Distribution and Population Density of the Russian Desman (Desmana Moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

Home , Alatyr (river), Desman, Insar (river), Kovylkino, Lopatino, Moksha (river)

Forestry Studies | Metsanduslikud Uurimused, Vol. 71, Pages 48–68

Research paper

Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

Alexey Andreychev*, Vyacheslav Kuznetsov and Alexandr Lapshin

Andreychev, A., Kuznetsov, V., Lapshin, A. 2019. Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia. – Forestry Studies | Metsanduslikud Uurimused 71, 48–68, ISSN 1406-9954. Journal homepage: http://mi.emu.ee/forestry.studies

Abstract. To date according to the registration work, the number of the Russian desman determined actually is 588 individuals in Mordovia. Based on extrapolation, the total desman population in Mordovia is about 1,400 individuals. We may assume that the most of the desman population is concentrated on floodplain lakes and reclamation canals. The highest density of the desman population in Mordovia was recorded in Krasnoslobodsky and Temnikovsky Districts (10.5 and 8.2 burrows per km, respectively). About 400 animals may live on the shore of the rivers Vad, Partsa, Yavas, Vindrey, Nuluy, and Kundybolka. The mean value of density of the desman population on the rivers was 0.9 burrows per km of the coastline, which corresponded to habitat quality class IV (0.5–5 burrows per km). The distribution of habitats across river basins is extremely uneven. The main part of the desman population is restricted to the Moksha basin (more than 1,350 individuals), and in the Alatyr basin (left tributary of the Sura River) just twenty individuals were recorded.

Key words: Red Data Book, desman, burrow, number of individuals, river, lake, Mordovia.

Authors’ addresses: Department of Zoology, Mordovian State University, Bolshevistskaya str. 68, 430005, Saransk, Republic of Mordovia, Russia; *e-mail: [email protected]

Introduction the Republic of Mordovia was initiated by well-known mammalogist L.P. Borodin. In The Russian desman Desmana moschata (L.) his monograph and articles L.P. Borodin is in the Red List of the International Union stated the following habitat areas of the des- for Conservation of Nature (Kennerley & man within the territory of Mordovia: Zubo- Turvey, 2016; Rutovskaya et al., 2017b), the vo-Polyansky, Temnikovsky, Tengushevsky, Red Data Book (2001) of the Russian Feder- Kochkurovsky, Bolshebereznikovsky, and ation, Appendix II of the Bern Convention, Dubensky Districts (Borodin, 1963, 1970). and Red Data Books of Russian regions, in- He noted that the desman was registered in cluding the Red Data Book (2005) of Mor- the Sura, Moksha, Vad, Vysha, Yuzga, Partsa dovia. The study of the Russian desman in Rivers and their tributaries.

DOI: 10.2478/fsmu-2019-0012

© 2019 by the authors. Licensee Estonian University of Life Sciences, Tartu, Estonia. This article is an open access article distributed under the terms and conditions of the Cre- ative Commons Attribution-NonCommercial-NoDerivatives (CC BY-NC-ND) 4.0 Inter- national license (https://creativecommons.org/licenses/by-nc-nd/4.0/). 48 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

L.P. Borodin stated that there had been In the past, the Sura basin was part of natural population of the species in the ter- the area of the Russian desman. According ritory of Mordovia for a long time, which to M.N. Bogdanov (1871), the desman was was only to some extent supported by re- numerous in the Sura valley up to Penza. introducing the animals in the 1930s. In B.M. Zhitkov (1898) notes that the des- October 1937, 98 animals (46 males and 52 man was especially abundant in the mid- females) were brought to lakes Inorka and dle reach of the Sura, and along the Alatyr Valza in the Mordovsky Nature Reserve and Pyana it lived only near the mouths from Kadomsky District of Ryazan Region, of these rivers. In 1925–1928, the desman that is, from the lower Moksha floodplain. lived throughout the floodplain of the Sura On September 10, 1938, another 97 animals (Paramonov, 1928). It was also noted that were brought from Kadomsky District and the desman was common in many areas released into Inorka, Tarmenka, and Tara- along the Sura, and along the Alatyr it was tino lakes (Borodina et al., 1970). found in the lower reaches near the river Basins of the rivers Moksha and Sura mouth (Anonymus, 1936). According to are in the desman range. According to the N.S. Kufeld (1939), the desman was nu- literature dating back to the first half of the merous in the upper flow of the Sura to 20th century, in the Moksha basin the des- Inzensky District of Penza Region. In low- man was common along the Moksha, start- er areas within the borders of Mordovia, ing from the Kovylkino station and up to the desman was seen occasionally near the the river mouth, along the Partsa River with villages of Sabaevo and Permisi and far- tributaries and on the Vysha River near the ther from Bolshiye Berezniki village to the village of Kirillovo (Kufeld, 1939; Borodin, Alatyr River. Along the Alatyr, the desman 1963). According to the data for 1954, the was common from the mouth to the bor- desman in Mordovia was spread along the ders of Ichalkovsky District of Mordovia Moksha from the town of Temnikov (Tem- (Borodin, 1963; Tereshkin, 1971). Accord- nikovsky District) to the village of Noro- ing to L.P. Borodin, until 1966 the desman vatovo (Tengushevsky District) and along was found in the floodplain of the Sura in the Vad with its tributary Partsa within the Kochkurovsky, Bolshebereznikovsky and borders of Zubovo-Polyansky District. In Dubensky Districts. In 1953–1956, desmans the second half of the 1940s, I.S. Tereshkin were captured in Kochkurovsky and Bol- witnessed repeated desman captures on shebereznikovsky Districts, where 56 and the Yuzga River (the Moksha tributary) 68 individuals were caught, respectively. below the road to the village of Tengushe- The desman was seen in Dubensky District vo. The desman also got into fishing creels of Mordovia, but there was no hunting ar- on the branch connecting the large forest ranged (Borodin, 1970). lake Piyavskoe (near the village of Ivanov- In 1961, L.P. Borodin and I.S. Tereshkin ka, Tengushevsky District) with the Yuzga set up three stations to study the desman riverbed. In 1950, a pond was built near the (two in the Moksha River floodplain, and village of Aleksandrovka on the small riv- one in the Sura River floodplain) (Borodin, er Vyazhga, which joins the Moksha down 1963; Tereshkin, 1971). The first station was the village of Narovatovo (Tengushevsky laid in the Moksha floodplain between the District). In 1953, I.S. Tereshkin caught a town of Temnikov and the village of Stary desman with a dragnet in this pond. All Gorod. The floodplain width in the obser- the literary facts mentioned above for the vation area reached 5 km in diameter. The Moksha and its tributaries confirm a rath- floodplain is right-sided, mostly open, with er widespread desman distribution in this a large number of lakes. Many of them river basin, and thus indicate the long- had a muddy bottom. Ten lakes (Shirchek, lived habitat of the animal. Bobrovoe, Ogorodnoe, Vatazhnoe, Cher-

