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Eastern European rodent (Rodentia, mammalia) faunas from the Early–Middle Pleistocene transition
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Quaternary International 131 (2005) 71–77
Eastern European rodent (Rodentia, mammalia) faunas from the Early–Middle Pleistocene transition Anastasia K. Markova Institute of Geography RAS, Staromonetny 29, 109017 Moscow, Russian Federation Available online 11 September 2004
Abstract The rodent faunas from nine sites on the Russian Plain and Taman Peninsula corresponding to the Early–Middle Pleistocene transition have been analyzed. These localities are situated in the Dnieper, Don, Dniester and Danube drainage basins and in the northern regions of the Black and Azov Seas. Seven of the sites are associated with the upper part of Matuyama reversed polarity Chron and located between the beginning of Jaramillo event and the Brunhes/Matuyama boundary. One of the sites was recovered from the Chauda marine deposits of the Black Sea and referred to the very beginning of Brunhes Epoch. The species composition of the rodent fauna sequence has been established. The faunas referred to the Jaramillo event (1.1–1.0 Ma) contain the advanced Mimomys, steppe lemmings Prolagurus pannonicus and Lagurodon arankae, and Allophaiomys pliocaenicus. Faunas were recovered from Korotoyak (Ostrogozhsk suite), Roksolany, Zapadnye Kairy and Ushkalka localities. The more advanced rodent faunas of the Morozovian assemblage (Port-Katon site) include also the first representatives of Terricola. Karai–Dubina and Shamin localities have been found below the Brunhes/Matuyama boundary (0.85 Ma), including the first representatives of Microtus (Microtus ex gr. oeconomus and Microtus arvalinus). The ‘‘Pitymys’’ like voles—Microtus (Stenocranius) hintoni, steppe lemming P. pannonicus and yellow steppe lemming Eolagurus argyropuloi compose the core of these faunas. The Litvin locality, located in the deposits of the Chauda transgression of the Black Sea and corresponding to the very beginning of the Middle Pleistocene, contains a rodent fauna closely resembling the Shamin one. It yielded the remains of Mimomys savini, advanced A. pliocaenicus, P. pannonicus, Microtus (Stenocranius) hintoni, Microtus ex gr. oeconomus and Mcirotus arvalinus. Thus, numerous data permit elucidation of the history of rodent faunas of the Russian Plain during Early–Middle Pleistocene transitions and reveal the definitive stages in their evolution. r 2004 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction The Middle–Early Pleistocene transition is increasingly recognized as a time of significant biotic changes in the Northern Hemisphere. The visible climatic changes (progressive cooling) are reflected in vegetation and animal assemblages. The open landscapes and the steppe mammal communities became widely distributed on the southern part of the Russian Plain in the second part of the Early Pleistocene and the beginning of the Middle Pleistocene. The well-pronounced evolutional changes in rodent fauna have been identified in material from E-mail address: [email protected] (A.K. Markova).
Eastern Europe for this same time interval. These data have been confirmed by geological as well as geochronological studies. The principal stages in rodent evolution are distinguished by the first appearance (FAD) of new taxa. These data permit the reconstruction of the sequence of evolutional changes in rodent faunas, and the identification of characteristic faunal assemblages, from the beginning of the Jaramillo paleomagnetic event until the beginning of the Brunhes paleomagnetic Epoch. The stratigraphical position of the sites was ascertained by direct geological and also paleomagnetic studies. Such studies are important for elucidating not only rodent evolution, but also the stratigraphy of Eastern Europe and correlation with other regions.
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2. Materials Rodent faunas from nine localities on the Russian Plain dating to the Early–Middle Pleistocene transition have been studied. All the localities are located between 511N and 451N. Most of the localities are associated with alluvium deposits of the main rivers of Russian Plain. Karai-Dubina, Zapadnye Kairy and Ushkalka faunas have been found in terrace deposits of the lower Dnieper drainage basin; Nagornoe 1 fauna have been recovered in the lower Danube drainage basin; the Roksolany site is in alluvium deposits of the Dniester river (Markova, 1982, 1990a,b; Mikailesku and Markova, 1992); Korotoyak (Ostrogozhsk suite) and Shamin faunas have been recovered in Don-river alluvium (Iosifova et al., 1992; Markova, 1992). Port-Katon and Litvin localities have been revealed on the Taman Peninsula in marine-liman deposits (Markova, 1990b, 1998). The faunas of all of these sites were studied (Markova, 1982, 1990a,b, 1998a; Bylinsky et al., 1993). Dr. A. Agadjanian and Dr. N. Kazantseva analyzed firstly the small mammals from the Korotoyak (Ostrogozhsk suite) locality (Agadjanian and Kazantseva, 1994).
3. Terminology This paper uses the terminology for the parts of occlusial surface of vole molars introduced by van der
Meulen (1973, 1974): L—length of molar; W—width of molar; AC 2—anterior cap; ACC—anteroconid complex; PL—posterior lobe; T1–T5—dentile fields of occlusial surface of molars; BRA—buccal re-entrant angle, BSA—buccal salient angle; LRA—lingual reentrant angle; LSA—lingual salient angle; B—shortest distance between AC 2 and T4 and T5; C—shortest distance between BRA 3 and LRA 3 angles. The SDQ index is used, after Heinrich (1978), which shows the ratio between the enamel thickness on posterior and anterior parts of the salient angles of the tooth. This work uses the measurements of the enamel of the three central angles of m1 (LSA2, LSA3, BSA2) and terms the SDQ index as SDQ3.
