Rodent Squad - rodentia bowdich, 1821 VI. SUBORMS MYOMORPHA Brandt, 1855 4. FAMILY HAMSTERS - CRICETIDAE Fischer, 1817 4. SUBSET. FIELDS - ARVICOLINAE Gray, 1821 12. GENUS BRANDY VOILS - LASIOPODOMYS Lataste, 1887
VI. SUBORMS MYOMORPHA Brandt, 1855
4. FAMILY HAMSTERS - CRICETIDAE Fischer, 1817
4. SUBSET. FIELDS - ARVICOLINAE Gray, 1821
12. GENUS BRANDY VOILS - LASIOPODOMYS Lataste, 1887
1. Brandt Vole - Lasiopodomys (Lasiopodomys) brandti Radde
Body length up to 148 mm, tail - 18-30 mm (about 1/5 of the body length). The color of the top is light, grayish-sand, mottled with rare blackish hairs. The tail is one-color, yellowish. Claws on the middle fingers of the forelimb are about 2/3 of their length (Fig. 183, 2). The soles in the hindquarters are covered with hair hiding small corns. In the karyotype, 2n = 34.
The skull is wide, with a well-developed interorbital crest, without depression in the posterior frontal bones. Hearing drums are enlarged, including due to mastoid bones. The incisor holes are relatively long. Front unpaired loop of the lower M1 with a uniformly rounded outer edge without a dentate in the posterior region.
Fig. 218. Skull of Brandt's vole (Lasiopodomys brandti)
VK - upper indigenous, NK - lower
The steppes of southern Transbaikalia to the north to about Aginskoye, were not found in the Selenga valley, but found to the west, on the Kosogolsky tract (Mondy). North Mongolia from the Basin of the Great Lakes to the east to the foothills of the Great Khingan, south to the latitude of Mandalgobi, North-East. China, Korean Peninsula
Lifestyle and importance to humans
It inhabits cereal and grass-wormwood steppes of plains and mountains up to an altitude of 2000 m above sea level. m. It reaches the highest abundance in the dry steppes, where it often settles on gravelly and sandy loamy soils, less often in lacustrine meadows. In contrast to most other lowland vole species, it leads a daytime lifestyle. It emits a characteristic sharp, circulating whistle. It settles in holes, the more complex of which have up to 10 holes, several "pantries" and nesting chambers. Family burrows can reach 30 m in length and, during periods of high abundance, merge to form “towns” that can stretch for many kilometers (Mongolia). Before mass vegetation begins, it feeds on the underground, and later on, the aboveground parts of plants. They predominate in stocks, the weight of which can exceed 10 kg. At their expense, animals exist in winter, when activity is greatly reduced. At the same time, autocoprophagy was noted, indicating low-fatness and poor digestibility of stored plants. Fertility is high, in favorable years there can be 4 or even 5 broods, and the number of young in a litter can reach 12-15 and even 17 (usually 6-8). The number is subject to sharp fluctuations, outbreaks of mass breeding are common, usually coinciding with years of persistent droughts, at which time it can reach 600 individuals per 1 ha.
In Mongolia, during the years of mass breeding, it becomes a serious pest of pastures, destroying valuable fodder plants in large areas, and through its intensive digging activity, it contributes to maintaining a fallow regime in the steppes.
It harms garden crops (Transbaikalia). The natural carrier of the causative agents of tularemia and salmonellosis, and in Mongolia one of the main carriers of the causative agent of plague.
There are 3 subspecies, it is believed that in the territory of the former USSR the nominative L. (L.) m. brandti Radde, 1861.
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Action spectrum: common vole (Microtus arvalis), Eastern European vole (Microtus rossiaemeridionalis), public (steppe) vole (Microtus (Sumeriomys) socialis), herd (narrow-cranium) vole (Microtus (Stenocranius) gregalis), large (vole) , field mouse (Apodemus agrarius), house mouse (Mus musculus), baby mouse (Micromys minutus), common forest mouse (Apodemus (Sylvaemus) sylvaticus), water vole (water rat) (Arvicola terrestris), Brandt vole (Lasiopodom , shelf (Myoxus glis), gray rat (Rattus norvegicus) and other mouse-like rodents.
