True lemmings (Lemmini, Arvicolinae, Rodentia) in the Eopleistocene (Calabrian) in the North of western Siberia

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Resumo

Based on zoological collections and published information, morphological characteristics of the teeth and mandibles of Lemmus cf. sibiricus from the Eopleistocene Skorodumian faunal complex of western Siberia are compared with those of modern and fossil Lemmini from Europe and Asia so as to clarify the taxonomic identity and determine the place of the fossil form under consideration in the evolutionary history of the Palaearctic Lemmini. Comparative morphological approaches based on qualitative traits and linear measurements of cheek teeth, traditionally used for taxonomic diagnostics of representatives of the tribe, are used. according to the combination of apomorphic and plesiomorphic features, lemmings from the Skorodumian faunal complex of the Eopleistocene of western Siberia are shown to occupy a position intermediate between the more ancient L. kowalskii and the modern species Lemmus sibiricus. According to this intermediate position between the ancestral widespread form and the modern regional taxa of the genus Lemmus, the West Siberian Eopleistocene lemmings correspond to the East Siberian Lemmus sheri Abramson, 1992. The differences between L. kowalskii and the Eopleistocene forms are clearly recognizable and reflect the evolutionary changes in the cheek teeth associated with the transition of the Eopleistocene forms to a higher level of specialization in bryophagy. At the same time, the morphological similarity of the western and eastern Siberian Eopleistocene lemmings to each other is no higher than that of each of the regional forms to modern Lemmus inhabiting the same regions. Based on this, within the Eurasian range of the Eopleistocene Lemmus, differentiation comparable to the intrageneric differentiation within modern Lemmus could presumably have been observed. The combination of similarities in some features and differences in others most likely reflects mosaic morphological variations across the Eopleistocene range of one form, possibly a polytypic taxon, which by right of priority should be termed as Lemmus sheri Abramson, 1992. Taking into account the presence of regional features, it seems appropriate to consider the Eopleistocene lemmings of the Olerian Formation of eastern Siberia as L. sheri sensu stricto, and the lemmings from the Skorodumian faunal complex of western Siberia as the polytypic taxon L. sheri sensu lato.

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Sobre autores

E. Markova

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

Autor responsável pela correspondência
Email: emrk@yandex.ru
Rússia, st. March 8, 202, Yekaterinburg, 620144

A. Borodin

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

Email: bor@ipae.uran.ru
Rússia, st. March 8, 202, Yekaterinburg, 620144; prospect Lenina, 51, Yekaterinburg, 620075

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2. Fig. 1. Scheme of evaluation of characters differentiating L. kowalskii from modern Palaearctic Lemmus and Myopus by the structure of M3. 1 – Orientation variants of M3 lophs: 1a – upper (mesial) lophs are oriented at an angle to the lower (distal) ones; 1b – lophs are conditionally parallel, i.e. four parallel lines can be drawn through them, and there are no structures that violate the lophodont plan; 1c – lophs are conditionally parallel and an unpaired distal prism is present. Straight arrows – lophs, crossed arrows – unpaired distal prism (= heel of M3), round frame – place of junction of AL and T2 (T2 has the shape of a triangle) in L. kowalskii with plesiomorphic orientation variant of lophs 1a. 2 – Asymmetry of reentrant angles of M3, estimated by the values of the IAs index. The arrows show the scheme of measuring the reentrant angles – lingual LRA2 and buccal BRA2. Variants of manifestation of features: 1a, 2a – plesiomorphic; 1b, 1c, 2b – apomorphic.

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3. Fig. 2. Mean and median values of the indices of the width of the chewing surface, IW (W/L*100), and the asymmetry of the reentrant angles, IAs (LRA2/BRA2*100) of the third upper tooth in modern and fossil Lemmini of the Palaearctic (1 – Tobienia kretzoi, N = 3 (Fejfar, Repenning, 1998); 2 – Plioctomys mimomiformis, N = 3 (Rebielice Królewskie, Kowalski, 1977); 3 – Lemmus kowalskii, N = 9 (Shernfeld (Carls, Rabeder, 1988) and Zamkowa Dolna (Kowalski, 1977)); 4 – Lemmus sheri, Krestovka, N = 21 (Abramson, 1992); 5 – Skorodum and Romanovo, N = 28; 6 – Lemmus sibiricus, N = 32 (Western Siberia, modern); 7 – Myopus schisticolor, N = 30 (Northern Cis-Urals, modern)).

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4. Fig. 3. Comparison of samples from the Skorodum and Romanovo localities (data combined) with the type series of Sher's lemming and modern Lemmus sibiricus (Western Siberia) and Myopus schisticolor (Northern Cis-Urals) by the values of the index of relative width of the chewing surface M3, IW.