4. Stages of rodent faunas 4.1. Faunas, approximately synchronous to the Jaramillo paleomagnetic event The earliest faunas of the discussed time interval have been found in fluvial deposits at the Roksolany site (lower Dniester basin). Small mammal remains have been collected during field works headed by Mikhailesku (Mikhailesku and Markova, 1992) and Dodonov (Dodonov et al., 1998). The fossiliferous layers have been recovered here below a thick loess-soil section where the Brunhes—Matuyama boundary and Jaramillo event were established (Dodonov et al., 1998) (Fig. 1). The bones were found in the channel alluvium of the
Fig. 1. Geological structure and paleomagnetic data of the sections with small mammal faunas: 1—soil; 2—loess; 3—loam; 4—clay; 5—sand; 6— limestone; 7—water level; 8—small mammal remains; 9—mollusks remains; 10—borrows of small mammals; 11—Brunhes/Matuyama boundary; 12—Jaramillo event.
ARTICLE IN PRESS 11.76–18.62–25.0 13.68–19.43–25.00 18.18–19.79–22.58 12.50–19.88–22.50 18.95–23.54–31.25 26.88–28.23–29.41 40.0–45.33–46.0 37.50–40.48–42.85 41.22–41.57–41.92 0.80–0.836–0.90 0.75–0.83–0.95 0.88–0.943–1.02 a
Abbazzi et al. (1998).
0.90–1.107–1.15 0.90–1.0–1.20 1.08–1.085–1.09
0.10–0.167–0.24 0.16–0.195–0.25 0.25–0.266–0.30 2.25–2.458–2.65 2.20–2.44–2.80 2.60–2.61–2.62 7 6 2
0.10–0.156–0.20 0.13–0.16–0.18 0.16–0.186–0.19
21.43–25.97–28.57 16.67–19.43–26.32 12.90–20.83–25.92 13.33–19.56–26.67 16.67–21.68–28.89 16.13–20.34–26.88 0.70–0.721–0.75 40.00–44.59–47.00 0.90–0.91–0.95 43.40–45.07–46.00 0.8803–0.9737–1.0434 42.11–44.73–47.56 2.10–2.36–2.50 0.90–1.020–1.15 0.10–0.126–0,20 2.45–2.50–2.65 1.10–1.137–1.15 0.15–0.197–0.26 2.4779–2.5975–2.7388 1.0434–1.1626–1.2390 0.1630–0.1957–0.228 7 4 6
0.15–0.187–0.20 0.15–0.19–0.25 0.1304–0.2012–0.228
16.25–20.98–25.71 16.66 11.11–18.92–30.00 15.79–20.83–25.00 22.50–29.105–35.71 22.58 11.76–17.38–37.50 11.67–21.16–31.25 43.75–43.875–44.00 44.16 41.82–43.65–46.00 40.82–44.31–49.02 0.70–0.75–0.80 0.93 0.80–0.88–0.95 0.75–0.84–0.95 1.05–1.07–1.10 1.06 1.05–1.14–1.20 1.00–1.10–1.25
0.18–0.215–0.25 0.21 0.10–0.183–0.25 0.105–0.175–0.25 2.40–2.45–2.50 2.40 2.50–2.614–2.75 2.45–2.537–2.65 2 1 7 5
Litvin Shamin Karai–Dubina Nagornoe 1 Port–Katon (Margaritovka) Zapadnye Kairy Korotoyak (Ostrogozhsk. suite) Ushkalka Roksolany Colle Curtia
0.13–0.155–0.180 0.155 0.10–0.191–0.27 0.12–0.16–0.20
B/W A/L W L
A
B N
C
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Name of locality
Table 1 Measurements and indexes of M/1 of Allophaiomys from different sites on the Russian Plain and at Colli Curtia (Italy)
Dniester terrace directly below the flood-plain loam with positive magnetization (Jaramillo paleomagnetic event). The rodent fauna from the Roksolany site is marked by the appearance of the lagurid P. pannonicus. A. pliocaenicus teeth from this site have the advanced enamel differentiation of 66.0–75.0–100 ðN ¼ 6Þ and rather low A/L ratio of 37.50–40.48–42.85 ðN ¼ 6Þ (Table 1) (Markova and Kozharinov, 1998). Eolagurus argyropuloi, Lagurodon arankae, Clethrionomys sokolovi also occur in this fauna. Mimomys is represented primarily by M. savini and Mimomys pusillus. Two other Eastern European localities (Ushkalka and Zapadnye Kairy; lower Dnieper basin) are very similar in species composition to that described above (Markova, 1998). Paleomagnetic analysis of the loesspalesol series of the Zapadnye Kairy locality, carried out by Dr. M. Pevzner, revealed the Brunhes/Matuyama boundary above two fossil soils and intermediate loams overlying the alluvial deposits containing bones of small mammals. A similar situation is found in Ushkalka site (Fig. 1) (Velichko et al., 1983). All these small mammal faunas are marked by the first appearance of the lagurid P. pannonicus which is direct descendant of P. ternopolitanus, found in earlier Nogaiskian faunas (Topachevski, 1973; Rekovets, 1994; Markova, 1992, 1998). Faunas of this evolutional level are abundant on the Russian Plain and are characterized also by A. pliocaenicus with advanced morphotypes of teeth structure, as well as E. argyropuloi, Lagurodon arankae and Clethrionomys sokolovi, Mimomys savini and Mimomys pusillus. The faunas of this stage have been named the Kairian assemblage, based on the fossil record from the locality Zapadnye Kairy (Markova, 1998). A very important locality of this evolutional stage is the Korotoyak locality (Don River basin). Iosifova, Krasnenkov and Semenov described the geology and magnetization of this locality (Iosifova and Krasnenkov, 1994; Iosifova et al., 1992). One stratum of this complicated section, the Ostrogozhsk suite, contains a small mammal fauna similar to the Roksolany one. Dr. A. Agadjanian and Dr. N. Kazantseva studied the Ostrogozhsk fauna and published a list of small mammals (Krasnenkov and Agadjanian, 1975; Agadjanian and Kazantseva, 1994). Recently, the morphology of Arvicolidae from this site was analyzed. From the paleomagnetic data, this layer corresponds directly with the zone of positive magnetization of the Jaramillo paleomagnetic event (Iosifova and Semenov, 1998). The rodent fauna from the Ostrogozhsk suite includes the remains of Mimomys pusillus; M. savini; Mimomys sp.; Clethrionomys ex gr. sokolovi; E. argyropuloi; P. pannonicus; and A. pliocaenicus, among others. The teeth of Allophaiomys are characterized by positive enamel of ‘‘microtus’’ type and the A/L ratio of 42.11–44.73–47.56 ðN ¼ 6Þ (Fig. 1: 1–3, 7; Table 1).