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Table of contents of the dissertation Doctor of Biological Sciences Rutovskaya, Marina Vladimirovna
Chapter 1. Material and methods
1.1. Technique for recording, processing and analysis of sound signals
1.2. Methodology for observing social and vocal behavior in a group
1.3. The volume of material and the methodology for studying the inheritance of signs of sound signals in interspecific hybrids
1.4. Methodology for constructing a hierarchical classification of species by
Chapter 2. Sound signaling of voles of the subfamily Arvicolinae
Lemming Vinogradov Dicrostonyx vinogradovi (Ognev, 1948)
Red Vole Myodes (Clethrionomys) glareolus (Schreber, 1780)
Tien Shan vole Myodes (Clethrionomys) centralis Miller, 1906
American Forest Vole Myodes (Clethrionomys) gapperi Vigors, 1830
Red Vole Myodes (Clethrionomys) rutilus Pallas, 1779
Red-gray vole Myodes (Clethrionomys) rufocanus Sundervall, 1846-1847
Shikotan vole Myodes (Clethrionomys) sikotanensis Tokuda, 1935
Steppe pestle Lagurus lagurus Pallas, 1773
Yellow pestle Eolagarus luteus Eversmann, 1840
Afghan Vole Blanfordimys afghanus Thomas, 1912
Brandt Vole Lasiopodomys brandti Radde, 1861
Chinese vole Lasiopodomys mandarinus Milne-Edwards, 1871
Herd (narrow-bore) vole Microtus (Stenocranius) gregalis Kastschenko, 1923
House-vole Microtus (Pallasiinus) oeconomus Miller ,! 899
Lake Vole Microtus (Pallasiinus) limnophilus Büchner, 1889
Middendorff vole Microtus (Alexandromys) middendorffii Poljakov, 1881
Far Eastern vole (large) Microtus (Alexandromys) fortis Thomas, 1911
Polevka Maksimovich (Uyghur) - Microtus (Alexandromys) maximowiczii
Evoron vole Microtus (Alexandromys) evoronensis Kovalskaja et Sokolov, 1981
Dagestan vole Microtus (Terrícola) daghestanicus Schidlovsky, 1919
Shrub vole of Shelkovnikov Microtus (Terrícola) scltelkovnikovi
Dark (plow) vole Microtus agrestis L., 1761
Common Vole Microtus arvalis Pallas, 1779
Eastern European Vole Microtus levis Miller, 1908
Trans-Caspian vole Microtus transcaspicus Satunin, 1905
Public or steppe vole Microtus (Sumeriomys) socialis Pallas, 1773
Kopetdag vole Microtus (Sumeriomys) irani Thomas, 1921
Gunther vole Microtus (Sumeriomys) guentheri Danford et Alston, 1880
Chapter 3. Intraspecific variability of sound signals
voles of the subfamily Arvicolinae
Chapter 4. The functional significance of squeaks and the formation
vocal repertoire of voles of the subfamily Arvicolinae
Chapter 5. Signs of sound signals of interspecific hybrids
Signs of sound signals of hybrids of red and red voles
Signs of sound signals of hybrids of red and Tien Shan voles
Signs of sound signals of hybrids of ordinary and Eastern European
Chapter 6. Species specificity of signs of sound signals and
phylogenetic relationships between species of voles
Chapter 7. The influence of environmental factors on the species
specific features of vole sound signals
Appendix 1. Material Volume
Appendix 2. Characteristics of sharp squeaks of different types
Appendix 3. Characteristics of silent squeaks of different types
Appendix 4. Characteristics of a hazard warning signal of different types
Appendix 5. Characteristics of singing of different types
Introduction of the dissertation (part of the abstract) on the topic “Variability and formation of sound communication of voles of the subfamily Arvicolinae”
The natural population has a complex internal structure (Naumov, 1963, Shilov, 1977), the formation and maintenance of which is based on communicative behavior (Nikolsky, 1984, Bradbury, Vehrencamp, 1998). The evolution of species, including their sociality, is inextricably linked with the evolution of intraspecific communicative processes, as described by C. Darwin (1953). E. Mayr (1974) in his work “Population of species and evolution” also notes the role of behavior, in particular communications, in speciation.