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5. Fig. 4. Examples of M3 of modern and fossil Lemmini with different values of the index of the relative width of the chewing surface, IW (W/L*100): black asterisks indicate index values in the range of 46–50 (typical for Lemmus), red asterisks indicate 58–60 (typical for Myopus), not marked with an asterisk indicates a range of intermediate index values (56–58). Modern: 1 – Myopus schisticolor, Northern Cis-Urals, id 1929`17; 2 – M. schisticolor, Northern Cis-Urals, id 1746`17; 6 – Lemmus sibiricus, Tyumen Region, IERiZh 19981; 7 – Lemmus sibiricus, Tyumen region, IERiZh 24522. Fossils: 3, 4, 8 – Romanovo 1s; 9 – Skorodum a; 5, 10 – L. sheri, type series, Krestovka, KLO-6. Scale bar – 1 mm.

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6. Fig. 5. Values of the discriminant function Y6 calculated from seven measurements of m3 from collection materials from the Skorodum locality (N = 13) and the type series of L. sheri (N = 11) in comparison with modern Lemmus and Myopus (frequencies for modern taxa according to: Chaline et al., 1988).

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7. Fig. 6. Examples of m3 from the Skorodum locality, for which the calculated values of the discriminant function Y6 are minimal (1) and maximal (2) for the studied sample (for comparison, m3 of L. sheri, corresponding to the proportions of Myopus (3) and Lemmus (4). 1 – id 1, Skorodum a`, Y6 = –12.6; 2 – Skorodum a, id 12, Y6=5.5; 3 – id 700, Krestovka, KLO-6, holotype of L. sheri; 4 – Krestovka, KLO-10, outcrop 6. Scale 1 mm.

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8. Fig. 7. Comparison of Pliocene, Pleistocene and modern Lemmini by complexity ranks of the third upper and third lower molars: 1 – modal class of distribution of complexity ranks of the basal genus Tobienia, Pliocene (based on a single find: Tesakov, Bondarev, 2021); 2 – Plioctomys, Pliocene (Sukhov, 1977; Kowalski, 1977); 3 – Lemmus kowalskii, Paleopleistocene = Gelasian (Kowalski, 1977; Carls, Rabeder, 1988)); 4 – Lemmus sheri, Eopleistocene = Calabrian (Abramson, 1992); 5a – Romanovo 1, Eopleistocene; 5b – Skorodum, Eopleistocene; 6 – modern Lemmus sibiricus (Markova et al., 2018): a – r. Kharasavey, Yamal Peninsula, b – Vorkuta, Polar Cis-Urals, c – r. Seyakha, Yamal Peninsula); 7 – modern Myopus schisticolor (Markova et al., 2018): a – Northern Cis-Urals, b – Sorumsky Reserve, Western Siberia, c – Sibirskie Uvaly, Western Siberia).

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9. Fig. 8. Lower jaws of lemmings from the Skorodum locality a: 1–3 – adult individual, view of the jaw from above (1), from the lingual side (2), and from the buccal side (3); 4–5 – young individual, view of the jaw from above (4) and from the lingual side (5). Scale bar 5 mm.

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10. Fig. 9. Visualization of the similarity of morphological features of the combined sample of Eopleistocene lemmings from the Skorodum and Romanovo localities (red rectangle) with modern genera of Palaearctic Lemmini (Lemmus, Myopus) and fossil Quaternary lemmings of Eurasia – L. sheri and L. kowalskii. Arrows are comparison options, numbers are numbers of features by which the considered forms are similar in pairwise comparisons: they have common variations (features 1, 5c, 7), do not differ from each other by the established boundary values (features 2, 6, 7) or do not demonstrate statistically significant differences (features 3–4, 5a, 5b). Drawings of tooth contours after: Smirnov et al., 1986; Carls, Rabeder, 1988; Abramson, 1992; Markova et al., 2018, preserving the style of the original source. Features: 1b – regular lophodont M3 with parallel lophs without additional prisms, 1c – lophodont M3 with subparallel lophs and a massive heel; 2 – minimum values of the IAS index (LRA2/BRA2*100) less than 80; 3 – values of the IW index (W/L*100) do not differ statistically; 4 – values of the discriminant function calculated by 7 measurements of m3 do not differ statistically; 5a, 5b – frequencies of complexity ranks of m3 (5a) and M3 (5b) do not differ statistically, 5c – simplified m1 are present (m1_rank-1); 6 – sizes of m1 in adults are on average more than 3 mm; 7 – the alveolar apex of the lower incisor is located at the level of the second loop of m3 in at least some adults.

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