International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.

1, March 2025

IMPACT OF LATITUDE ON SEXUAL SIZE

DIMORPHISM IN GROUND BEETLE
POECILUS CUPREUS L
Raisa Sukhodolskaya 1,2,3, Teodora Teofilova 4, Vladimir Langraf 5,
Kirill Maximovich 6, Tamara Avtaeva 7,8, Nadezhda Ukhova 9
1
Institute of Ecology and Subsoil Use of Tatarstan Academy of Sciences,
Kazan, Russian Federation
2
Kazan State Medical University, Kazan, Russian Federation
3
Volga-Kama State Reserve, Republic of Tatarstan, Zelenodolsk Disctrict,
Russian Federation
4
Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of
Sciences, Sofia, Bulgaria
5
Constantine the Philosopher University, Nitra, Slovakia
6
Siberian Federal Scientific Centre of Agro-BioTechnologies of the Russian
Academy of Sciences, Krasnoobsk, Russian Federation
7
Kh. Ibragimov Complex Institute of the Russian Academy of Sciences,
Grozniy, Russian Federation Chechen State Pedagogical University,
Grozniy, Russian Federation , Visim State Reserve, Kirovograd, Russian
Federation

ABSTRACT
Sexual dimorphism is a common phenomenon in insects, and it may be expressed in various
traits including body size and body shape. Sexual selection is the main factor shaping such
differences between males and females in insects, even if they depend also on food
availability during larval stages. Body size can differ geographically between particular
populations of one animal species, both with latitude and altitude, and it was undoubtedly
shown for ground beetles. The latter are considered excellentbioindicators and model
organisms for evolutionary studies. We have investigated ground beetle Pocilus cupreus
body size variation practically throughout its entire range and have found that the curve of
its body size variation was saw-tooth in latitude gradient and in longitude as well. Using
that base set we estimates sexual size dimorphism (SSD) variation in geographical
gradients.We have measured more than 8000 specimen of males and females in ground
beetle Poecilus cupreus L. sampled in 14 large regions of Eurasia. Sexual Size
Dimorphism changed drastically in latitude gradient not showing any trend to decrease or
to increase. We conclude that several dominating factors affected SSD presentation in
species studied, among which climatic factors at the sites where beetles have been sampled.

KEYWORDS
Sexual Size Dimorphism, Ground Beetles, Latitude Gradient, Body Size Variation, Sexual
Selection, Environmental Factors, Climate Change.

DOI: 10.5121/ijbb.2025.15102 13
 International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.1, March 2025

1. INTRODUCTION
The patterns and causes of body size variation in geographical gradients are fundamental themes
in studies on life-history evolution [1]. Their importance is high because climate changes and
trends in space may be highly relevant for predictions of changes over time [2]. Body of
publications in this theme grows but the diversity of ecogeographic body size clines remains not
fully understood, particularly in ectotherms [3, 4].

The most known are latitudinal and altitudinal clines in body size [5]. Temperature is often
assumed to be the principal determinant of ecogeographic body size clines, because temperature
covaries consistently with latitude and altitude, and it strongly affects vital processes in the
organisms [6]. Yet, a number of recent studies have found that water availability (precipitation,
humidity), and particularly seasonality (within-year variation in temperature or precipitation),
often explain a higher proportion of body size variation than does mean temperature [7, 8]. For
each of these factors, multiple mechanistic hypotheses have been proposed (see below).
However, rigorous testing of such hypotheses is often impeded by collinearity between climatic
variables [9] which is particularly common within limited geographic areas.

No less significant in studies of body size is the phenomenon of differences observed between
females and males, the so-called sexual-size dimorphism (SSD). A lot of research has been done
to elucidate the evolutionary processes that underlie sexual dimorphism patterns betweenanimals
[10]. The direction and extent of the sexual differences in body shape and size in many insect
species are greatly influencedby sexual dimorphism and particular developmental processes [11].
Beetles (Coleoptera) are cosmopolitan in distribution andexhibit great sexual dimorphism among
different species.The evolutionary foundation for the study of sex differences is provided by
Darwin’s sexual selection theory, whichincludes intrasexual competition for mates and
discriminative mating partner choice (intersexual choice). Although nichedifferentiation between
sexes through ecological character displacement can cause sexual dimorphism, its development
hastraditionally been attributed mostly to differences in the strength of sexual selection acting on
males and females [12].Competition and selection dynamics not only result in the evolution of
sex variations in trait size or degree of elaboration,but they also make these traits more sensitive
to social and ecological pressures.