In recent decades, interest in the evolution of communications, including acoustic, has been very high (Hauser, 1996, Noble, 1998, Searcy, Nowicki, 2005, Blumstein, 1999, 2007). The technical ability to record and then analyze sound in the middle of the last century led to the rapid development of interest in the acoustic communication channel (Ilyichev et al., 1975). Much work has been devoted to the evolution of communication in birds (Panov, 1978, Searcy, Andersson, 1986, Ecology and Evolution. 2009), and less research has been conducted on mammals (Nikolsky, 1984), with particular attention paid to the evolution of alarm, (Maynard Smith, 1965, Hirth , McCullough, 1977, Shelley, Blumstein, 2005, etc.). The evolution of mammalian sound signals is often considered in connection with the origin of human speech (Lieberman, 1968, Fitch, 2000, etc.).
Considering the various issues of the evolution of communication, one must be aware that the system is formed under the influence of a number of factors. N. Tinbergen (1952) defined four main directions in the study of animal communications:
1. Mechanistic: the study of the mechanisms (eg, nervous, physiological, psychological) underlying the demonstrations (signals), or what factors regulate behavior?
2. Ontogenetic: the study of genetic factors and the influence of the environment in the development of signs of signals, or in what way is the behavior formed in ontogenesis?
3. Functional: the study of the effectiveness of communication for survival and reproduction, or what is its adaptive value?
4. Phylogenetic: the study of the evolutionary history of a species, so as to understand what features of demonstrations evolved from ancient ancestors, or how behavior is formed in phylogenesis?
In each of these areas, there is already a lot of research, including for acoustic communication.
Sound communication arose in terrestrial vertebrates based on the breathing system: sound occurs when a stream of air from the lungs passes through oscillating valves with a pressure of 100 to 2000 Pa, which depends on the strength of the respiratory muscles and lung volume, which, in turn, depend on the linear size animal (Fletcher, 2007). This valve, the larynx (larynx), originally appeared in air-breathing fish to protect delicate membranes and epithelium from damage and ingress of water and foreign particles when swallowing into the lungs (Negus, 1929). The larynx consists of cartilage and paired closing muscles. The latter have folds of loose connective tissue and are called vocal cords. Sound is produced by modulating the flow of air from the lungs (Berke, Long, 2010). In mammals, they are controlled by 5 muscles that control the closure of ligaments to protect the bronchus and control breathing. During vocalizations, the respiratory and laryngeal muscles are involved in the work, with the help of which the air pressure, mass and elasticity of the ligaments are regulated, which is necessary to control the frequency and amplitude of sound (Höh, 2010).
Vibrations of the mammalian vocal cords determine the fundamental frequency of sound, which usually corresponds to the lowest band of the spectrum, and multiple harmonics corresponding to the vibration frequencies of their parts. The sound formed when passing through the vocal tract (pharynx, oral and nasal cavities and additionally the trachea in birds) changes: at the output we have maximum frequencies (formants), which are determined by the length of the vocal tract: (üL / c = (2п-1) to / 2 in the sequence 1,3,5 where c is the speed of sound, co is the angular velocity, L is the length of the vocal tract (Fletcher, 2007). Asynchronous vibrations of the vocal cords lead to the appearance of a noise component in the signal (Fitch et al., 2002 ), amplitude modulation - to the appearance of close-packed side frequencies that form practical Eski uniform spectrum filling (Nikolsky 2007a, 2011).
An important component of the vocalization process is its nervous regulation. In mammals, a limited number of brain regions are involved in the production of sounds. This is primarily the limbic system, especially the cingulate gyrus of the lower surface of the hemisphere and the hypothalamus. The limbic system is responsible for emotions and memory. The midbrain innervates the larynx, and the caudal brainstem coordinates respiration and sound production (Newman, 2010).
C. Darwin (Darwin, 1953), in addition to various expressions of emotions in animals and humans, described the emotional vocalizations of animals as a way to convey their emotional status. It is possible that the very first sound signal was a cry of pain, which became the basis for the development of various vocal adaptations to warn other individuals of danger (Panksepp, 2010). Emotional signals have wide variability, which, however, has its own laws.
