Quaternary mammalian faunal complexes

Main article: Paleolithic in Russia

Six large faunal complexes have been identified that are genetically related to each other. Complexes are common in vast territories. In time, faunal complexes correspond to the Quaternary period and the end of the Neogene. However, some researchers recognize the quaternary system in a wider scope. With this understanding, all faunal complexes should be attributed to the Quaternary period. From bottom to top (from ancient to young) these complexes are as follows:

2.6-2.2 million hp: Khaprovsky (Eopleistocene, Akchagyl) faunal complex

Haprovsky Faunal Complex (2.6-2.2 Ma, Paleopleistocene, Middle Villafrunk, Late Villania)

It is the oldest complex. Along with the ancient elephant, Stenon's horse, camel, it includes such relic forms as the Auvergne mastodon, hipparion and saber-toothed tiger.

The Khaprovsky faunal complex was established by V.I. Gromov in the 1930s on the northern coast of the Sea of ​ ​ Azov near the Khapry station.

The Hapra faunal complex is compared by almost all researchers with the Villafranca fauna of Western Europe.

The species composition of the Haprov fauna indicates that at that time steppe and forest-steppe spaces were widespread in the south of the European part of the former USSR. The reconstructed climate of this time is quite warm, with a well-defined hot dry season and a mild winter period.

The basis of the isolation of the complex is a representative collection of vertebrate remains from the Haprovsky, Liventsovsky, Mokro-Chaltyrsky quarries of the North-Eastern Azov region, as well as from a number of coastal and ravine outcrops (Merzhanovo, Morskaya, Volovaya Balka, etc.). Subsequently, oriktocenoses similar in age were discovered in other regions of Eastern Europe (for example, the locations of Etulia-3, Cherevichnoye-2, Kryzhanovka-2, -3, Northwestern Black Sea region; Saber, Stavropol Territory; Mozdok, North Caucasus; Kushkuna in Transcaucasia).

As part of the Haprov fauna, animals occupying precisely forest-steppe biotopes (about 11% of species) are widely represented. This group includes animals, the ranges of modern analogues of which are associated mainly with rare woody vegetation, pegs and shrub thickets, -

• Gromov's elephant Archidiskodon meridionalis gromovi,
• rhinoceros Stephanorhinus,
• camel Paracamelus alutensis,
• олень Eucladoceros dicranios tanaitensis, Arvernoceros cf. verestchagini, Metacervoceros rhenanus,
• the root-toothed vole of the genus Pityimomys and the first forest vole of Clethrionomys primitivus.

Typical representatives of steppe landscapes (45% of species) are the mole-worm Spalax sp., hamster Allocricetus sp., Borsodia praehungarica praehungarica and pliomys Pliomys ucrainicus, Equus (Allohippus) livenzovensis and Equus sp., antelopes Gazellospira gromovae and Gazella sp. The presence of Struthio ostrich in the fauna indicates the absence of snow cover in cold periods. In a diverse fauna of predators are represented

• Canis dog cf. senezensis,
• Pliocrocuta perrieri hyena,
• Lynx issiodorensis steppe lynx,
• cheetah Acinonyx pardinensis,
• saber-toothed cat Homotherium crenatidens.

Haprovsky orictocenosis is characterized by a small number of bovid remains - Gazella, Gazellospira, Tragelaphinae, Leptobos (about 1%), common for the steppe and forest-steppe landscapes of the Pliocene and Pleistocene of the entire steppe and forest-steppe zone from Western Europe to China. Inhabitants of floodplain and bayrac forests and edges are considered

• tuberculate mastodon Anancus arvernensis alexeevae,
• boar Sus strozzii,
• лось Cervalces (Libralces) gallicus,
• giraffe Palaeotragus priasovicus,
• Ursus bear cf. etruscus,
• raccoon dog Nyctereutes megamastoides,
• representatives of the Kuny family.

