrustes distance,
P–value: value of permutation test
FIGURE 13. Shape dierences of scales by age. Numbers represent ages
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Upon reviewing the DFA results, signicant differences were found
between age groups 2–4, 5–6, 3–4, 5–6, and 4–6, while sucient
differences were not observed among other age groups (TABLE VI;
FIG. 13). Signicant differences were found between the seasonal
TABLE VI
DFA results f of scales by age
Age
2 3 4 5 6
3
T
2
19,4271
Param. P 0,1293
Perm. P (Proc./T
2
) 0,1200 / 0,1300
4
T
2
18,6414 14,1287
Param. P 0,1456 0,1960
Perm. P (Proc./T
2
) 0,0140 / 0,1500 0,0600 / 0,1820
5
T
2
46,2855 26,5816 9,7766
Param. P
0,0170 0,0246 0,4679
Perm. P (Proc./T
2
) 0,0140 / 0,0170 0,0650 / 0,0210 0,8200 / 0,4600
6
T
2
101,0029 43,3982 28,1028 16,1055
Param. P 0,1505 0,0065 0,0438 0,3656
Perm. P (Proc./T
2
) 0,0070 / 0,1550 < 0.0001 / 0,0060 0,0090 / 0,0440 0,0140 / 0,3740
7
T
2
37,6469 20,6773 9,6780 9,0769 8,4726
Param. P 0,7085 0,1513 0,5808 0,7534 0,8535
Perm. P (Proc./T
2
) 0,3530 / 0,6840 0,3060 / 0,1610 0,8400 / 0,6250 0,7250 / 0,7140 0,3660 / 0,4090
T
2
: T–square, Param. P: Parametric P–values, Perm. P: Permutation P–value, Bolded: signicant
groups Autumn–Summer and Spring, but there was not enough
difference observed between Summer and Spring (TABLE VII;
FIG.14). No signicant differences were observed between sexes
(TABLEVIII; FIG. 15).
FIGURE 14. Shape dierences of scales by season. Au: autumn, Sp: Spring, Sm: summer
FIGURE 15. Shape dierences of scales by sex F: female, M: male
Variation in scale characteristics of Acanthobrama marmid / Bilici et al. ____________________________________________________________
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TABLE VIII
DFA result of scales by sex
Female
Male
T
2
11,2785
Param. p 0,2719
Perm. P (Proc./T
2
) 0,4550 / 0,2470
T
2
: T–square, Param. P: Parametric P–values, Perm. P: Permutation P–value
can be signicant based on the physicochemical parameters of the
environment and feeding [34]. In this sense, changes in the shape
of sh scales can allow for differentiation in populations [35, 36].
Additionally, inter/intraspecic morphological variability may indicate
genetic differences among samples or can respond to environmental
conditions within the framework of phenotypic plasticity [37, 38].
Geometric morphometrics is important in sh scales studies
because it allows for the quantitative analysis of shape and size
variation in a way that traditional morphometrics cannot achieve
[38, 39]. This method provides a detailed and comprehensive
understanding of the shape and size changes in sh scales, which
can be used to address questions related to taxonomy, evolution,
and ecology. Additionally, geometric morphometrics allows for the
visualization and analysis of complex patterns of shape variation,
making it a valuable tool for researchers studying sh scales [12,
40]. Çiçek et al. [41] applied geometric morphometric methods
successfully on Capoeta trutta and Capoeta umbla species. In the
present study, it was achieved on Acanthobrama marmid species
at the same success. In the size analysis performed according to
sex, it was seen that female samples were larger than males. These
results show that sh species can be successfully distinguished by
morphometric geometric analysis.
This type of analysis has been used successfully in previous studies.
For example, studies on sh scale and otolith morphometry and
geometry [13, 34, 41, 42, 43, 44, 45, 46] have yielded important results
in this eld. In addition, studies examining the relationship between
sh size and otolith morphometry [47, 48] were also effective in
determining the species.
CONCLUSIONS
Geometric morphometric analyses are highly accurate in species
discrimination and detecting diversity, offering a significant
advantage in future studies due to their ease, effectiveness, low
cost, reliability, and simplicity. Fish scales are essential for species
identication, making geometric morphometric analysis a vital tool
in future biological research. Procrustes ANOVA showed a signicant
size difference between sexes but no difference in shape. Signicant
variations in both size and shape were also found among groups based
on season and age. Additionally, PCA, CVA, and DFA analyses revealed
distinct patterns in scale size and signicant differences within age
and seasonal groups, but not between sexes.
Conict of interest
The authors have no declaration of competing interests.
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