49 A. Andreychev et al. nomutskoe, Starica, Nemcovo, Krasnenkoe, (Shidlovskaya, 2018). According to these Linevo, Olyushka) with a total coastline results, the average population density in length of 8 km were chosen as references. the station was 3.5 individuals per coast- Over the years of observation, the average line km or 3.2 burrows per coastline km. number of desman burrows per coastline In view of the value of the results obtained kilometer varied from 2.9 to 4.1. for this station, the distribution of burrow The second station was set up in the densities over specific lakes should be Moksha floodplain within the territory of made since the variation is significant. For the Mordovsky Nature Reserve. The width example, the density at Lake Zaresheno of the floodplain in this area reached 4 km was 0.33 burrows per km, and at Mochilki in diameter. The floodplain is two-sided. and Ovechiy Poldnik lakes it was 22.5 bur- Nine lakes (Kuniha, Gorbatenkoe, Korlysh- rows per km and 30 burrows per km, re- ki, Krapivnoe, Chernenkoe, Chernye Lu- spectively. Moreover, the report provides zhki I, Chernye Luzhki II, Chernye Luzhki information about banding of the desman III, Dikaya) with a total coastline length of on non-station lakes, namely the Telim- 8 km were chosen as references. erka, Baklusha, and Uzhovo lakes. The third station was set up in the Sury In 1982, V.S. Vechkanov, Head of the Zo- floodplain to the east of the village of Per- ology Department, V.M. Smirnov, Head of misi, to the south of Lake Inerka, Bolshe- the Zoology Department Laboratory, and bereznikovsky District. The station lakes a group of students A.A. Ananyev, A.V. were located within Bolshebereznikovsky Shevorakov, V.M. Matyaev, Yu.A. Tomi- and Kochkurovsky Districts. Ten lakes lov, G.F. Grishutkin, S.A. Mikova per- (Pendelyukha, Chudomaz, Sarazerka, Bu- formed registration work on 163 lakes of kach, Bulyk, Kriusha krestovaya, Shirokoe Mordovia. According to the survey and bolotnoe, Dolgenkoe, Kriusha Arapovska- registration results, scientists noted the ya, Bezymyannoe) with a total coastline desman’s dwelling in water reservoirs length of 8 km were chosen as referenc- of six districts: Zubovo-Polyansky, Kras- es (Borodin, 1970). In 1968, I.S. Tereshkin noslobodsky, Elnikovsky, Temnikovsky, registered 4.7 desman burrows per coast- Ardatovsky, Kochkurovsky (Report, 1982). line km at the reference station in the Sura The results of the surveys revealed 5.25 floodplain (Tereshkin, 1971). burrows per coastline km along the Vad In addition to the three stations set up from the mouth of the Yavas near the vil- by L.P. Borodin, there is another desman lage of Ozyorny to the mouth of the Tast, station in Mordovia. It was set up by N.K. 1.5 burrows per km along the Moksha in Shidlovskaya in Tengushevsky District in the protected area of the Mordovsky Na- 1961 and named “Krasny Yar”. It was lo- ture Reserve, 2.1 burrows per km along the cated in the vicinity of Krasny Yar, Staraya Alatyr from the village of Anyutino down- Kocheevka, and Vedenyapino villages. stream to the village of Redkodubiye, and This station was located 35–40 km from 2.8 burrows per km along the Sura in Koch- the Mordovsky Nature Reserve. In 1961, 19 kurovsky District. The authors noted that lakes were surveyed, 9 (Obshirki, Mochil- the desman’s settlement in Kochkurovsky ki, Zaresheno, Melkoe, Uksala, Ovechiy District was the most southerly point in the Poldnik, Muromka, Bolshoe Rassoshnoe, Sura floodplain within the borders of Mor- Maloe Rassoshnoe) of which were includ- dovia. The desman was not registered on ed in a control and registration point. The the territory of Bolshebereznikovsky and total coastline length of these lakes was 12 Dubensky Districts. In 1982, according to km. A total of 38 desman burrows were the extrapolation made, the total desman found on these lakes in the fall of 1961, number in Mordovia was one thousand which corresponded to 42 individuals individuals. In fact, 149 desman indi-

50 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia viduals were registered in lake burrows, that period. Only oral reports on desmans’ 64 of which were documented in Zubo- habitats from state hunting inspection vo-Polyansky District (Staryy Vad, Kru- specialists and foresters were received. gloe, Mokshayevo, Siyarka, Popovskoye, Therefore, we used the following two con- Narehrke, Lokseyevo, Kondraehrke, Uti- trol dates to estimate the distribution of noe lakes), 42 – in Kochkurovsky District the Russian desman within the territory (Malyy Kamysh, Maloye Martianovo, Bol- of Mordovia in the past: 1) the period of shoye Martianovo, Selevyarka, Stychnoye, L.P. Borodin’s studies before 1970, and 2) Cherepashnoye, Bolshoy Kamysh, Bol- the period of two-year registration work shoy Chadamas, Cheremukha, Poperech- by V.S. Vechkanov and others (Report, noye lakes), 25 – in Ardatovsky District 1982, 1983). The latest report on the des- (Kochalino, Terlyaka, Egadam, Mukorka, man in Mordovia was not available to a Kamyshovoe, Glubokoye, Bolotnoye, Ses- wide range of scientists who studied the ke lakes), and 18 – in Temnikovsky Dis- desman. Therefore, the subsequent pub- trict (Kanal, Krasnenkoe, Linevo, Lipovoe, lications contained information on the Rubezhnoe, Starica lakes). desman in Mordovia borrowed by other In 1983, V.S. Vechkanov et al. carried authors from the available works of L.P. out research on introducing the desman Borodin, in particular for the years of 1963 into two floodplain lakes Dolgoye and and 1970. V.S. Vechkanov’s data were in- Krugloye on the territory of the biologi- cluded in the monograph of G.V. Khakhin cal station of the Mordovian University in & A.A. Ivanov (1990); however, 1985 was Bolshebereznikovsky District. In total, ten indicated there, and the registration work desmans captured on the Staryy Vad and was carried out in 1982. This situation with Utinoe lakes in Zubovo-Polyansky District typing errors in the literature, lack of data were released. 5 desmans were released on and, in general, termination of the study of the lake of Dolgoye. 5 desmans were re- the desman was typical not only of Mor- leased on the lake of Krugloe. According dovia, but also of other regions of Russia. to the report, no desmans were observed As a result, A.S. Onufrenya and M.V. Onu- during registration in Kochkurovsky and frenya reported the reliable presence of the Elnikovsky Districts (Report, 1983). species only in 6 specially protected natu- In the autumn of 2001, the republic’s ral areas of Russia in 2007, and, notably, hunting administration performed work in the Mordovsky Nature Reserve was not six districts, including in the Moksha basin included (Onufrenya & Onufrenya, 2008a). within the territory of Temnikovsky and At one time, 4 desmans in specially pro- Krasnoslobodsky Districts. The abundance tected natural areas were organized on the of the desman was 2.7 and 0.6 burrows territory of the former USSR: Oka Nature per km, respectively. The extrapolation of Reserve, Khopyor Nature Reserve, Mor- the data performed by G.V. Khakhin to all dovsky Nature Reserve, and Klyazminsky floodplain lands revealed that the species Nature Reserve. The Klyazminsky Nature stock in Mordovia was about 500 animals Reserve closed. The Oka Nature Reserve (Khakhin, 2009), with the estimated number and Khopyor Nature Reserve are among of desmans in the Moksha floodplain with- these 6 specially protected natural areas in in the borders of Mordovia equal to 250 in- desman territories. dividuals (Onufrenya & Onufrenya, 2016). Our research on the desman in Mordo- Considerable time has passed since via has been conducted since 2009. First of L.P. Borodin’s active study of the Russian all, at the initial stage of research, we ex- desman in the republic, but there were tradited the spread of the species and the practically no studies on the habitats and state of the population. The purpose of registration of the desman population for the presented work is the analysis of the