2. MATERIAL AND METHODS

2.1. Sampling Regions

The material was obtained under scientific cooperation agreements from researchers from fifteen
regions of Russia and abroad. Ground beetles were caught using the standard Barber trap method.
The sample from each region included beetles caught in a range of sites that differed in the
degree of anthropogenic impact and vegetation: cities, suburbs, agrocenoses, and natural
biotopes. The number of individuals from each raft was at least 35 individuals (Table 1).

14
 International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.1, March 2025
Table 1. Description of sampling localities of P. cupreus

Region Latitude, ◦N Longitude, Number of Type of Sample

◦
E Sites Habitats Size
1 Switzerland 47,0016° N 8,0013° E 1 rape 148
2 Germany 51,0916° N 9,0013° E 4 rape 569
3 Bulgary 42.3243° N 25.0232° E 3 rape 698
4 Romania 44.2518° N 26.3703° E 9 rape 1174
5 Krasnodar 44.3833° N 38.9667° E 1 rape 154
Territory
6 Kostroma 57,4606° N 40,5559° E 1 meadow 100
Oblast
7 Stavropol 48,46° N 35,04° E 1 meadow 71
Territory
8 Chechen 43,1719° N 45,4534° E 1 arable 187
Republic
9 Republic of 55,4731° N 49,0720° E 9 barley, pea, 2643
Tatarstan spring wheat,
rye, vetch & oat,
alfalfa, carrot,
lawn
10 Sverdlovsk 55,1719° N 68,4534° E 1 meadow 200
Oblast
11 Kemerovo 55,2520° N 86,70° E 1 meadow, lawn 67
Oblast
12 Novosibirsk 55,27° N 79,33° E 2 meadow 548
Oblast
13 Slovakia 45,02° N 41,55° E 1 meadows, 146
pasture
14 Ukraine 48,77° N 35,08° E 3 arable lands 145

2.2. Study Organism

P. cupreus is a widespread Palaearctic species 8.5-13.5 mm in length. The humeral tooth of the
elytra is indistinct. The head is densely dotted. The elytra are somewhat wider than the base of
the pronotum. The upper surface is copper-red, bronze, green, black, blue, etc.; sometimes the
femora, rarely the legs, are red. The coloration is very diverse, sometimes two-colored, the legs
are always black. This is a meadowfield species with a high population in agrocenoses. It is of
great importance as an entomophage. It finds favorable conditions for living in urban areas also
and has a wide range of requirements for the habitat.

2.3. Study Design

Individuals were photographed 5-6 specimen at a time and the parameters were measured on a
self-written program in Python 2.7 using the numpy and openCV libraries. The program is
necessary to achieve greater measurement accuracy, automate the process and reduce
subjectivity. They are measured according to six dimensional traits (Fig. 1.). A total of 6,909
insect specimens were measured, with the last column indicating the number of specimens
measured in each region (Table 1). The data are collected in Excel tables for subsequent
statistical analysis.

15
 International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.1, March 2025

Figure 1. The scheme of beetles measurements. A –elytra length, B – elytra width, C – pronotum length, D
– pronotum weight, E – hat length, F – distance between eyes.

2.4. Statistical Analysus

Statistical processing was carried out in the Excel program. Methods of onedimensional
statistical analysis were used. The mean, standard deviation, error of the mean, and the variation
coefficient were calculated. The variation coefficient is calculated using the formula standard
deviation/mean*100. Graphs were constructed based on the data obtained, and the results were
entered into a table. SSD was calculated according the formula: mean trait in females/mean trait
in males -1. Thus, if calculated value of SSD was positive, females in the sample studies were
larger than males. And vice versa: when SSD value was negative, then we thought that the
conditions of that region were unfavorable, or some microevolutionary changes were taking place
in that region, since negative SSD values (when females are smaller than males) are not typical
for ground beetles