C/W
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Several of the M/1 have a rather simple structure of anteroconid complex (Fig. 2). Two M/1 have a more evolved feature of anteroconid complex resembling Microtus oeconomus (Fig. 1: 9-10). These two teeth possibly belong to one animal and represent morphological variation. The remnants of more evolved species of the Allophaiomys–Microtus phylogenetic line, such as Terricola and Microtus, have not been found in this layer. Some comparisons have been made with the Allophaiomys remains from the Colle Curti locality (Central
Italy). The small mammal remains have been found in deposits with normal polarity (Jaramillo event), which are located between the reversed polarity thicknesses of the Matuyama Chron. According to Coltorti et al. (1998, Fig. 5), the fossiliferous layer is located in the lowest part of the normal polarity event. According to some of the morphological features (A/ L index) the Allophaiomys remnants from the Ostrogozhsk suite are more evolved than Allophaiomys sp. from Colle Curti. Several other characteristics (L, W, C/ W) suggest that they are similar. The B/W index differs
Fig. 2. The structure of m/1 Allophaiomys pliocaenicus from Korotoyak (Ostrogozhsk suite).
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in the Ostrogozhsk suite and Colle Curti samples and shows more primitive morphology in Allophaiomys from Colle Curti (Table 1) (Abbazzi et al., 1998). A mixture of primitive (low development of anteroconid complex) and derived (microtine differentiation of enamel) features of Allophaiomys teeth was recognized from Colle Curti. On the whole, the Colle Curti fauna is more similar to the Roksolany one, which was recovered from deposits directly below the Jaramillo positive event. More primitive features of Allophaiomys from Colle Curti compared to Korotoyak (Ostrogozhsk suite) can be explained in two ways: (1) these two faunas correspond to different parts of Jaramillo event—the Colle Curti site corresponds to the very beginning of this interval, the Ostrogozhsk suite—a younger; (2) the very few remnants of Allophaiomys teeth from the Colle Curti site (only 2 m/1) prevent sufficient morphological comparison. Thus, the Eastern European rodent faunas corresponding to the Jaramillo event are characterized by the presence of the following Arvicolidae: Mimomys savini, Mimomys pusillus, Clethrionomys sokolovi, A. pliocaenicus (more advanced type), Borsodia fejervaryi, P. pannonicus, Lagurodon arankae and E. argyropuloi. 4.2. Faunas for the period between the Jaramillo event and the Brunhes/Matuyama boundary Small mammal faunas of this evolutional level are distinguished by the first appearance of more advanced voles Microtus (Stenocranius) hintoni and M. (Terricola) sp., which evolved from the voles of the genus Allophaiomys. These faunas were first described by Alexandrova (1976) from Morozovka 1 locality, near Odessa town. Later, faunas similar in species composition were found in the Port-Katon section, Taman peninsula (Markova, 1982, 1990b) (Fig. 1). Dr. A. Chepalyga and Dr. V. Udartsev recovered the small mammal remains. A similar fauna was found in the fluvial-liman deposits of the seventh Danube terrace near Nagornoe village. Besides rare Allophaiomys remains, bones of Mimomys savini, Pliomys episcopalis, Microtus (Terricola) arvalidens have been determined here (Mikhailesku and Markova, 1992). All the faunas of this evolutionary level feature the presence of an advanced type of Allophaiomys and late Mimomys (M. savini and Mimomys pusillus). Steppe lemmings are represented by P. pannonicus, Lagurodon arankae and E. argyropuloi. A/L Allophaiomys ratio is about 45 (Table 1). The position of the Port-Katon locality is shown in Fig. 1. The geological and paleomagnetic studies have been carried out by Velichko et al. (1983). The fossiliferous layer is located under the Brunhes/Matuyama boundary and is divided from it by four soils with hydrogenous characters. The faunas of this species