51 A. Andreychev et al. past and modern distribution of the Rus- bution of precipitation across the territory sian desman. The data was obtained by us is not very diverse. The average long-term for over 10 years (2009–2018). The results value of evaporation is calculated to be in of our work had been collected for a long the range of 390–460 mm. time. The data were not published until Mordovia is located in the south-west- completion of research. Only brief sum- ern part of the Volga River basin. 47% of maries were published in the collections of the area is located in the basin of Sura and the regional Red Book (Andreychev et al., 53% in the basin of Moksha. Moksha is the 2009, 2010, 2012, 2014). main river in western Mordovia that flows in the meridional direction to the confluence of the Urkat River, where its direction Material and Methods changes to the sublatitudinal. The Mok- sha is the right tributary of the Oka River The Republic of Mordovia is located in the and empties into it. The channel is 60–90 centre of the European part of Russia. Its m wide and 5–8 m deep. The absolute wa- extreme points are defined by geographi- ter level is 89–126 m. The Moksha’s largest cal coordinates 42º 11’ – 46º 45’ E and 53º38’ tributaries in the territory of Mordovia are – 55º 11’ N (Figure 1). The maximum dis- the Vad, the Satis, the Urkat, the Sivin, and tance from west to east is 298 km and the the Issa. distance from north to south is 57 to 140 The Sura River flows along the south- km. The area of the republic is 26.2 thou- eastern border of Mordovia. The width of sands km2. Features of the geological struc- the channel varies from 100 to 150 m and ture of Mordovia are determined by its its depth is 10 m. It flows 0.3–1.0 m/s and location in the central part of the Russian its absolute water level is 90–115 m. The Platform and the north-western slopes of largest tributaries of the Sura include the the Volga Upland. In the western part of Bolshaya Ksha, the Bolshaya Sarka, the Pi- the Republic of Mordovia, the Volga Up- ana, the Menya, and the Alatyr. land reaches the Oka-Don Lowlands. The density of the river network in the The climate of the region is moderate- entire area is 0.23 km of river per km2. By ly continental with pronounced seasons the nature of intra-flow distribution, the throughout the year. The influx of direct republic’s rivers are of Eastern Europe- solar radiation in Mordovia varies from 5.0 an type. The water regime of the rivers is in December to 58.6 kJ/cm2 in June. The to- characterized by the presence of summer tal radiation throughout the year is 363.8 and winter seasons, spring floods and au- kJ/cm2; the radiation balance is 92.1 kJ/ tumn floods. The most high-water month cm2. The average annual air temperature is April. 3–15% of the annual runoff oc- varies from 3.5 to 4.0 °C. The average tem- curs during the low-flow period. The river perature of the coldest month (January) is length is used in the study as a criterion for in the range of -11.5...-12.3 °C. Tempera- classifying rivers. We validate this method ture drops down to -47 °C occur. The aver- because most reference books on surface age temperature of the warmest month, i.e. water resources are based on that classi- July, is in the range of +18.9...+19.8 °C. Ex- fication (Sokolov, 1964; Sokolovsky, 1968; treme temperatures in the summer reach 37 Yevstigneyev, 1990; Chalov, 1994). Accord- °C. The average annual precipitation in the ing to the classification, large rivers are riv- territory of Mordovia is 480 mm. Over the ers the length of which varies from 301 to course of observation lasting many years, 1,000 km. Medium rivers extend from 101 periods of high and low humidity were to 300 km. The length of small rivers rang- noted, ranging between the minimum and es from 26 to 100 km. The smallest rivers maximum values of 120–180 mm. Distri- are usually rivers with a length of up to