Of the representatives of microteriofauna, indicators of forested landscapes are the Clethrionomys forest vole. Mimomys praepliocaenicus and Dolomys voles were associated with near-water stations. For woody and shrubby the vegetation of beams and watersheds was characterized by such inhabitants as moles and shrews, avoiding both heavily waterlogged and very dry places [Titov, 2008].

Thus, the landscapes of the Azov region and the Ciscaucasia region, adjacent to the Kuyalnitsky Sea (Akchagyl Sea in the Caspian Sea), during the existence of the Haprovsky theriocomplex, were forest-steppe savannoy-like landscapes - vast steppe spaces on watersheds interspersed with pegs of woody and shrubby vegetation, as well as floodplain and bayrac forests. The faunal community in its appearance and the presence of forms such as like elephants, rhinos, horses, giraffes and antelopes, resembles the association of modern African savannas. However, from ancient and recurrent African associations, the fauna of southern Eastern Europe differs significantly at the generic and species levels [Alekseeva, 1977; Titov, 2008; Arambourg, 1979; Geraads, 1997; Geraads, Amani, 1998].

The structure of the Haprovian association reveals the originality of this community, consisting in a combination of the forms usual for the beginning of the early Pleistocene (middle villafrank) of the entire Palearctic (Nyctereutes, Canis, Pannonictis, Pliocrocuta, Homotherium, Acinonyx, Lynx, Anancus, Equus (Allohippus), Gazella, Eucladoceros) and the Black Sea endemic form of the small camel Paracamelus alutensis.

The geographical location of the Azov region led to the existence in this territory during the early Pleistocene, along with Asian forms (Elasmotherium, large Paracamelus, Gazellospira), animals characteristic mainly of the territory of Central and Southern Europe (Archidiskodon, Hipparion, large Equus (Allowippus), Sus strozzii, Arvernoceros) [Titov, 2008].

There are no unambiguous findings indicating the existence of a person at this time. However, in the outcrops of Haprovsky layers opening in the area of ​ ​ the village. Matveev Kurgan (Rostov region), separate finds of stones were discovered with suspected traces of human exposure [Khokhlovkina, 1940; Kachevskiy, Litvinenko, 2010].

2.2-1.6 million hp: Psekup faunal complex

Psekup faunal complex (2.2-1.6 Ma, Paleopleistocene, late villafrank, end of late villania - beginning of early biharia).

Initially, this association was described from the territory of the North Caucasus (p. Psekups, the location of Baku, Saratov), but subsequently its finds have been identified in other locations from southern Eastern Europe and adjacent regions (Bolshaya Kamyshevakha, Northern Black Sea region; Georgievsk, Palan-Tyukan, Mukhkai II, North Caucasus) [Garutt, Safronov, 1965; Alekseeva, 1977; Titov, Shvyryova, 2016; Amirkhanov et al., 2016]. Typical representatives of this association are

• слоны Archidiskodon meridionalis meridionalis, Phanogoroloxodon mammonthoides,
• rhino Stephanorhinus aff. etruscus,
• Equus horses (Allohippus) cf. major, E. (A.) ex gr. stenonis,
• camel Paracamelus alutensis,
• the giraffe Palaeotragus cf. priasovicus,
• олени Eucladoceros orientalis orientalis, Megaloceros stavropolensis,
• bull Leptobos cf. etruscus,
• the antelope Gazellospira torticornis and
• Gallogoral meneghinii goat.

In ancient Psekup rodent fauna, relatively high-crown root-toothed voles of lagurins of the group Borsodia newtoni-arankoides, Mimomys pliocaenicus and Pitymimomys pitymyoides were common; primitive mole moles Ellobius (Ellobius) cf appear. kujalnikensis. Clethrionomys kretzoii forest voles become numerous. In the late Psekup (Odessa) faunas (Tizdar), the first non-corniparous voles Allophaiomys deucalion and Lagurodon/Prolagurus appear.