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composition were determined as a Morozovian assemblage (Aleksandrova, 1976; Markova, 1992, 1998). The most advanced Early Pleistocene faunas are distinguished by the first appearance of Microtus (Pallasiinus) ex gr. oeconomus. The faunas have been named after the Petropavlovka locality in the Don drainage basin and are described by Alexandrova (1976) and Agadjanian (Krasnenkov and Agadjanian, 1975). A similar evolutionary level, rich fauna of small mammals has been found in the lower reaches of the Dnieper River, near the Karai–Dubina village (Markova, 1982, 1998, Markova and Kozharinov, 1998; Rekovets, 1994). In addition, another fauna of this evolutionary level has been discovered on the Taman Peninsula from Priozernoe, and is being examined. Steppe and yellow lemmings P. pannonicus, E. argyropuloi and voles Microtus (Stenocranius) hintoni formed the core of the Petropavlovian faunas. The presence of rhizodont voles Mimomys savini and Mimomys pusillus, as well as very few remnants of late Borsodia (B. fejervaryi) are also typical for these faunas. Scarce Allophaiomys teeth recovered from the localities are distinct for their advanced morphotypes (Table 1). Their enamel is differentiated according to Microtus type. Pevzner (Velichko et al., 1983) carried out the paleomagnetic studies of the sections with the Petropavlovian faunas. He revealed that the fossil-bearing strata belonged to the upper part of the Matuyama Chron. In the Karai–Dubina section the Brunhes/ Matuyama boundary is documented in the floodplain alluvium overlying the channel alluvial facies with the micromammals (Fig. 1). The Shamin locality (Don Basin) has yielded teeth of Microtus arvalinus as well as remains of Microtus (Stenocranius) hintoni and P. posterius. This site is correlated with the very end of the Matuyama Chron. The Shamin fauna is more advanced than the Petropavlian ones and has been attributed to the Early Tiraspolian fauna (Markova, 1982, 1998). In the Bolshevik 2 locality (layers II and III) near Odessa the Brunhes/Matuyama boundary has been established between layers III (lower) and II (Rekovets and Nadachowski, 1995). The small mammals from these layers (lagurids and microtins) resemble the Shamin and Litvin’ assemblages, but differ from Karai–Dubina and Petropavlovka ones. In the latter lagurids are more primitive in appearance. Unfortunately the Bolshevik assemblages contain significantly high proportion of redeposit remains of small mammals. 4.3. Faunas located above the Brunhes/Matuyama boundary Some small mammal localities were found above the Brunhes/Matuyama boundary. The earliest of them is
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an Early Tiraspolian fauna from the Litvin locality (Taman Peninsula). It contains rodents closely resembling the Shamin fauna. This fauna has been recovered from the Chauda marine deposits dated to the beginning of Brunhes Epoch. Both marine (Didacna pseudocrassa Pavl., Didacna pazvula Naliv., Dreissena palymorpha Pallas) and freshwater (Unio pictorum L., Viviparus viviparous L., V. tiraspolitanus Pavl.) mollusks have been found in these deposits (Chepalyga, personal communication). This layer also yields the remains of small mammals: Mimomys savini, A. pliocaenicus, P. pannonicus, E. simplicidens gromovi, Microtus (Stenocranius) hintoni and Microtus (Microtus) arvalinus (Markova, 1992). The core of this fauna consists of steppe and yellow lemmings and Microtus (Stenocranius) hintoni and Microus arvalinus.
5. Conclusions The numerous data provide original information about rodent history during the Early–Middle Pleistocene transition in Eastern Europe. The rodent faunas, synchronous and near the Jaramillo event (1.1– 1.0 Ma), are characterized by the presence of Mimomys savini, Mimomys pusillus, Borsodia fejervaryi, P. pannonicus, Lagurodon arankae and Allophaiomys pliocaenicus. More advanced faunas of Morozovian assemblage are distinguished by the FAD of Terricola and the ‘‘pitymys’’—like vole Microtus (Stenocranius) hintoni. These faunas are correlated with the upper part of the Matuyama Chron. The first occurrence of Microtus ex gr. oeconomus and Microtus arvalinus have been marked at the very end of Matuyama Chron. The arhizodont cementless voles P. pannonicus (advanced type) and E. argyropuloi, and rhizodont voles Mimomys savini and Mimomys pusillus coexisted in these faunas with the very evolved and rare Allophaiomys. The ancient representative of the Stenocranius lineage—Microtus (Stenocranius) hintoni, became abundant in these faunas. Rodent faunas recorded from deposits from the beginning of the Brunhes positive polarity Chron are very similar to the earlier ones and include several extinct species. The revealed rodent sequence permits long-distant correlation of mammalian data and geological deposits from the other parts of Europe.
Acknowledgements Dr. Yu. Iosifova is deeply acknowledged for her permission to study the Rodentia fauna from Korotoyak (Ostrogozhsk suite). I would also like to thank Dr. A. Dodonov for the geological information about the Roksolany section and Dr. A. Chepalyga for his
constructive comments about the geology and chronology of the Litvin site. I am very grateful to T. Adrian for her careful edition of the English translation. The author thanks the organizers of the SEQS International Conference in Bari (Italy, 2000) focusing on Pliocene– Pleistocene and Early–Middle Pleistocene transitions.