52 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

25 km. In the territory of Mordovia, there daylight hours by going through the coast- are about 1,520 watercourses with the total line of water bodies overland along the wa- length of 9,250 km. The major share (96%) ter edge on foot or by boat. At least 70% of of the river network of the republic falls the examined reservoirs were covered by on the smallest rivers. These constitute the survey. Surveys were conducted in dif- 65% of the total length of all rivers. Small ferent areas over different years from 2009 rivers make up 4% of the total and 22% of to 2018. We did not identify any stationary the length of all rivers. The share of me- water bodies, but on some lakes the repeat- dium-sized rivers (the Issa, the Sivin, the ed surveys were carried out several years Insar, the Partsa, the Vad, the Visha, and later. First of all, our task was to cover the the Alatyr) in the total length is negligible largest possible area to determine the dis- (0.5%). Two rivers – the Moksha and the tribution and population density of the Sura – are more than 500 km long (Water Russian desman in Mordovia. resources, 1999). Small rivers and streams The total number of desmans was cal- are distributed unevenly in the basins of culated by extrapolating the surveyed data large rivers. In the river Sura, there are 24 over the entire territory using the follow- small rivers and 286 very small rivers and ing formula: streams. 30 small rivers and 385 very small Lnm rivers and streams flow into the Moksha X = K (1) River. 100L1 It is typical of small rivers to have a 5 to 10 m wide channel, with extensions up where X = the absolute desman population to 35 m; for average rivers, from 25 to 50 size, K = the conversion factor (average m with extensions up to 70 m. Large rivers number of animals per burrow), L = the are 100 to 150 m wide and in the mouths, length of the coastline of all reservoirs in 300 m. Rivers are shallow and the prevail- the area; n = the number of shelters count- ing depths of small rivers are 0.4 to 1.2 m, ed during the survey; m = the length of the for average ones 1.5 to 2.0, and for large control coastline expressed as a percentage ones 2.5 to 3.5 m. In rundowns, the depth compared to that measured on the map; LI throughout is less than 0.5 m. The flow = the length of the surveyed coastline. rate varies from 0.2 m/s to 1.2 m/s in run- In areas inhabited by the desman only, downs (Water resources, 1999). the conversion factor had the following There are approximately 500 lakes in values: September – 1.86; October (before Mordovia, five of which are large, and the freeze-up) – 1.10, and October-November rest are medium- or small-sized. Almost all (clear ice period) – 0.68. In areas inhabited lakes are floodplain, located in the valleys by both desmans and muskrats, conversion of the main rivers of the region being typi- factors were calculated for September-Oc- cal habitats of the desman. tober (before freeze-up): 0.6 for desmans, Surveys of possible habitats of the Rus- and 1.0 for muskrats (Kudryashov, 1976; sian desman in the areas of the Republic of Onufrenya & Onufrenya, 2016). The hab- Mordovia have been conducted. Tradition- itat quality of lands was traditionally de- al guidelines on the study of the desman termined by the population density of the were used and taken into account during desman as follows: quality I – > 25 burrows the research (Kudryashov, 1976; Khakh- per km, quality II – 15 to 24 burrows per in & Ivanov, 1990; Marchenko, 2011). The km, quality III – 6 to 14 burrows per km, desman survey was carried out mainly by quality IV – 0.5 to 5 burrows per km, and calculating the inhabited desman burrows quality V – < 0.5 burrows per km. (Kudryashov, 1976; Onufrenya & Onufren- ya, 2016). Surveys were carried out during

53 A. Andreychev et al.

Figure 1. Past and present distribution of the Russian desman in Mordovia. Note: the dotted line indicates the current range of the species within the region, the black fill indicates the distribution area until 1970 (Borodin 1963, 1970). – Stationary observation points for ★ desmans set up by L.P. Borodin (1970); I.S. Tereshkin (1971); N.K. Shidlovskaya (2018). ■ – Desman registration points in 1982 (Report, 1982) to – Desman registration points according to the results of this research; the point numbering is given in the Table 1. ① ⑳

54 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

Results given by districts. In Tengushevsky Dis- trict, 38 lakes are habitable for desmans, Altogether, 174 desman burrows were with the total coastline length of 60 km found in the reservoirs surveyed. The (Figure 2). These lakes have a coastline abundance was 1.6 burrows per coastline length of mainly 1 to 2 km. Considering km. The conversion factors and the total the fact that the mean relative density is coastline length of the reservoirs where the 2.4 burrows per km on the lakes studied in research was carried out let us conclude Tengushevsky District, we can assume that that at least 588 individuals inhabited the 86 to 268 individuals inhabit the lakes in surveyed territory. It should be noted that this area. This is due to the fact that accord- in this calculation we were dealing with ing to the results of the work carried out by the minimum population size of the spe- A.S. Onufrenya & M.V. Onufrenya (2008b) cies since only desman burrows were taken on the territory of Meschera National Park, into account. Considering the fact that the not all reservoirs may be inhabited by the reservoirs where the desman’s presence desman. This fact was also confirmed by us was detected also had beaver’s burrows when different situations were observed in most cases, the desman numbers will on the neighboring lakes: there were both be much larger since the desman probably burrows and feeding boards on some uses beavers’ shelters, as well. However, lakes, and there were no desman traces in this work we did not take this circum- on others. In particular, this is character- stance into account, and considered only istic of the Telimerka and Takushevskoe desman burrows. In addition, the situation lake margins and reclamation canals near in the western Mordovia consists in the these lakes. The reclamation canal adja- fact that there are no traces of the musk- cent to Lake Telimerka (south-western rat registered on many lakes. Therefore, it part) was found to have desman burrows is logical to use higher conversion factors, and some evidence of feeding activity. September – 1.86; October (before freeze- 130 meters away from the lake on the ca- up) – 1.10, and October-November (clear nal approaching perpendicular to the lake, ice period) – 0.68. fresh traces of vital activity of the Russian To assess the total desman stock in the desman were found. The surveys of recla- water bodies of Mordovia, we carried out mation canals to the north and east of the an analysis of water bodies habitable for the village Standrovo in 2011 did not yield Russian desman using OziExplorer soft- positive results in relation to the desman. ware and satellite images. In this analysis, The search for traces of desmans’ presence we considered only those areas in which on the artificially straightened riverbed of the desman had been registered. The riv- the Yuzga to the east of Standrovo was also erbed coastline was analyzed separately. unsuccessful. The exploratory research on The riverbeds of the Nuluy (35.1 km) and the reclamation canals to the south of the Kundybolka (8.2 km) were studied along village Shelubey over the Yuzga revealed the entire length. The riverbeds of the Vad, the presence of the desman and burrows. Partsa, Yavas, and Vindrey were partially In two cases, traces of desman poaching studied along 110 km of the coastline. The were found with traps from nesting cham- mean value for the rivers was 0.9 burrows bers. Excavated beaver burrows were also per coastline km, which corresponded to found on this site. According to local res- quality class IV (0.5–5 burrows per km). idents, poaching of the Russian desman Based on the results obtained, we may is common in Tengushevsky District. The assume that about 400 animals live in the survey of the coastline of Lake Beloye lo- above-mentioned rivers. cated 2.8 km southeast of the village of The distribution of floodplain lakes is Shiromasovo on the Moksha left bank, on