M. Kiley (1972) showed that most sounds made by domestic ungulates are typological continua expressing animal “excitation levels” or “main motivational states”. M. Kili believes that the sounds specific to certain situations, at least among ungulates, are negligible, and even the extreme elements of continua can be connected by transitional forms.Basically, sound reactions transmit information not about a specific condition of an animal, such as, for example, an aggressive or sexual state, but rather about the level of interest of an animal in a stimulus, which is a reflection of the main motivational level of animal excitement. According to M. Keeley, one of the main conditions causing and enhancing sound reactions is a state of frustration, "when the animal wants something, waits for something, or cannot take possession of something."
Another approach to the definition of communication is “Animal communication takes place if it is shown that one animal influenced the behavior of another. As a rule, the influence falling under the category of “communication” is mediated by the signals that the animal receives from the sensory organs. ” (Corsini, Auerbach, 2006). This regulatory approach was proposed by R. Dawkins and J. Krebs (Dawkins, Krebs, 1978), who believed that natural selection should support the behavior most favorable for the survival and reproductive success of an individual to a greater extent than cooperative behavior. Therefore, they proposed replacing the cooperative concept of communication (based on group selection) with the concept of manipulating the behavior of the recipient (supporting selection at the individual level). The essence of the concept is that the individual - the source of the signal with his help changes the behavior of the recipient to his advantage. According to this approach, the signal contains information that is not necessarily pragmatic, but may be false. However, in this case, if the signal does not benefit the recipient, then the latter will stop responding to it at all. And then communication as such will not take place.
We will follow an informational approach in which communication exists when animals begin to exchange information. The exchange of information assumes that the signal should be perceived by the recipient, and, therefore, the range of the received signal should correspond to the sensitivity of the recipient’s hearing aid, the ability to perceive and differentiate the frequency spectrum, as well as the rhythmic structure of sound (Ehret and Kurt, 2010). In addition, the acoustic characteristics of the environment, such as humidity, strong winds, temperature gradients, as well as the acoustic properties of biotopes, can impose significant restrictions on the propagation of sound signals, especially transmitted over long distances. So sound attenuation is maximum at low humidity of less than 20% and the higher the frequency, the greater it is (Ingard, 1953). Therefore, in the arid zones, lower frequencies spread better in the daytime. For example,
associated with habitats in deserts is associated with a lower than what could be expected from the size of the animal, the frequency of sound signals of the large gerbil (Nikolsky, 1973, Golydman et al., 1977). Sound propagation in the natural environment can be influenced by wind at a speed exceeding 4 m / s, at which turbulence develops, and the medium becomes inhomogeneous. The probability of signal loss in this situation can be reduced by repeated repetition: when a signal consisting of a series of pulses passes, the probability of a signal falling at times with the least attenuation increases. Such a “windproof” signal, for example, is possessed by two semi-desert species of ground squirrels - small and yellow (Nikolsky, 1984).
The commonality of the laws underlying the formation of vocal reactions in mammals allowed E. Morton (Morton, 1977) to formulate “motivational-structural rules”: in the “behavioral spectrum” E. Morton identified “endpoints” - “hostility” and friendliness ”. The former are expressed by sharp broadband sounds, the latter by pure and relatively higher-frequency sounds. At the same time, E. Morton notes that these rules work with close contacts. With increasing transmission distance of acoustic information, there is a need to increase noise immunity, which leads to an exception to the rule. According to E. Morton, low sounds have a repellent effect, as initially characteristic of large animals. Tall, on the other hand, are attractive because they are typical for cubs. G. Eret (2006) formulated the rules for the perception of sound signals, including the perception of the biological meaning of the issued signals. They almost completely coincide with the generalizations of E. Morton (Morton, 1977): (1) relatively high-frequency tonal sounds expressing appeasement or emotional fear, and cause interest, are perceived as "attractive", (2) soft, low-frequency rhythmic sounds expressing " friendly "emotions that usually accompany the peaceful interactions of animals in groups - are perceived as a sign of" cohesion ", (3) sharp, loud, noisy and noisy sounds that express aggressiveness cause evasion behavior, thus, are perceived to aka repulsion.