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Eastern European rodent (Rodentia, mammalia) faunas from the Early–Middle Pleistocene transition Anastasia K. Markova Institute of Geography RAS, Staromonetny 29, 109017 Moscow, Russian Federation Available online 11 September 2004
Abstract The rodent faunas from nine sites on the Russian Plain and Taman Peninsula corresponding to the Early–Middle Pleistocene transition have been analyzed. These localities are situated in the Dnieper, Don, Dniester and Danube drainage basins and in the northern regions of the Black and Azov Seas. Seven of the sites are associated with the upper part of Matuyama reversed polarity Chron and located between the beginning of Jaramillo event and the Brunhes/Matuyama boundary. One of the sites was recovered from the Chauda marine deposits of the Black Sea and referred to the very beginning of Brunhes Epoch. The species composition of the rodent fauna sequence has been established. The faunas referred to the Jaramillo event (1.1–1.0 Ma) contain the advanced Mimomys, steppe lemmings Prolagurus pannonicus and Lagurodon arankae, and Allophaiomys pliocaenicus. Faunas were recovered from Korotoyak (Ostrogozhsk suite), Roksolany, Zapadnye Kairy and Ushkalka localities. The more advanced rodent faunas of the Morozovian assemblage (Port-Katon site) include also the first representatives of Terricola. Karai–Dubina and Shamin localities have been found below the Brunhes/Matuyama boundary (0.85 Ma), including the first representatives of Microtus (Microtus ex gr. oeconomus and Microtus arvalinus). The ‘‘Pitymys’’ like voles—Microtus (Stenocranius) hintoni, steppe lemming P. pannonicus and yellow steppe lemming Eolagurus argyropuloi compose the core of these faunas. The Litvin locality, located in the deposits of the Chauda transgression of the Black Sea and corresponding to the very beginning of the Middle Pleistocene, contains a rodent fauna closely resembling the Shamin one. It yielded the remains of Mimomys savini, advanced A. pliocaenicus, P. pannonicus, Microtus (Stenocranius) hintoni, Microtus ex gr. oeconomus and Mcirotus arvalinus. Thus, numerous data permit elucidation of the history of rodent faunas of the Russian Plain during Early–Middle Pleistocene transitions and reveal the definitive stages in their evolution. r 2004 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction The Middle–Early Pleistocene transition is increasingly recognized as a time of significant biotic changes in the Northern Hemisphere. The visible climatic changes (progressive cooling) are reflected in vegetation and animal assemblages. The open landscapes and the steppe mammal communities became widely distributed on the southern part of the Russian Plain in the second part of the Early Pleistocene and the beginning of the Middle Pleistocene. The well-pronounced evolutional changes in rodent fauna have been identified in material from E-mail address: [email protected] (A.K. Markova).
Eastern Europe for this same time interval. These data have been confirmed by geological as well as geochronological studies. The principal stages in rodent evolution are distinguished by the first appearance (FAD) of new taxa. These data permit the reconstruction of the sequence of evolutional changes in rodent faunas, and the identification of characteristic faunal assemblages, from the beginning of the Jaramillo paleomagnetic event until the beginning of the Brunhes paleomagnetic Epoch. The stratigraphical position of the sites was ascertained by direct geological and also paleomagnetic studies. Such studies are important for elucidating not only rodent evolution, but also the stratigraphy of Eastern Europe and correlation with other regions.
1040-6182/$ - see front matter r 2004 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2004.07.020
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2. Materials Rodent faunas from nine localities on the Russian Plain dating to the Early–Middle Pleistocene transition have been studied. All the localities are located between 511N and 451N. Most of the localities are associated with alluvium deposits of the main rivers of Russian Plain. Karai-Dubina, Zapadnye Kairy and Ushkalka faunas have been found in terrace deposits of the lower Dnieper drainage basin; Nagornoe 1 fauna have been recovered in the lower Danube drainage basin; the Roksolany site is in alluvium deposits of the Dniester river (Markova, 1982, 1990a,b; Mikailesku and Markova, 1992); Korotoyak (Ostrogozhsk suite) and Shamin faunas have been recovered in Don-river alluvium (Iosifova et al., 1992; Markova, 1992). Port-Katon and Litvin localities have been revealed on the Taman Peninsula in marine-liman deposits (Markova, 1990b, 1998). The faunas of all of these sites were studied (Markova, 1982, 1990a,b, 1998a; Bylinsky et al., 1993). Dr. A. Agadjanian and Dr. N. Kazantseva analyzed firstly the small mammals from the Korotoyak (Ostrogozhsk suite) locality (Agadjanian and Kazantseva, 1994).
3. Terminology This paper uses the terminology for the parts of occlusial surface of vole molars introduced by van der
Meulen (1973, 1974): L—length of molar; W—width of molar; AC 2—anterior cap; ACC—anteroconid complex; PL—posterior lobe; T1–T5—dentile fields of occlusial surface of molars; BRA—buccal re-entrant angle, BSA—buccal salient angle; LRA—lingual reentrant angle; LSA—lingual salient angle; B—shortest distance between AC 2 and T4 and T5; C—shortest distance between BRA 3 and LRA 3 angles. The SDQ index is used, after Heinrich (1978), which shows the ratio between the enamel thickness on posterior and anterior parts of the salient angles of the tooth. This work uses the measurements of the enamel of the three central angles of m1 (LSA2, LSA3, BSA2) and terms the SDQ index as SDQ3.