55 A. Andreychev et al. the border of the Republic of Mordovia general, Krasnoslobodsky District should and Ryazan Region, did not reveal desman be recognized as less studied for the des- presence. The lake is supposedly of mete- man before we started the work. Perhaps orite impact origin; the shores of the reser- it was of less interest in Borodin’s days due voir are open, not bushy. to the distance from the Mordovsky Na- In Krasnoslobodsky District, 25 lakes ture Reserve, where L.P. Borodin worked are habitable for desmans; the total coast- or perhaps also due to fewer lakes in the line length of these lakes is 30 km (Fig- floodplain. However, the results -demon ure 3). The lakes have a coastline mainly strate that despite all the circumstances, length of less than 1 km. Considering that the territory of the district is of great im- the mean relative density is 8.9 burrows portance in preserving the desman, since per km on the lakes studied in Krasnoslo- the population density here is the highest. bodsky District, we can assume that 160 In particular, the highest density of the individuals inhabit the lakes of this area. desman population in Mordovia being 10.5 In Krasnoslobodsky District, as well as in burrows per coastline km was registered in Tengushevsky District, burrows and feed- 2009 on reclamation canals adjacent to the ing boards were often recorded on recla- northeastern part of Lake Churilka. mation canals rather than on the lakes, to In Temnikovsky District, 34 lakes are which they were adjacent. This situation habitable for desmans. The total coastline was common in the vicinity of Nagornoye length of these lakes is 62 km. These lakes Shenino and Novaya Karga villages. In mainly have a coastline length of about 2

Figure 2. Records of the Russian desman in Tengushevsky District. Note: water bodies are shown in white, – desman registration points. ★

56 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

Figure 3. Records of the Russian desman in Krasnoslobodsky District. Note: water bodies are shown in white, – desman registration points. ★ km. Considering the fact that the mean rel- Surveys at the first and second station ative density is 0.6 burrows per km on the set up by L.P. Borodin were successful lakes studied in Temnikovsky District, we (Figure 4, 5). The desman was still present can assume that 22 to 69 individuals inhab- in those areas after 57 years of setting the it the lakes in this area. We started our re- stations up. We surveyed floodplain lakes search on habitats in Temnikovsky District on the left bank of the Moksha to the west in 2009. According to S.K. Potapov’s oral of Temnikov Town (first station). The sur- report, he personally watched the desman veys were started from the mouth part of on Lake Valza. We investigated the east- the Kundybolka where according to N.N. ern part of Lake Valza. No vital activity Chuprunov, State Hunting Inspectorate of the desman was found. The next reser- specialist, the desman was previously voirs that were surveyed were two ponds seen. As a result of the river survey, we near Pushta Settlement. Studies on the also found desman burrows in the Europe- entire coastline of the ponds did not give an beaver’s burrows and passages, which any positive results in relation to the des- was consistent with the literature data on man. The southwestern part of the coast- the symbiotic relationship between the line of Lake Inorskoye and the coastline of desman and beaver (Barabash-Nikiforov, Lake Picherki were investigated. Feeding 1959). High population of the European boards and desman burrows were found beaver is favorable for the desman in Mor- on Lake Picherki. According to L.P. Boro- dovia in this respect (Andreychev, 2017). din, Lakes Valza and Inorskoye were the Then we studied the lakes to the north of main reservoirs in which the desman was the Kundybolka mouth. released (Borodin, 1963).

57 A. Andreychev et al.

Figure 4. Records of the Russian desman in Temnikovsky District. The first station. Note: water bodies are shown in white, – desman registration points. ★

Figure 5. Records of the Russian desman in Temnikovsky District. The second station. Note: water bodies are shown in white, – desman registration points. ★

58 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

The easternmost desman registration allows us to classify Elnikovsky District as points in Temnikovsky District are lakes in a low quality V area. the vicinity of Mitryaly and Nizhniye Bor- In Zubovo-Polyansky District, more ki villages. At the latter point in 2017, the than 100 lakes are habitable for desmans. desman was registered by E.N. Potapkin The total coastline length of these lakes (Mordovian Pedagogical Institute) visually is 110 km (Figure 6). These lakes mainly and by burrows. Registrations in the east- have a coastline length of less than 1 km. ern parts of Temnikovsky District and the Considering the fact that the mean relative information received from hunting inspec- density is 2.5 burrows per km on the lakes torate specialists in Elnikovsky District studied in Zubovo-Polyansky District, we suggested a widespread habitat of the spe- can assume that 165 to 512 individuals in- cies in the Moksha floodplain within the habit the lakes in this area. There is a stable territory of Elnikovsky District. However, desman settlement on Belyye Lakes, which our survey of the Nului River in the river are floodplain water bodies of the Vindrey mouth showed no evidence of the Russian River. The research revealed burrows and desman’s presence. Earlier in winter 2010, feeding boards of the desman. A large a desman was hunted (shot) by a hunt- number of burrows and feeding boards er on this river. This river is swift-flow- were found on the coastal line of the Yavas ing, the banks are sandy shale, sloughing River from the village of Ozyorny at the shale. Most likely the desman appeared mouth of the Vad. It should be mentioned here from the nearest floodplain lakes. The that poaching capture is carried out in the studies carried out by V.S. Vechkanov et al. rivers of Zubovo-Polyansky District with (Report, 1982) suggest low registration fre- the use of fixed nets and other fishing gear. quency for the desman in the area. All this Moreover, we carried out a search for the

Figure 6. Records of the Russian desman in Zubovo-Polyansky District. Note: water bodies are shown in white, – desman registration points. ★

59 A. Andreychev et al. desman on the Vysha within the area be- on one of the feeding boards. tween the villages of Crimea (Penza Re- The only area where the desman is gion) and Gorodische. The river banks are currently preserved in the eastern part steep, thickly grown with vegetation. No of Mordovia is Ardatovsky District (the signs of the desman were found. Alatyr basin) (Figure 7). Twenty lakes are On floodplain lakes of the Vad in the habitable for the desman in Ardatovsky area between the villages of Kiselyovka District, in the floodplain from the borders and Vadovo-Sosnovka, local fishermen of Ichalkovsky District to Redkodubye rarely observed cases of desman capture. A Village. The total coastline length of these survey on a part of the Udev River (south- lakes is 47 km. These lakes mainly have a ern part of Zubovo-Polyansky District) did coastline length of about 2 km. Consider- not give a positive result. The area sur- ing the fact that the mean relative density veyed represented a meandering section is 0.6 burrows per km on the lakes studied of the river thickly grown with shrubs, in Ardatovsky District and that the musk- and the banks were mostly flat. The flood- rat is common here, we can assume that 17 plain lakes at the time of the survey were individuals only inhabit the lakes in this covered with aquatic vegetation, no mir- area. ror-like surface observed, and the shores In 2010, we carried out a survey on the were also bushy. lakes at L.P. Borodin’s third station in the Torbeyevsky District, where the des- floodplain of the Sura (Figure 8). Unfortu- man was also discovered, adjoins Zubo- nately, it has been revealed that the des- vo-Polyansky district from the east. Habi- man does not currently inhabit these lakes. tats were located on floodplain lakes of the Old uninhabited burrows of the desman rivers Partsa and Vindrey there. Feeding were found on lakes Bolshoy Chadamas boards and desman burrows were found. and Bulyk only. Traces of the vital activ- Seventy-three gastropod shells were found ity of the previous desmanon these lakes