3. RESULTS AND DISCUSSION

The SSD values changed significantly in the latitudinal gradient (Fig. 2): they became more or
less stable closer to the northern latitudes - this is the northern boundary of the range of the
studied species. In the south, the SSD values changed abruptly when moving from one region to
another, acquiring negative values in the Krasnodar Territory and in Ukraine. The latter
phenomenon (when females are smaller than males) is not typical for ground beetles in general.
This fact supports the opinion that such environmental conditions developed there. Considering
that P. cupreus is a field species, this may well be explained by some agricultural practices that
were used in the places of capture. However, this is not the only explanation. SSD studies have
shown that its value can be determined by a number of factors: mean summer temperature,
summer precipitation, seasonality, and reproductive mode/lineage identity on female size, male
size, and SSD. For example the investigators found a moderate effect of reproductive mode and
precipitation and a strong but complex effect of seasonality [13]. The latter differed drastically
16
 International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.1, March 2025
between the lineages, being also modulated by precipitation and especially by temperature.
Female size and SSD varied stronger than male size, and virtually all the effects were strongly
female-biased. That study operated with lizards.

Studies in ground beetles SSD variation are scarce. So, it was shown, that SSD in them also
varied. In different species its value fluctuated between 0.02 and 0.12, and according to various
characteristics those values varied even more [14]. Moreover, intra-specific variation of SSD
values was great. Latitudinal variability of SSD in six species of ground beetles was studied and
the authors concluded that despite SSD occurred in some analyzed latitudes and mainly at the
margins of area, modeling results did not confirm opinion than SSD systematically changes in
latitude gradient [15, 16]. Moreover, sometimes SSD varied according to the type of biotope [17].

Figure 1. Variation of SSD in P. cupreus in different points. 1) Bulgaria 2) Chechen Republic 3) Romania
4) Krasnodar Province 5) Ukraine 6) Stavropole Province 7) Switzerland 8) Slovakia 9) Germany 10)
Kostroma Oblast 11) Kemerovo Oblast 12) Novosibirsk Oblast 13) Tatarstan Republic 14) Sverdlovsk
Oblast

4. CONCLUSIONS
Despite the relevance of the topic, the problem of SSD is far from being resolved. To solve it, it
is necessary to comply with a lot of conditions, both when collecting material and when choosing
an object and research methods. Similar to the conditions for conducting experiments to assess
the variation of animal sizes in environmental gradients, when studying SSD, it is necessary to
keep in mind that the object must be massive, well studied in biological and ecological terms, and
a unified measurement method must be used. In our study, the SSD value changed in the same
saw-tooth pattern as shown in earlier studies on size variability in this species of ground beetles.
However, the approximation showed a weak trend toward an increase in the SSD value toward
the north. At the same time, in ground beetles of other genera, the SSD values either did not
change in the latitudinal gradient or were tied to the population size in a given region. This
suggests that the expression of SSD may be genus-specific, which should be taken into account in
further studies.

17
 International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.1, March 2025

ACKNOWLEDGEMENTS
The research is funded by the State Task Theme 730000P.16.1.OH17AA106000 “Biological
diversity of Eastern Europe under the influence of natural and climatic factors in historical
and modern contexts”.