An essential detail in the idea of motivational-typological gradients was introduced by J. Eisenberg (Eisenberg, 1974), using the example of sound signaling of representatives of the porcupine suborder, he showed that the repertoire of sound signals in them is represented by two systems. One reflects the three main "moods". The second is an “indicator of arousal,” reflecting the motivational levels of the source. Thus, the signs of the signal reflect the level of excitation of the source, but at the same time the typological continuum can be torn into “functional classes” having their own characteristic signs, and within them the signs continually vary, reflecting the level of excitation of the animal.
The early ethologists explain the appearance of communication by the origin of signals from nonspecific behavior, which in the process of animal interactions gains informational and communicative value (Tinbergen, 1952, Smith, 1977). This is accompanied by a ritualization process, as a result of which the resulting signal has a number of characteristics that allow the recipient to accurately identify and adequately evaluate it.
The term “ritualization” was proposed by J. Huxley (1914), who was the first to note that some actions in the process of phylogenesis lose their own original function and turn into purely symbolic ceremonies that carry a communicative function. His ideas were developed by N. Tinbergen (Tinbergen, 1952), developing a theory of ritualization. He believed that communicative signals could come from the so-called intentional movements, biased activity, or redirected behavior.
In order to fulfill its communicative function, the signal must attract the attention of the recipient, be adequately perceived by him and, when transferred through a medium separating partners, be minimally distorted. Therefore, under the pressure of natural selection in the process of ritualization, those complexes of behavior that begin to bear a communicative function undergo significant modifications. First, ritualized behavior becomes more expressive and exaggerated than the original form of activity. Secondly, ritualized movements become strictly regulated in speed and amplitude, that is, they acquire a fixed intensity. Due to this, the invariance of the signals is created, which increases the efficiency of information transfer. Third, ritualized movements often become stereotyped, incomplete, and repeatedly repeated (Tinbergen, 1952).
Thus, the parameters of the signal are determined by the capabilities of the sound source and can be modified under the influence of directional selection in order to optimize its passage through the medium and its perception by the recipient. Directional selection is possible primarily because the signs of sound signals in most terrestrial mammals are genetically inherited. Inheritance of signs of sound signals has been shown in a number of works. The first was F. Frank (Frank, 1967), who found a mutation in hybrid lines of the common vole (Microtus arvalis) that changes cries in aggressive situations. Voles emitted a trill, or did not emit sounds at all, while normal animals emitted single signals. F. Frank also showed that the inheritance of acoustic behavior obeys the laws of Mendel. Another option for proving the inheritance of signs of an audio signal was proposed by P. Winter et al. (Winter et al., 1973). In his experiment, squirrel saimiri (Saimirí sciureus) cubs were raised by females, operatively devoid of voice. In adolescents, a normal vocal repertoire characteristic of this species was formed. Ability
mammals (we do not consider such specialized groups as marine mammals and bats) are poorly developed to learn and imitate sound signals, even in close relatives of humans - monkeys (Kozarovitsky, 1965, Ladygina-Kote, 1965). However, there are serious differences between the ability of animals to imitate new, and even more speech-like sounds, and sounds that differ little from those that are included in the acoustic repertoire of the species (Ladygina-Kote, 1965). According to her observations, a chimpanzee easily identifies with a person when the latter plays sounds borrowed from the everyday life of the chimpanzee itself. For some species, imitation of the sounds of relatives is a characteristic feature of their behavior. First of all, this is a phenomenon of fusion of individual characters, in which sound signals are executed synchronously by a group of individuals of the same species. Such synchronization was described in the Palawan tupa group (Tupaia palovanensis) (Williams et al., 1969), in the siamang duets (Symphalangus syndactylus) (Lamprecht, 1970), in the group of jackals (Canis aureus) (Nikolsky, Poyarkov, 1979) and in a number of other social mammals.
Inheritance of signs of sound signals in hybrids, as a rule, is of an intermediate nature; this was seen in sound signaling in hybrids of large cats (Peters, 1978). Intermediate, in relation to the parameters of the signals of the parental species, are warning signals of the hybrids of the steppe and gray marmots recorded from the natural hybridization zone (Nikolsky et al., 1982, 1983), as well as signs of sound signals of the hybrids of cattle and bison living in Askania-Nova (Rutovskaya, 1983).