Figure 7. Records of the Russian desman in Ardatovsky District. Note: water bodies are shown in white, – desman registration points. ★ 60 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

Figure 8. Records of the Russian desman in Kochkurovsky District. Note: water bodies are shown in white. only testify to the fact that some factor con- desman. Also, no results were obtained tributed to the extinction of the desman. It during the examination of lake margins at should be noted that Lake Pendelyukha a biological station of the Mordovian Uni- has turned into an ephemeral swamp since versity in Bolshebereznikovsky District. the studies on L.P. Borodin’s stations. L.P. Borodin considered that the muskrats’ settlement in desman areas was the con- Discussion tributing factor. He stated that the grow- ing muskrat population leads to a dramatic Thus, to date, the number of the Russian decrease in the number or extinction of the desman in Mordovia determined actually desman within relationships of these two as a result of registration work is 588 indi- species (Borodin, 1970). It should be not- viduals (Table 1). According to estimates, ed that burrows and traces of the muskrat the total desman population is about 1,400 were found in the southern part of Lake individuals in Mordovia (Figure 9). As we Krivulka. The situation is similar on other can see from the desman population dy- lakes of Prisurye. namics diagram for the period from 1966 A survey on other lakes in the Prisurye to 2018, there is an increase in its popula- area that are not part of L.P. Borodin’s station. However, a detailed study of the mation, such as Lake Selevyarka (4 km south terials provided by us demonstrates that of Lake Inerka) in Kochkurovsky District, this is not the case. The fact is that the data Lake Makarikha (8 km south of Engalyche- for 1966 are given only according to the vo Village) in Dubensky District, did not data of the well-known stations set up by yield positive results with regard to the L.P. Borodin, and this is a small territory

61 A. Andreychev et al. in Temnikovsky and Kochkurovsky Dis- already known that desman trapping ex- tricts. No data was available on the total clusively for fur production amounted to species population either for these districts 2,493 individuals in the period from 1953 or for other districts. In 1982, V.S. Vech- to 1956 in Mordovia. It should be noted kanov presented data for several districts that the largest quantity of desman skins by extrapolating registration data obtained in Mordovia was stocked in Zubovo-Poly- on areas in four districts. In fact, 149 des- ansky, Temnikovsky and Tengushevsky man individuals were registered during Districts (71%), that is, the Moksha basin. fi eldwork in burrows on lakes. We may It is noteworthy that the situation has not conclude that to date the population of the changed and these areas are currently the species has not increased, but the number main ones in desman conservation in the of known desman habitats has increased region. However, the situation is not clear due to surveys of wide territories in the in Krasnoslobodsky District where the des- districts during expeditionary studies in man is also favorably distributed through- Mordovia. Most scientists see a tendency out the reservoirs. And at the time of L.P. of a dramatic decrease in the species pop- Borodin’s work there was no information ulation in Russia (Onufrenya & Onufren- on the district at all, not only on fur pro- ya, 2016; Rutovskaya et al., 2017b). We can duction (stock), but also in general. The only guess the desman population in the desman probably settled extensively in the past, but clearly, it was several times high- fl oodplain of Krasnoslobodsky District af- er than now. Because during the studies ter 1970. performed by L.P. Borodin (1970) it was

1 Figure2 9. Changes in the number of Mordovian population of the Russian desman during the last 50 years. Data for 1966 – according to Borodin (1970). Data for 1982 – according to Vechkan- ov et al. (1982). Data for 2018 – according to our data for 2009–2018.

62 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia

Table 1. Relative numbers of desmans and their total reserve by results of surveys in 2009–2018.

Year Point District Name of the reservoir or The total Population Desman num- the nearest settlement length of the density, reserve, ber (georeferencing) coastline, burrows/km indi km (quality class) viduals 2009 1 Temnikovsky Lake Picherki 2.1 1.1 (IV) 4 2 Temnikovsky Lake Tucherki 1.7 <0.5 (V) 2 3 Temnikovsky River Kundybolka and Lakes 18.1 8.2 (III) 89 Zhegalovo, Mochalnoe, Bolshoe Palkino, Maloe Palkino 4 Krasnoslobodsky Reclamation canal northwest of 2.3 7.1 (III) 30 the village Nagornoye Shenino 5 Krasnoslobodsky Reclamation canal at Lake 2.2 10.5 (III) 43 Ramenskoye 6 Torbeyevsky River Vindrey and Lake near the 58.1 0.8 (IV) 20 village Windray 7 Zubovo-Polyansky Lake Belye 2.2 4.4 (IV) 18 8 Zubovo-Polyansky River Yavas from the village 75.3 2.5 (IV) 79 of Ozerny to the mouth at the confluence of River Vad 9 Ardatovsky Lake Parma 4.6 0.6 (IV) 5 2010 10 Krasnoslobodsky Lake Churilka and reclamation 4.4 9.3 (III) 76 canals 2011 11 Elnikovsky River Nuluy 35,1 <0.5 (V) 8 12 Tengushevsky Lake Telimerki and reclamation 7.2 2.2 (IV) 30 canals 13 Tengushevsky Reclamation canals northeast of 10.1 2.2 (IV) 41 the village Standrovo 14 Zubovo-Polyansky River Partsa near the village 92.4 <0.5 (V) 60 Romanovka 15 Zubovo-Polyansky River Vad 114.1 <0.5 (V) 74 16 Zubovo-Polyansky River Partsa between the villages 92.4 <0.5 (V) 60 of Romanovka and Partsa 2012 7 Zubovo-Polyansky Lake Belye 2.2 6.1 (III) 18 2013 17 Temnikovsky Lake near the village Mitryaly 2.5 <0.5 (V) 2 2014 10 Krasnoslobodsky Lake Churilka and reclamation 4.4 10.1 (III) 76 canals 2015 6 Torbeyevsky River Vindrey and Lakes near the 58.1 0.6 (IV) 20 village Vindrey 2017 18 Zubovo-Polyansky River Partsa between the villages 92.4 <0.5 (V) 60- of Romanovka and Vasilevka 19 Temnikovsky Lake north of the village 2.1 <0.5 (V) 2 Nizhniye Borki 2018 12 Tengushevsky Lake Telimerki and reclamation 7.2 2.2 (IV) 30 canals 13 Tengushevsky Reclamation canals northeast of 10.1 3.1 (IV) 41 the village Standrovo 20 Tengushevsky Lake Bolshoye, south of the 4.1 2.1 (IV) 5 village Kuraevo On average 1.6 Total 588