4. Stages of rodent faunas 4.1. Faunas, approximately synchronous to the Jaramillo paleomagnetic event The earliest faunas of the discussed time interval have been found in fluvial deposits at the Roksolany site (lower Dniester basin). Small mammal remains have been collected during field works headed by Mikhailesku (Mikhailesku and Markova, 1992) and Dodonov (Dodonov et al., 1998). The fossiliferous layers have been recovered here below a thick loess-soil section where the Brunhes—Matuyama boundary and Jaramillo event were established (Dodonov et al., 1998) (Fig. 1). The bones were found in the channel alluvium of the
Fig. 1. Geological structure and paleomagnetic data of the sections with small mammal faunas: 1—soil; 2—loess; 3—loam; 4—clay; 5—sand; 6— limestone; 7—water level; 8—small mammal remains; 9—mollusks remains; 10—borrows of small mammals; 11—Brunhes/Matuyama boundary; 12—Jaramillo event.
ARTICLE IN PRESS 11.76–18.62–25.0 13.68–19.43–25.00 18.18–19.79–22.58 12.50–19.88–22.50 18.95–23.54–31.25 26.88–28.23–29.41 40.0–45.33–46.0 37.50–40.48–42.85 41.22–41.57–41.92 0.80–0.836–0.90 0.75–0.83–0.95 0.88–0.943–1.02 a
Abbazzi et al. (1998).
0.90–1.107–1.15 0.90–1.0–1.20 1.08–1.085–1.09
0.10–0.167–0.24 0.16–0.195–0.25 0.25–0.266–0.30 2.25–2.458–2.65 2.20–2.44–2.80 2.60–2.61–2.62 7 6 2
0.10–0.156–0.20 0.13–0.16–0.18 0.16–0.186–0.19
21.43–25.97–28.57 16.67–19.43–26.32 12.90–20.83–25.92 13.33–19.56–26.67 16.67–21.68–28.89 16.13–20.34–26.88 0.70–0.721–0.75 40.00–44.59–47.00 0.90–0.91–0.95 43.40–45.07–46.00 0.8803–0.9737–1.0434 42.11–44.73–47.56 2.10–2.36–2.50 0.90–1.020–1.15 0.10–0.126–0,20 2.45–2.50–2.65 1.10–1.137–1.15 0.15–0.197–0.26 2.4779–2.5975–2.7388 1.0434–1.1626–1.2390 0.1630–0.1957–0.228 7 4 6
0.15–0.187–0.20 0.15–0.19–0.25 0.1304–0.2012–0.228
16.25–20.98–25.71 16.66 11.11–18.92–30.00 15.79–20.83–25.00 22.50–29.105–35.71 22.58 11.76–17.38–37.50 11.67–21.16–31.25 43.75–43.875–44.00 44.16 41.82–43.65–46.00 40.82–44.31–49.02 0.70–0.75–0.80 0.93 0.80–0.88–0.95 0.75–0.84–0.95 1.05–1.07–1.10 1.06 1.05–1.14–1.20 1.00–1.10–1.25
0.18–0.215–0.25 0.21 0.10–0.183–0.25 0.105–0.175–0.25 2.40–2.45–2.50 2.40 2.50–2.614–2.75 2.45–2.537–2.65 2 1 7 5
Litvin Shamin Karai–Dubina Nagornoe 1 Port–Katon (Margaritovka) Zapadnye Kairy Korotoyak (Ostrogozhsk. suite) Ushkalka Roksolany Colle Curtia
0.13–0.155–0.180 0.155 0.10–0.191–0.27 0.12–0.16–0.20
B/W A/L W L
A
B N
C
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Name of locality
Table 1 Measurements and indexes of M/1 of Allophaiomys from different sites on the Russian Plain and at Colli Curtia (Italy)
Dniester terrace directly below the flood-plain loam with positive magnetization (Jaramillo paleomagnetic event). The rodent fauna from the Roksolany site is marked by the appearance of the lagurid P. pannonicus. A. pliocaenicus teeth from this site have the advanced enamel differentiation of 66.0–75.0–100 ðN ¼ 6Þ and rather low A/L ratio of 37.50–40.48–42.85 ðN ¼ 6Þ (Table 1) (Markova and Kozharinov, 1998). Eolagurus argyropuloi, Lagurodon arankae, Clethrionomys sokolovi also occur in this fauna. Mimomys is represented primarily by M. savini and Mimomys pusillus. Two other Eastern European localities (Ushkalka and Zapadnye Kairy; lower Dnieper basin) are very similar in species composition to that described above (Markova, 1998). Paleomagnetic analysis of the loesspalesol series of the Zapadnye Kairy locality, carried out by Dr. M. Pevzner, revealed the Brunhes/Matuyama boundary above two fossil soils and intermediate loams overlying the alluvial deposits containing bones of small mammals. A similar situation is found in Ushkalka site (Fig. 1) (Velichko et al., 1983). All these small mammal faunas are marked by the first appearance of the lagurid P. pannonicus which is direct descendant of P. ternopolitanus, found in earlier Nogaiskian faunas (Topachevski, 1973; Rekovets, 1994; Markova, 1992, 1998). Faunas of this evolutional level are abundant on the Russian Plain and are characterized also by A. pliocaenicus with advanced morphotypes of teeth structure, as well as E. argyropuloi, Lagurodon arankae and Clethrionomys sokolovi, Mimomys savini and Mimomys pusillus. The faunas of this stage have been named the Kairian assemblage, based on the fossil record from the locality Zapadnye Kairy (Markova, 1998). A very important locality of this evolutional stage is the Korotoyak locality (Don River basin). Iosifova, Krasnenkov and Semenov described the geology and magnetization of this locality (Iosifova and Krasnenkov, 1994; Iosifova et al., 1992). One stratum of this complicated section, the Ostrogozhsk suite, contains a small mammal fauna similar to the Roksolany one. Dr. A. Agadjanian and Dr. N. Kazantseva studied the Ostrogozhsk fauna and published a list of small mammals (Krasnenkov and Agadjanian, 1975; Agadjanian and Kazantseva, 1994). Recently, the morphology of Arvicolidae from this site was analyzed. From the paleomagnetic data, this layer corresponds directly with the zone of positive magnetization of the Jaramillo paleomagnetic event (Iosifova and Semenov, 1998). The rodent fauna from the Ostrogozhsk suite includes the remains of Mimomys pusillus; M. savini; Mimomys sp.; Clethrionomys ex gr. sokolovi; E. argyropuloi; P. pannonicus; and A. pliocaenicus, among others. The teeth of Allophaiomys are characterized by positive enamel of ‘‘microtus’’ type and the A/L ratio of 42.11–44.73–47.56 ðN ¼ 6Þ (Fig. 1: 1–3, 7; Table 1).