Genetic inheritance of signs of sound signals suggests that in the process of evolution, directed selection can act on them if the evolving communication system increases the survival and reproductive success of the species. And this is contrary to the fact that in any communication the animal spends energy on producing a signal, spends time (instead of, for example, feeding behavior or grooming), while issuing and perceiving a signal, the risk of being caught by a predator increases (Burlak, 2011).
However, the association of individuals makes it possible to optimize the use of fodder resources or shelters, in a group it is easier to detect danger and protect oneself from predators and competitors in terms of resources, partners and offspring, living in a common nest contributes to energy saving during collective thermoregulation, etc. Payment for combining individuals is associated with losses as a result of increased competition for resources or partners, the risk of parasitic diseases and attractiveness for predators (Chabovsky, 2006).
The communicative system evolves in conjunction with the social system of the species. Thus, E. Shelley and D. Blumstein (Shelley, Blumstein, 2005) for 209 species of rodents showed that
the appearance of an alarm and the degree of its complexity are correlated with the daily way of life and the degree of sociality of the species.
The acoustic communication of mammals, as well as olfactory, forms and maintains the structure of populations. At the same time, signals can work as consolidating and regulating intraspecific integration. For example, the acoustic activity of a wolf synchronizes the motivational state of association in the schooling period, and regulates spatial intra-group relationships (Nikolsky and Frommolt, 1989). Another example: a fur seal rookery, the population structure of which is largely determined by sound communication - a set of male sounds supports the territorial structure of the rookery (Lisitsyna, 1981), contact cries make it possible to maintain mother-child relationships and find each other after forced separation, for example, if the mother goes to feed at sea (Lisitsyna, 1980).
Using an informational approach to the study of communication is not easy to explain cases where communicative signals are not “honest”. For example, using a general pattern: the larger the animal, the lower the formant and fundamental frequencies — you can virtually increase your size by underestimating the signal frequency. For example, in males of a koala, a pair of indescribable large ligaments was found inside the pharyngeal mouth, allowing them to publish a frequency 20 times lower than the 8 kg weight predicted for the animal. This, according to the authors, facilitates the identification of the size of an individual, which can serve as an indicator of quality
male (Charlton et al., 2007). Studying the role of signals in sexual choice, A. Zahavi (1975) tried to solve the dilemma of honest communication: the female chooses a high-quality male according to his signals, but a low-quality male can also give a “dishonest” signal, indicating its high quality, and will selected by female. Then the signal ceases to carry information about the high quality of the male and will not be supported by selection. A. Zahavi (1975) proposed a handicap theory in which only a high-class male can afford to give hypertrophied information about his quality, since this is costly and increases the risk of death from a predator. However, in this case, his signal is already becoming "honest." Currently, there are several theories that consider the evolution of communicative signals in terms of their “honesty” and price, and the conditions of natural group or individual selection (Searcy, Nowicki, 2005).
All of these theories consider the evolution of a signal in which directional natural selection plays a decisive role, leading to the specialization of communicative signals. Of particular interest are hazard warning signals, the model group of which turned out to be ground squirrels (Nikolsky, 1980, Bluimstein, 2007). An interesting model of the role of the phylogenetic history of the species in the evolution of communication turned out to be subspecies of red deer, in which
acoustic signals (roars) reflect the history of propagation (Nikolsky et al., 1979). This example suggests that the variability of the parameters of the audio signal is not necessarily the result of a directed selection of their attributes. P. Cumbell et al. (Campbell et al., 2010) proved using the example of a singing mouse (Scotinomys teguina and S. xerampelinus) that the geographical variability of an advertisement song is the result of gene drift rather than adaptation to living conditions. Thus, geographical variability, the species specificity of a species’s vocal repertoire can be either a result of directed selection or a by-product of a species’s evolution that is not directly related to the communication process (for example, due to gene drift or changes linked to morphological adaptations of the species).
Traditionally, evolutionary constructions are based on the results of comparative anatomy and morphology, and now molecular genetic methods that consider animals of different degrees of affinity. Once phylogenetic relationships between taxonomic groups have been established, a comparison of behavioral traits can provide information about their evolution (Heind, 1975). To attempt to describe the evolution of the communicative system, the subfamily Arvicolinae is ideally suited - a large group of species of voles, with varying degrees of kinship.