63 A. Andreychev et al.

We may assume that the main desman individuals) and low population density. population is concentrated on floodplain The registration point for the desman near- lakes and reclamation canals. About 400 est to Lake Parma (Ardatovsky District) animals may live on riverbeds of the in Mordovia is Lake Staritsa south-west Vad, Partsa, Yavas, Vindrey, Nuluy, and of the village Atrat (Alatyrsky District) in Kundybolka. The distribution of habitats Chuvashia. The distance between registra- across river basins is extremely uneven. tion points in a straight line is 52 km, the The desman majority is in the Moksha ba- distance along the riverbed is 93 km, with sin (more than 1,350 individuals), and the 81 km along the Alatyr riverbed and 12 km Alatyr basin (left tributary of the Sura) has along the Sura riverbed. We cannot rule about twenty individuals. out the possibility of subsequent registra- We cannot consider the Elnikovsky tion of the desman at intermediate points, floodplain of the Moksha as the key one for example at the Alatyr mouth. A small in conservation of the desman at present, group of individuals could not survive in in comparison with Tengushevsky, Tem- isolation for several decades in the flood- nikovsky and Krasnoslobodsky floodplain of the Alatyr on Lake Parma. The plains. The desman population density latter assumption is justified since desman in Torbeyevsky District is way below the habitats are scattered under Mordovian density in Zubovo-Polyansky District. conditions in the Moksha and Alatyr ba- Therefore, we do not consider Torbe- sins. The distance between Lake Parma yevsky District as an important one with (Ardatovsky District) and the easternmost regard to desman conservation. The situ- registration point in the Moksha basin ation with the desman in Ardatovsky Dis- (Lake Ramenskoye in Krasnoslobodsky trict is worse than in other districts where District) is 132 km in a straight line. the desman was detected. The Russian desman is not preserved in The study results demonstrate a frus- the upper reaches of the Sura in Penza Re- trating situation with the desman in Mor- gion (Ilyin, 2003). Thus, the probability of dovian Prisurye. However, there is rea- finding the desman is extremely low up to son for optimism due to the fact that the the Alatyr mouth on the Sura. In Penza Re- desman has persevered at present in the gion, the desman was recorded only in the lower reach of the Sura according to our Don basin on two steppe rivers in 2002: the colleagues’ registration work in 2016. Ac- Syuvernya River in Belinsky District near cording to M.V. Rutovskaya et al., (2017a) Lopatino Village and the Serdoba River in the total number of the Russian desman Maloserdobinsky District near Ogaryovka was more than 500 individuals in the Sura Village. There were two desman settle- basin in Alatyrsky, Poretsky, Shumer- ments per 1 km of the riverbed. linsky and Yadrinsky Districts of Chu- A somewhat better situation with the vashia and Pilnensky Districts of Nizhny population density of the Russian desman Novgorod Region. The average relative in comparison with other neighboring re- density of this population was 2.4 holes at gions of Mordovia is observed in Ryazan 1 km of the coastline. It is noteworthy that and Vladimir Region (Onufrenya & Onu- the desman was registered by research- frenya, 1992, 2005; Okulova et al., 2008; ers on the Sura in areas after meeting the Rutovskaya et al., 2015). The surveys con- Alatyr (Alatyrsky District), that is, taking ducted by A.S. Onufrenya and M.V. Onu- into account the last registration sites in frenya in 2008–2009 in the floodplain lands Mordovia in the floodplain of the Alatyr, in the middle reaches of the Oka (Spassky the southernmost population group of the and Shilovsky Districts) allow to conclude species is located in the Sura basin. This that the habitats of the desman are natural- explains such a low population (about 20 ly high-quality (class I and II). The current

64 Distribution and population density of the Russian desman (Desmana moschata L., Talpidae, Insectivora) in the Middle Volga of Russia state of the desman areas in most of the frenya (2012), the desman lives in Ryazan territory surveyed by the authors corre- Region (Kadomsky and Shatsky Districts) sponds, at best, to class IV or even class V, areas bordering Mordovia, thus forming a the lowest quality class. The relative num- continuous range of the species with tran- ber of the desman population for the entire sition to Tengushevsky and Zubovo-Poly- floodplain expansion averages 3.9 shelters ansky Districts of the republic. per coastline km. At the same time, this In the material presented in this article, number varies from 10.9 to 6.2 shelters per we tried to fill in the missing information coastline km in permanently protected or on the Russian desman population densi- inaccessible areas, and does not exceed 1.0 ty in Mordovia, so that there would be no in other areas. The total desman popula- misconception or typing errors in the liter- tion within the borders of Spassky and Shi- ature regarding the small number of regis- lovsky Districts is 1,800 to 2,000 individu- tration points in the region. These data are als, with about 80% of the total stock being important for assessing the entire range of within the territory of the protected area the Russian desman. It is no coincidence of the Oka Nature Reserve and the “Yer- that C. Nores et al. (1998) in their work on akhtursky” hunting reserve (Onufrenya & the Pyrenean desman (Galemys pyrenaicus Onufrenya, 2012). According to data avail- Geoffroy, E.), whose range is many times able for Ryazan Region, the desman pop- smaller than the range of the Russian ulation is 5,960 individuals (Onufrenya & desman, report that, despite studying for Onufrenya, 2016), which is several times several decades (Stone & Gorman, 1985; higher than the desman population (1,400) Stone, 1987), information on the density of in Mordovia. the population of the Pyrenean desman is According to some latest data, the extremely insufficient. This conclusion is Russian desman population in Russia is also valid for the Russian desman, where estimated at 8,000 to 10,000 individuals information is collected by researchers for (Rutovskaya et al., 2017b); 6,000 individ- each individual region. According to the uals (Rutovskaya et al., 2014); 4,000 indi- results obtained by foreign colleagues, the viduals (Onufrenya et al., 2011), 25,000 population density of the Pyrenean des- individuals (Khakhin, 2009). According to man of 2.8–7.3 individuals per km is com- A.S. Onufrenya & M.V. Onufrenya (2016), parable to the population density of the the desman population in eight regions Russian desman. (Oryol, Kaluga, Ryazan, Vladimir, Iva- novo, Penza, Nizhny Novgorod regions Acknowledgements. We are grateful to and the Republic of Mordovia) of the Oka G.F. Grishutkin, G.V. Shkolov, E.A. Lo- basin is 4,110 individuals. G.V. Khakhin bachov, S.N. Spiridonov, S.V. Susarev, V.I. and A.A. Ivanov (1990) give old numbers Astradamov, S.G. Erofeev, E.N. Potapkin for Mordovia – 250 individuals. Based on for their support in carrying out field stud- the data we obtained for Mordovia (plus ies. 1,100 individuals), the number of 5,210 individuals for the Oka basin should be considered. The Moksha basin (Tengu- References shevsky, Temnikovsky, Zubovo-Polyan- sky and Krasnoslobodsky Districts) should Andreychev, A. 2017. Population density of the be taken into account since the desman is Eurasian beaver (Castor fiber L.) (Castoridae, Rodentia) in the Middle Volga of Russia. indigenous to the lakes and rivers here, de- – Forestry Studies | Metsanduslikud spite the decrease in its number since L.P. Uurimused, 67(1), 109–115. Borodin’s studies in Mordovia. Moreover, according to A.S. Onufrenya & M.V. Onu-