C/W
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Several of the M/1 have a rather simple structure of anteroconid complex (Fig. 2). Two M/1 have a more evolved feature of anteroconid complex resembling Microtus oeconomus (Fig. 1: 9-10). These two teeth possibly belong to one animal and represent morphological variation. The remnants of more evolved species of the Allophaiomys–Microtus phylogenetic line, such as Terricola and Microtus, have not been found in this layer. Some comparisons have been made with the Allophaiomys remains from the Colle Curti locality (Central
Italy). The small mammal remains have been found in deposits with normal polarity (Jaramillo event), which are located between the reversed polarity thicknesses of the Matuyama Chron. According to Coltorti et al. (1998, Fig. 5), the fossiliferous layer is located in the lowest part of the normal polarity event. According to some of the morphological features (A/ L index) the Allophaiomys remnants from the Ostrogozhsk suite are more evolved than Allophaiomys sp. from Colle Curti. Several other characteristics (L, W, C/ W) suggest that they are similar. The B/W index differs
Fig. 2. The structure of m/1 Allophaiomys pliocaenicus from Korotoyak (Ostrogozhsk suite).
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in the Ostrogozhsk suite and Colle Curti samples and shows more primitive morphology in Allophaiomys from Colle Curti (Table 1) (Abbazzi et al., 1998). A mixture of primitive (low development of anteroconid complex) and derived (microtine differentiation of enamel) features of Allophaiomys teeth was recognized from Colle Curti. On the whole, the Colle Curti fauna is more similar to the Roksolany one, which was recovered from deposits directly below the Jaramillo positive event. More primitive features of Allophaiomys from Colle Curti compared to Korotoyak (Ostrogozhsk suite) can be explained in two ways: (1) these two faunas correspond to different parts of Jaramillo event—the Colle Curti site corresponds to the very beginning of this interval, the Ostrogozhsk suite—a younger; (2) the very few remnants of Allophaiomys teeth from the Colle Curti site (only 2 m/1) prevent sufficient morphological comparison. Thus, the Eastern European rodent faunas corresponding to the Jaramillo event are characterized by the presence of the following Arvicolidae: Mimomys savini, Mimomys pusillus, Clethrionomys sokolovi, A. pliocaenicus (more advanced type), Borsodia fejervaryi, P. pannonicus, Lagurodon arankae and E. argyropuloi. 4.2. Faunas for the period between the Jaramillo event and the Brunhes/Matuyama boundary Small mammal faunas of this evolutional level are distinguished by the first appearance of more advanced voles Microtus (Stenocranius) hintoni and M. (Terricola) sp., which evolved from the voles of the genus Allophaiomys. These faunas were first described by Alexandrova (1976) from Morozovka 1 locality, near Odessa town. Later, faunas similar in species composition were found in the Port-Katon section, Taman peninsula (Markova, 1982, 1990b) (Fig. 1). Dr. A. Chepalyga and Dr. V. Udartsev recovered the small mammal remains. A similar fauna was found in the fluvial-liman deposits of the seventh Danube terrace near Nagornoe village. Besides rare Allophaiomys remains, bones of Mimomys savini, Pliomys episcopalis, Microtus (Terricola) arvalidens have been determined here (Mikhailesku and Markova, 1992). All the faunas of this evolutionary level feature the presence of an advanced type of Allophaiomys and late Mimomys (M. savini and Mimomys pusillus). Steppe lemmings are represented by P. pannonicus, Lagurodon arankae and E. argyropuloi. A/L Allophaiomys ratio is about 45 (Table 1). The position of the Port-Katon locality is shown in Fig. 1. The geological and paleomagnetic studies have been carried out by Velichko et al. (1983). The fossiliferous layer is located under the Brunhes/Matuyama boundary and is divided from it by four soils with hydrogenous characters. The faunas of this species
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composition were determined as a Morozovian assemblage (Aleksandrova, 1976; Markova, 1992, 1998). The most advanced Early Pleistocene faunas are distinguished by the first appearance of Microtus (Pallasiinus) ex gr. oeconomus. The faunas have been named after the Petropavlovka locality in the Don drainage basin and are described by Alexandrova (1976) and Agadjanian (Krasnenkov and Agadjanian, 1975). A similar evolutionary level, rich fauna of small mammals has been found in the lower reaches of the Dnieper River, near the Karai–Dubina village (Markova, 1982, 1998, Markova and Kozharinov, 1998; Rekovets, 1994). In addition, another fauna of this evolutionary level has been discovered on the Taman Peninsula