The general appearance of the Psekup fauna is similar to that of the Haprovsky. Which suggests a wide distribution of forest-steppe landscapes of the savannah-like type and a generally warm climate.

The end of the Paleopleistocene is characterized by a number of locations indicating the existence of representatives of the genus on the territory of the Caucasus at this level Homo. Tools of labor together with bone remains of animals are known, for example, from the monuments of Kermek (section of Tizdar, Taman Peninsula) and Mukhkai II (Dagestan) [Amirkhanov et al., 2016; Shchelinsky et al., 2016]. This level is well consistent with the age-close location of Dmanisi (Transcaucasia), from which the remains of Homo erectus ergaster are known [Lordkipanidze et al., 2013].

Camel metapody fragment with notch marks [Sablin, Girya, 2010] ^[1]," originating from the vicinity of Rostov-on-Don, is an indirect evidence of the penetration of ancient people on the northern coast of the Kuyalnitsky Sea.

Paracamelus alutensis is a typical representative of the Eastern European fauna of the late Pliocene

The find was made by N.K. Vereshchagin in 1954 in the Liventsov career, when the quarry was just beginning to be developed.

According to the testimony of V.S. Baigusheva (oral message, 2010), at the time of the find only the upper layers of the alluvial stratum were penetrated. From this top pack floodplain sand facies subsequently discovered the remains of rodents of the Psekup faunal complex [Tesakov, 2004]. While the bulk of the findings of the Haprovsky faunal complex from this quarry comes from the lower channel strata of Haprovsky alluvium [Titov, 2008]. Therefore, the artifact with explicit traces of processing should be attributed to the existence of the Psekup theriocomplex.

Incisions were made with a chopper made of sintered limestone on fresh and dry bone

1.6-0.8 million hp: Taman (Absheron) faunal complex

Taman faunal complex (1.6-0.8 Ma, eopleistocene, the end of the late villafranca - the beginning of galeria, early biharia).

It is characterized by the appearance of the southern elephant, Caucasian elasmotherium, Taman boar, Taman beaver, etc. Ancient relics are disappearing. The Taman faunal complex is located on the Taman Peninsula by V.I. Gromov. Bone remains lie in alluvia.

In addition to the Taman Peninsula, the finds of this faunal complex are known in a number of regions of Ukraine and the North Caucasus. In more northern areas, animal remains from this era are unknown. This complex in Western Europe corresponds to the late stages of the development of the Villafranca fauna.

The remains of animals of the Taman theriocomplex are known from a number of locations of the Taman Peninsula (Tsimbal, Akhtanizovskaya, Blue Balka/ Bogatyri), Northeast Azov Region and Lower Don (Port Katon, Semibalki, Nogaisk, Sarkel) [Vereshchagin, 1957; Topachevsky, 1965; Rekovets, 1994; Baigusheva, 2000; Baigusheva, Titov, 2008; Tesakov et al., 2007; Sotnikova, Titov, 2009; Vislobokova, Titov, 2020]. Fauna of this level are also known from the Northwestern Black Sea region: Western Cairo, Ushkalka, Roksolany, Chishmikioy [Markova, 1982; Tesakov, 2004].

Elasmotherium

Characteristic forms of association are

• Canis tamanensis, Canis (X.) lycaonoides dogs,
• otter Lutra simplicidens tamanensis,
• Pachycrocuta brevirostris hyena,
• leopard Panthera sp.,
• saber-toothed cat Homotherium latidens,
• слон Archidiskodon meridionalis tamanensis,
• Elasmotherium caucasicum rhinoceros,
• Equus ( A.) cf. major, E. (A.) cf. suessenbornensis,
• олень Eucladoceros orientalis pliotarandoides,
• bison tamanensis and
• antelope Pontoceros ambiguus.

