Dental enamel differentiation quotient of modern water voles of the genus Arvicola in Azerbaijan: variation and diagnostic value

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Abstract

Based on a study of dental characteristics of the water vole genus Arvicola from 7 regions of Azerbaijan, captured in 1926–1965 and stored in the collection of the Institute of Zoology, Ministry of Science and Education of Azerbaijan, the range of variability of the tooth enamel differentiation index SDQ is determined. The index is calculated as the ratio of the thickness of leading to trailing cutting edges of the enamel prisms of the cheek tooth crowns. SDQ serves as a criterion for identifying chronospecies of Arvicola in the fossil record, and it has recently been proposed for diagnosing modern Eurasian and Persian water voles. As a result of measuring the SDQ index in 55 individuals from the Shemakha, Saatli, Imishli, Goygol (= Khanlar), Ordubad, and Adjikabul regions, the water voles are shown to have a plesiomorphic type of enamel differentiation, in which the SDQ index values are greater than 100. In the specimens under consideration, the values of SDQ indices assessed for the first lower and second upper teeth vary from 100 to 178. In contrast to the dimensional characteristics of the teeth, the index values do not demonstrate directional changes in postnatal development and retain the plesiomorphic pattern of dental enamel differentiation in all age classes distinguished by the degree of skull maturity. The results obtained confirm the difference in the dental characteristics of the water voles occurring in Azerbaijan from the Eurasian A. amphibius and the proximity to the west Asian water voles currently considered as an independent species or species group, A. persicus (Persian water vole). Revision of data on differential chromosome staining and an analysis of molecular markers, both mitochondrial and nuclear, are necessary to answer the question concerning the taxonomic status of water voles in Azerbaijan and other regions of Western Asia, where water voles with a negative type of differentiation of tooth enamel occur. Until a final decision is made on the taxonomic status, composition and distribution of the Persian water vole, the water voles of Azerbaijan with a plesiomorphic type of tooth enamel differentiation are proposed to be referred to as Arvicola cf. persicus.

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About the authors

E. A. Markova

Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences

Email: emrk@yandex.ru
Russian Federation, st. March 8, 202, Yekaterinburg, 620144

L. V. Nadirli

Institute of Zoology, Ministry of Science and Education, Republic of Azerbaijan; Western Caspian University

Author for correspondence.
Email: leyla_h-va@list.ru
Azerbaijan, st. A. Abbaszade 1128, Baku, AZ1004 Azerbaijan; st. Istiglaliyyat, Baku, AZ1001 Azerbaijan

S. V. Zykov

1 Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences

Email: svzykov@yandex.ru
Russian Federation, st. March 8, 202, Yekaterinburg, 620144

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2. Fig. 1. Nomenclature of elements of the first lower and second upper cheek teeth and the scheme for assessing dental features. A – Designations of tooth prisms (according to: Van der Meulen, 1973), metric features of the chewing surface and areas 1–2, where the enamel thickness was additionally assessed taking into account its microstructure using an electron scanning microscope; 1 – anterior wall, 2 – posterior wall, scale 20 μm. B – Scheme for measuring the enamel differentiation index SDQ on one prism (using T3 as an example), C – Locations of enamel thickness measurement for calculating indices m1_SDQ7, m1_SDQ3, M2_SDQ3. D – Types of chewing surface wear using one tooth prism as an example: left – lunate type, right – stepped type. D – formulas for enamel differentiation indices on the first lower and second upper cheek teeth. Abbreviations – see text.

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3. Fig. 2. Scheme of assessment of maturity of cranial structures of non-rhizomedentary Arvicolinae according to characters I–IX, expressed for each character as a percentage of the total (sum) maturity of the skull (Larina, Lapshov, 1974, with changes). Designation of the skull bones on which the characters of cranial maturity are assessed: 1 – frontal, 2 – parietal, 3 – squamosal part of the temporal bone, 4 – petrous and mastoid parts of the temporal bone, 5 – occipital, 6 – interparietal.

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4. Fig. 3. Age-related changes in the skull of the water vole, using three individuals with different degrees of cranial maturity assessed as a percentage using the method of Larina and Lapshov (1974): side view of the skull (a), top view (b), and back view (c). 1 – ID 95, total skull maturity 20%, Nakhichevan, sex and date of capture unknown; 2 – ID 2, male, skull maturity 60%, Imishli district, Bahramtepe, March 1965; 3 – ID 8, male, skull maturity 100%, Saatli district, Lake Sary-Su. Scale 1 cm.

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5. Fig. 4. Chewing surface of cheek teeth of water vole with skull maturity of 20% (1,2) and 100–100+% (3–5). White arrows – sections m1, M1 and M3 of juvenile individuals that have not reached definitive outlines, red arrows – senile folding of enamel edges, black arrow – “mimomisis fold”, found singly and being a manifestation of individual variability. 1, 2 – id 95 and id 96, Nakhichevan, 20% skull maturity, sex and date of capture unknown; 3 – id 44/3, female, Goygol district, 100+% skull maturity, June 23, 1964 (* lower row of teeth, ** M3), 4 – id 2854 (899), Chukhur-Yurt, Shemakha district, sex unknown, October 11, 1926, 100% skull maturity, 5 – id 7, Lake Sary-Su, Saatly district, March 18, 1965, female, skull 100+%.

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6. Fig. 5. Mean values ​​and 95% confidence interval of the length and width of m1 and M2 and the length of the anteroconid m1 (m1_A) in age classes of the water vole from Azerbaijan, distinguished by the degree of maturity of the skull. The range of variation of the absolute values ​​of the length and width of m1 and M2 of the Eurasian water vole, known from literature data (Borodin, 2009), is shown in gray. Age classes (by skull maturity, %): 1 - 20–45, 2 - 50–95, 3 - 100–100+.

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7. Fig. 6. Mean values ​​and 95% confidence interval of the values ​​of three indices of differentiation of cheek tooth enamel in water voles from Azerbaijan by age classes (left) and by regions of capture for animals of age class 3 (right). The overlapping area of ​​the range of absolute values ​​of the SDQ indices of the Eurasian and Persian water voles, established from the literature data (Röttger, 1987; Fadeeva et al., 2021; Maul et al., 2021), is shown in gray. Age classes (by skull maturity, %): 1 - 20–45, 2 - 50–95, 3 - 100–100+; regions: 1 - southern slope of the Greater Caucasus, 2 - Kura-Araks Lowland, 3 - Lesser Caucasus and Transcaucasian Plateau.

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8. Fig. 7. Comparison of modern water vole samples from Azerbaijan with literature data (Königswald, Kolfschoten, 1996; Masini et al., 2020; Fadeeva et al., 2021) on modern species (including A. sapidus, A. amphibius and the controversial taxon A. italicus) and the Middle Pleistocene form A. mosbachensis based on the values ​​of the m1_SDQ7 and m1_A/L indices. Dots are the average values ​​of the two indices, lines are the range of variability of the m1_SDQ7 index. Age classes - see text, data by region - excluding the age of animals.

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9. Values ​​of enamel differentiation indices m1_SDQ7, m1_SDQ3, M2_SDQ3 of water vole from Azerbaijan depending on cranial maturity of individuals

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10. Variants of the structure of the posterior section of the lower jaw of the water vole from Azerbaijan in the age group 100+: id 44/3, female, Goygol district, 100+% skull maturity, June 23, 1964 (left) and id 7, Lake Sary-Su, Saatly district, March 18, 1965, female, skull 100+% (right)

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