from Priozernoe, and is being examined. Steppe and yellow lemmings P. pannonicus, E. argyropuloi and voles Microtus (Stenocranius) hintoni formed the core of the Petropavlovian faunas. The presence of rhizodont voles Mimomys savini and Mimomys pusillus, as well as very few remnants of late Borsodia (B. fejervaryi) are also typical for these faunas. Scarce Allophaiomys teeth recovered from the localities are distinct for their advanced morphotypes (Table 1). Their enamel is differentiated according to Microtus type. Pevzner (Velichko et al., 1983) carried out the paleomagnetic studies of the sections with the Petropavlovian faunas. He revealed that the fossil-bearing strata belonged to the upper part of the Matuyama Chron. In the Karai–Dubina section the Brunhes/ Matuyama boundary is documented in the floodplain alluvium overlying the channel alluvial facies with the micromammals (Fig. 1). The Shamin locality (Don Basin) has yielded teeth of Microtus arvalinus as well as remains of Microtus (Stenocranius) hintoni and P. posterius. This site is correlated with the very end of the Matuyama Chron. The Shamin fauna is more advanced than the Petropavlian ones and has been attributed to the Early Tiraspolian fauna (Markova, 1982, 1998). In the Bolshevik 2 locality (layers II and III) near Odessa the Brunhes/Matuyama boundary has been established between layers III (lower) and II (Rekovets and Nadachowski, 1995). The small mammals from these layers (lagurids and microtins) resemble the Shamin and Litvin’ assemblages, but differ from Karai–Dubina and Petropavlovka ones. In the latter lagurids are more primitive in appearance. Unfortunately the Bolshevik assemblages contain significantly high proportion of redeposit remains of small mammals. 4.3. Faunas located above the Brunhes/Matuyama boundary Some small mammal localities were found above the Brunhes/Matuyama boundary. The earliest of them is
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an Early Tiraspolian fauna from the Litvin locality (Taman Peninsula). It contains rodents closely resembling the Shamin fauna. This fauna has been recovered from the Chauda marine deposits dated to the beginning of Brunhes Epoch. Both marine (Didacna pseudocrassa Pavl., Didacna pazvula Naliv., Dreissena palymorpha Pallas) and freshwater (Unio pictorum L., Viviparus viviparous L., V. tiraspolitanus Pavl.) mollusks have been found in these deposits (Chepalyga, personal communication). This layer also yields the remains of small mammals: Mimomys savini, A. pliocaenicus, P. pannonicus, E. simplicidens gromovi, Microtus (Stenocranius) hintoni and Microtus (Microtus) arvalinus (Markova, 1992). The core of this fauna consists of steppe and yellow lemmings and Microtus (Stenocranius) hintoni and Microus arvalinus.
5. Conclusions The numerous data provide original information about rodent history during the Early–Middle Pleistocene transition in Eastern Europe. The rodent faunas, synchronous and near the Jaramillo event (1.1– 1.0 Ma), are characterized by the presence of Mimomys savini, Mimomys pusillus, Borsodia fejervaryi, P. pannonicus, Lagurodon arankae and Allophaiomys pliocaenicus. More advanced faunas of Morozovian assemblage are distinguished by the FAD of Terricola and the ‘‘pitymys’’—like vole Microtus (Stenocranius) hintoni. These faunas are correlated with the upper part of the Matuyama Chron. The first occurrence of Microtus ex gr. oeconomus and Microtus arvalinus have been marked at the very end of Matuyama Chron. The arhizodont cementless voles P. pannonicus (advanced type) and E. argyropuloi, and rhizodont voles Mimomys savini and Mimomys pusillus coexisted in these faunas with the very evolved and rare Allophaiomys. The ancient representative of the Stenocranius lineage—Microtus (Stenocranius) hintoni, became abundant in these faunas. Rodent faunas recorded from deposits from the beginning of the Brunhes positive polarity Chron are very similar to the earlier ones and include several extinct species. The revealed rodent sequence permits long-distant correlation of mammalian data and geological deposits from the other parts of Europe.
Acknowledgements Dr. Yu. Iosifova is deeply acknowledged for her permission to study the Rodentia fauna from Korotoyak (Ostrogozhsk suite). I would also like to thank Dr. A. Dodonov for the geological information about the Roksolany section and Dr. A. Chepalyga for his
constructive comments about the geology and chronology of the Litvin site. I am very grateful to T. Adrian for her careful edition of the English translation. The author thanks the organizers of the SEQS International Conference in Bari (Italy, 2000) focusing on Pliocene– Pleistocene and Early–Middle Pleistocene transitions.
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