The locations of this level are characterized by a combination of root-toothed voles of the genus Mimomys, as well as non-root-toothed voles of Allophaiomys pliocaenicus, pied Prolagursus pannonicus и Lagurodon arankae. Pliocrocuta, Anancus, Paracamelus, Arvernoceros, Palaeotragus, Gazellospira and Leptobos, common for paleopleistocene, become extremely rare or completely disappear during the existence of the Taman complex. Eolagurus, Panthera, Pontoceros, Bison become common. In late Taman time, Stenocranius voles migratively appear.

Paleoteriological and palynological materials indicate the distribution of steppe and forest-steppe landscapes in the Azov region along the banks of the Gurian basin and the rivers flowing into it [Shchelinsky et al., 2010]. The climate in southern Eastern Europe in the Eopleistocene remained quite warm and probably without severe winters. However, at this time, in the zone of open and semi-open landscapes in southern Eastern Europe, the increasing influence of aridization began to manifest itself to a greater extent.

On the Taman Peninsula to a number of locations of the Taman faunal the complex is dedicated to the finds of the tools of the oldest man (Springs-1, -2; Tsimbal). In particular, numerous early Paleolithic stone products come from the type location of the Taman Blue Beam/Bogatyri faunal association [Shchelinsky et al., 2010]. Here in total, according to N.K. Vereshchagin [1957], I.A. Dubrovo [1964] and the authors, at the time of 2020 were discovered remains of at least 100 elephants and 45 elasmotheria.

The formation of this unique tafocenosis was probably due to the caldera of one of the large mud volcanoes typical of the Taman Peninsula. As a result of a number of factors, the skeletons of large, mainly thick-skinned animals - elephants and rhinos - were killed and buried in this place. Animals died in this place, bogged down in a furnace of mud, unable to get out of the basin with steep banks, became easy prey for predators or died from poisonous gases periodically emitted by the volcano. The accumulation of carcasses did not occur simultaneously, but over a relatively long time.

Animals of different individual ages, both young and old individuals, died here. However, the most numerous remains belong to sexually mature and adult animals. A similar ratio of age groups is observed in herds of modern elephants and rhinos. Thus, no selectivity in the death of animals is observed. Among the numerous bones of elephants and elasmotheria, separate remains of predators - wolves and homotherium [Kazanov et al., 2019; Sotnikova, Titov, 2009]. All these facts suggest that the accumulation of carcasses attracted, along with predators and scavengers, people who, with the help of their massive stone tools, could extract valuable food resources here [Titov et al., 2018].

Tiraspol (Lower Pleistocene) faunal complex (Q1)

It is installed in the lower Dniester in sand and gravel deposits (in Tiraspol gravel). Gravel deposits with fauna make an ancient hollow that stretches along the left bank of the Dniester near Tiraspolyai.

The main representatives of the Tiraspol faunal complex are the elephant Wust, Mosbach's horse, Merck's rhino, red deer, Shetenzak's bison, broad-throated elk, complex deer, Deninger's bear.

The Tiraspol faunal complex of the Russian Plain, compared to the older ones (Haprovsky, Taman), is distinguished by the absence of a number of animals (mastodon, southern elephant, mahayrodus). At the same time, new species of animals appear in it (Wusta elephant, Mosbach horse, Shetenzak bison).

Tiraspol fauna in the south of the Russian Plain and the North Caucasus lived in steppe and forest-steppe landscapes. To the north (in the vicinity of Moscow) at that time, forest landscapes probably existed.

Khazar (Middle Pleistocene) faunal complex of the Libyan interglacial period (Q12)

It is identified by V.I. Gromov in the Lower Volga region under the name "Volga fauna." For the Khazar complex, the trogonterian elephant, the Khazar horse, the long-horned bison, the Knobloch camel, the Merck rhino are the most typical.

The finds of the Khazar complex are known on the vast territory of Eurasia (Transbaikalia, France, British Isles). This complex (its individual representatives) occupies a belt between 45 and 60 ° N.