REFERENCES

[1] Arendt, J., & Fairbairn, D. J. (2012). Reproductive allometry does not explain the temperaturesize
rule in water striders (Aquarius remigis) Evolutionary Ecology, Vol. 26, pp745–757.
https://doi.org/10.1007/s10682-011-9524-4
[2] Teplitsky, C., & Millien, V. (2014). Climate warming and Bergmann's rule through time: is there
any evidence? Evolutionary Applications, Vol. 7, pp156–168. https://doi.org/10.1111/eva.12129
[3] Hjernquist, M. B., Söderman, F., Jönsson, K. I., Herczeg, G., Laurila, A., &Merilä, J. (2012).
Seasonality determines patterns of growth and age structure over a geographic gradient in an
ectothermic vertebrate Oecologia, Vol.170, pp641–649. https://doi.org/10.1007/s00442-012-23384
[4] Rypel, A. L. (2014). Cold-water connection: Bergmann's rule in North American freshwater fishes
The American Naturalist,Vol.183, pp147–156.https://doi.org/10.1086/674094
[5] Blanckenhorn, W. U., & Demont, M. (2004). Bergmann and converse Bergmann latitudinal clines
in arthropods: two ends of a continuum? Integrative and Comparative Biology V. 44, pp413–424.
https://doi.org/10.1093/icb/44.6.413
[6] Angilletta, M. J. (2009). Thermal adaptation: a theoretical and empirical synthesis. Oxford:Oxford
University press. https://doi.org/10.1093/acpro f:oso/97801 98570 875.001.1
[7] Çağlar, S. S., Karacaoğlu, Ç., Kuyucu, A. C., & Sağlam, İ. K. (2014) Humidity and seasonality
drives body size patterns in males of the bush cricket Isophya rizeensis Sevgili, 2003 (Orthoptera:
Tettigoniidae: Phaneropterinae) Insect Science V. 21, pp213–226.
https://doi.org/10.1111/17447917.12027
[8] Stillwell, R. C., Morse, G. E., & Fox, C. W. (2007). Geographic variation in body size and sexual
size dimorphism of a seed-feeding beetle The American Naturalist V. 170, pp358–369.
https://doi.org/10.1086/520118
[9] Millien, V., Kathleen, L. S., Olson, L., Smith, F. A., Wilson, A. B., & Yom-Tov, Y. (2006).
Ecotypic variation in the context of global climate change: revisiting the rules. Ecology Letters Vol.
9, pp853–869. https://doi.org/10.1111/j.1461-0248.2006.00928.x
[10] Benítez, Hugo, Sukhodolskaya, Raisa, Órdenes-Clavería, Rodrigo, Avtaeva, Tamara, Kushalieva,
Shapaat, Saveliev, Anatoliy (2020).Measuring the Inter and Intraspecific Sexual Shape Dimorphism
and Body Shape Variation in generalist Ground Beetles in Russia Insects V.11(6), pp1–13.
https://doi.org/10.3390/insects11060361.
[11] Budečević, S, Savković, U, Đorđević, M, Vlajnić, L, Stojković, B. (2021) Sexual Dimorphism and
Morphological Modularity in Acanthoscelides obtectus (Say, 1831) (Coleoptera: Chrysomelidae): A
Geometric Morphometric Approach Insects V.12(4), pp1–12.
https://doi.org/10.3390/insects12040350.
[12] Baur, J, Roy, J, Schäfer, M.A, Puniamoorthy, N, Blanckenhorn, W.U, Rohner, P.T. (2020)
Intraspecific mating system evolution and its effect on complex male secondary sexual traits: Does
male–male competition increase selection on size or shape? Journal of Evolutionary Biology V.
33(3), pp297–308. https://doi.org/10.1111/jeb.13565.
[13] Roitberg, Evgeny S., Orlova Valentina F., Bulakhova, Nina A., Kuranova, Valentina N.,| Eplanova,
Galina V., Zinenko, Oleksandr I., Arribas, Oscar, Kratochvíl, Lukáš, Ljubisavljević, Katarina,
Starikov, Vladimir P., Strijbosch, Henk, Hofmann, Sylvia, Leontyeva, Olga A., Böhme, Wolfgang
(2020) Variation in body size and sexual size dimorphism in the most widely ranging lizard: testing
the effects of reproductive mode and climate Ecology and Evolution V.10, pp4531–4561.
[14] Sukhodolskaya, Raisa, Saveliev,A., Muhammetnabiev T. (2016) Sexual Dimorphism of Insects and
Conditions of Its Manifestation Research Journal of Pharmaceutical, Biological and Chemical
Sciences V. 7(2), pp1992 – 2001.
[15] Sukhodolskaya, Raisa & Saveliev, Anatoliy (2017) Geographical variation of sexual size
dimorphism in ground beetle Pterostichus melanarius (Coleoptera, Carabidae) Proceedings of 18
European Carabidologists Meeting. Rennes, France pp 61.

18
 International Journal on Bioinformatics & Biosciences (IJBB) Vol 15, No.1, March 2025
[16] Sukhodolskaya, Raisa, Saveliev, Anatoliy, Gordienko, Tatiana, Vavilov, Dmitriy (2018) Sexual size
dimorphism in Ground Beetles and its modeling in latitude gradient GSC Biological and
Pharmaceutical Sciences V. 3(1), pp 11-18.
[17] Sukhodolskaya R. A., Avtaeva T. A, Saveliev A. A., Vavilov D. N. 2019. Sexual Size Dimorphism
in Ground Beetle Carabus cumanus Fischer von Waldheim, 1823 (Coleoptera, Carabidae) and its
Variation in Different Traits. Baltic J. Coleopterol., 19(1): 89 - 100.

AUTHOR

Raisa Sukhodolskaya, Senior Researcher Ph.D in Biology , Institute of Ecology and

Subsoil Use of Tatarstan Academy of Sciences, Russia

19