Voles of the subfamily Arvicolinae are small rodents of the Palearctic, most of which are represented on the territory of the former Soviet Union. All species of voles are herbivorous, active throughout the year. Most of the species are adapted to a normal lifestyle.Due to this, voles have a valky body shape with a shortened tail, limbs and outer ear and weaker differentiation of the fur cover (Gromov and Polyakov, 1977). A similar body structure determines a fairly close repertoire of motor reactions and elements of social interactions (Johst, 1967). However, the widespread distribution throughout the northern hemisphere within the Holarctic (Gromov, Polyakov, 1977) determines a wide variety of living conditions, and as a result, a wide variety of spatial distribution and social structure of populations (Gromov, 2008) from single-territorial to family-group, with high density clusters of which their settlements can be called colonies.
The immediate ancestors of microtins were voles-toothed hamsters from three extinct genera living in Europe in the Myo-Pliocene. In the late Pliocene-Holocene, modern habitats of living species were formed. In the Pleistocene, we observe the rapid evolution and generic differentiation of microtins, especially in the Old World (Gromov and Polyakov, 1977). The wide distribution and, as a consequence, the great geographical variability of many species, the ongoing evolution and radiation of this group are the reasons for the fact that many debatable issues remain in the taxonomy of this group of mammals. Various
many disputed species are attributed to different subgenera and sometimes genera. So in modern taxonomy (Pavlinov, 2006, Carleton, Musser, 2005), the mole vole (genus Ellobius), which I.M., was included in the subfamily Arvicolinae Gromov and I.I. Polyakov (Gromov, Polyakov, 1977) was considered as a separate subfamily. THEM. Gromov and I.Ya. The Poles were distinguished by the currently existing 7 tribes: Prometheomyni (Prometheus voles), Ondatrini (muskrats), Clethrionomyni (forest voles), Lagurini (pied petals), Dicrostonyxini (hoofed lemmings), Lemmini (real lemmings) and Microtini (voles). AND I. Pavlinov (2006) reduced the number of tribes. The tribe of real lemmings remained unchanged. From the Microtini tribe, he isolated a separate tribe Phenocomys (North American voles). The remaining gray voles are combined with the Ondatrini tribe in the Arvicolini tribe. And all other tribes are combined with prometheus voles in the tribe Prometheomyni. Together with mole voles (tribe Ellobiusini), 5 tribes are obtained. J. Carleton. and M. Musser (Carleton, Musser, 2005), on the contrary, increased the number of tribes to 12, having allocated a few groups into independent tribes: Neofibrini, Pliomyni, Phenacomyni and Phenacomyne. In total, the subfamily unites about 150 species of voles belonging to 28 genera. Moreover, subgeneric taxa can often be distinguished within genera. The genus Microtus is especially large and complex, including 14 subgenera and 64 species.
Differences in the taxonomy of the group among different authors is a consequence of the lack of a clear idea of the phylogenetic relationships in this group. New molecular data considered in a paleontological context (Abramson et al., 2009a) allow us to distinguish the following main stages of group diversification. The first radiation of the subfamily: basal has a late Miocene age, which is consistent with paleontological data, according to which the most primitive indisputable representatives of the group appeared about 7.0 million years ago in the Pontic deposits of Eastern Europe. The second radiation corresponds to the isolation of the ancestors of modern Clethrionomyini, the time of which, according to molecular data, corresponds to the very end of the Miocene - the beginning of the Pliocene. The divergence of cellrinomyin from the common vole trunk was after the first (basal) stage of radiation, but before the radiation of microtin-lagurin-mole rat. Based on the distribution of modern representatives and paleontological data, it can be assumed that East Asia was the center of origin of the group, and the plain and mountain forests were the initial habitat type. The third radiation of voles includes the divergence Lagurini / Eliobiusini / Arvicolini, the time of which is supposedly related to the Early Pliocene. From this it follows that the common ancestor of this group could be the Palaearctic voles of a promisimis level of organization. Lagurini and Arvicolini developed along the main path of evolution of the subfamily, increasingly adapting to the nutrition of vegetative parts of grassy plants and mastering mainly meadow (Arvicolini) and steppe (Lagurini) landscapes. Ellobiusini shows a great example of fast evolutionary adaptive