The species composition of the Khazar fauna indicates the existence of open steppe spaces with a sharply continental climate in the Lower Volga region at that time. In the Middle Volga region (at the mouth of the Kama), forest faunal forms (elk, beaver) are noted as part of the Khazar fauna, indicating the presence at that time of zonality close to modern. In time, the complex corresponds to the first half of the Middle Pleistocene era.

Upper paleolytic (mammoth) faunal complex (Q 22 - Q3)

In time, it corresponds to the second half of the Middle Quaternary and the entire Upper Quaternary era. At this time, the Great (maximum) glaciation causes a wide distribution of cold-loving mammals. These include the early mammoth form and woolly rhino. In addition, animals appear that are characteristic of the tundra. The late Upper Paleolithic complex is most fully represented. For him, the late type of mammoth, horse, short-horned bull, reindeer, saiga, etc. are common.

The Upper Paleolithic faunal complex is so named because the predominant number of its locations is associated with Upper Paleolithic human sites and partly Mousterian monuments. The remains of this fauna are known from a vast area. Geomorphological conditions for the occurrence of bone residues are diverse.

The most characteristic feature of the upper paleotic complex is that it contains animals of various ecological groups: forest, steppe and tundra. Northern animals live together with temperate and southern. In this regard, the complex in question is often referred to as a "mixed fauna."

This complex of animals, peculiar in its species composition, was formed at the beginning of the Dnieper glaciation. It existed in the vast spaces of Eurasia until the end of the Valdai glaciation. In the Holocene, it was replaced by modern fauna.

The Upper Paleolithic complex differs from the older complexes in that it contains many northern animals. At that time, these animals lived much further south (reindeer, arctic fox, musk fox). At the same time, some other animals of the Upper Paleolithic faunal complex lived north than we see now. So, a horse, a saiga, a bison are found in Taimyr and the Novosibirsk Islands. Now they're not there.

By the time of the formation of the Upper Paleolithic complex within the European part of the former USSR, a number of animals that were characteristic of the previous Khazar complex disappeared. This is the trogonterian elephant, Merck's rhino, elasmotherium, long-horned bison. Instead, new species are widespread, more adapted to cold climatic conditions. These are mammoth, woolly rhino, etc., such animals arose as a result of evolutionary transformations of animals of the Khazar faunistic complex. At the beginning of the formation of the faunal complex, the mammoth belonged to an early type. It was then replaced in the second half by a late-type mammoth.

The Upper Paleolithic faunal complex remains more or less constant in species composition throughout its existence. At the same time, vegetation has changed repeatedly and significantly. This was a consequence of significant climate change. As you can see, the fauna of mammals compared to flora is less sensitive to changes in physical and geographical conditions. The different "sensitivity" of fauna and flora is one of the main causes of the disagreements and inconsistencies that exist in the paleogeographic constructions of some paleozoologists and paleobotanics.

Modern (Holocene) faunal complex

The modern complex includes all modern and some endangered Pleistocene animals. The Holocene faunal complex is characterized by three features:

1) extinction of large Pleistocene forms,

2) the appearance of pets,

3) the formation of modern biocenoses.

The modern fauna is depleted compared to the Upper Paleolithic complex. This was due to the extinction of the mammoth, woolly rhino, cave lion. At the same time, the range of other animals (reindeer, musk fox, arctic fox, saiga) has significantly narrowed.

In general, it should be noted that the fauna of the anthropogen is closely related to the older Neogene fauna. The main difference is the appearance of real elephants, bulls, horses, etc.

On the territory of the former USSR, most of the finds of mammal fauna are located in the southern regions. In the middle lane, the number of them is sharply reduced. At the same time, in the middle strip, almost all finds belong to the Upper Paleolithic complex. The northern regions of the European part of the former USSR are poor in fauna finds. The fauna found here is most often represented by single bone remains. In the north of the Russian Plain, faunistic remains belong only to the Upper Paleolithic complex. The older fauna of the area remains unknown. In this regard, the Quaternary deposits of the northern regions are more often divided on the basis of glaciation data, and to the south - by fauna.

Notes