65 A. Andreychev et al.

Andreychev, A.V., Kuznetsov, V.A., Lapshin, Borodin, L.P. 1970. The desman in floodplains A.S. Grishutkin, G.F. 2009. Preliminary of the Moksha and Sura Rivers. (Выхухоль в materials about distribution of Russian поймах рек Мокши и Суры). Works of the desman (Desmana moschata L.) in the Republic Mordovian State Reserve of P.G. Smidovich. of Mordovia. (Предварительные материалы Saransk, Mordovian Book Publishing House, по распространению русской выхухоли 5, 61–90. (In Russian). (Desmana moschata L.) в Республике Borodina, M.N., Borodin, L.P., Tereshkin, I.S., Мордовия). Rare Animals of the Republic of Shtarev, Yu.F. 1970. Mammals of the Mordovia Mordovia: Materials for the Red Data Book of Reserve. (Млекопитающие Мордовского the Republic of Mordovia for 2009. Saransk, заповедника). Works of the Mordovian Izdatel’stvo Mordovskogo Universiteta, 4–7. State Reserve of P.G. Smidovich. Saransk, (In Russian). Mordovian Book Publishing House, 5, 5–60. Andreychev, A.V., Kuznetsov, V.A., Lapshin, (In Russian). A.S. 2010. On the problems and prospects of Chalov, R.S. 1994. Of riverbed mode of the rivers preserving the Russian desman in specially of northern Eurasia. (Русловой режим рек protected natural territories of Mordovia. (О Северной Евразии). Moscow, Izdatelstvo проблемах и перспективах сохранения Moskovskogo Universiteta. 336 pp. (In русской выхухоли на особо охраняемых Russian). природных территориях Мордовии). Ilyin, V.Yu. 2003. Materials for the spread of – Problems of the Rational use of Natural desman (Desmana moschata L.) in the Penza Resources and Environmental Protection: region. (Материалы к распространению Materials of the International Scientific- выхухоли (Desmana moschata L.) в Пензенской Practical Conference. Makhachkala, 389–390. области). – Protection of flora and fauna of the (In Russian). Volga region and adjacent territories. Penza, Andreychev, A.V., Kuznetsov, V.A., Lapshin, A.S. 69–71. (In Russian). 2012. Results of inspection of possible habitats Kennerley, R., Turvey, S.T. 2016. Desmana of Russian desman (Desmana moschata L.) in moschata. – The IUCN Red List of Threatened the western part of Mordovia. (Результаты Species 2016: e.T6506A22321477. 11 pp. обследования возможных местообитаний Available online: http:// dx.doi.org/10.2305/ русской выхухоли (Desmana moschata L.) в IUCN.UK.2016-2.RLTS.T6506A22321477.en. западной части Мордовии). Rare Animals [Downloaded 22 April 2017]. of the Republic of Mordovia: Materials for the Khakhin, G.V. 2009. Russian desman in danger: Red Data Book of the Republic of Mordovia dynamics of numbers and problem of for 2011. Saransk, Izdatel’stvo Mordovskogo protection. (Русская выхухоль в опасности: Universiteta, 3–5. (In Russian). динамика численности и проблемы Andreychev, A.V., Lapshin, A.S., Kuznetsov, V.A., охраны). Moscow, Publishing House of the Lobachov E.A., Shkolov G.V. 2014. Information Biodiversity Conservation Centre. 104 pp. (In on the registration of rare mammals of Russian). the Republic of Mordovia. (Сведения о Khakhin, G.V., Ivanov, A.A. 1990. Desman. регистрации редких видов млекопитающих (Выхухоль). Moscow, Agropromizdat. 191pp. в Республике Мордовия). Rare Animals of (In Russian). the Republic of Mordovia: Materials for the Kudryashov, V.S. 1976. Methodical instructions on Red Data Book of the Republic of Mordovia account desmans and muskrats in inundated for 2014. Saransk, Izdatel’stvo Mordovskogo grounds. (Методические указания по учету Universiteta, 4–5. (In Russian). выхухоли и ондатры в пойменных угодьях). Anonymus. 1936. Desman. (Выхухоль). Moscow, Moscow, Kolos. 10 pp. (In Russian). Gudok. 169 pp. (In Russian). Kufeld, N.S. 1939. Reacclimatization of Desman in Barabash-Nikiforov, I.I. 1959. Symbiotic links the Kuibyshev Region. (Реакклиматизация of the population of the beaver’s burrow. – выхухоли в Куйбышевской области). Zoologicheskii Zhurnal, 38(5), 767–771. (In – Scientific Methodological Notes of the Russian with English summary). Commission on the Reserve. Moscow. 3 pp. Bogdanov, M.N. 1871. Birds and animals of the (In Russian). black earth strip of the Volga region and valleys Marchenko N.F. 2011. Methodical peculiaries of the Middle and Lower Volga. (Птицы of Russian desman counts. (Методические и звери черноземной полосы Поволжья особенности учёта выхухоли русской). – и долины Средней и Нижней Волги). – Povoljskii Ecological Journal, 1, 97–102. (In Proceedings of the Society of Naturalists of the Russian). Imperial Kazan University. Kazan, V. 1. 229 Nores, C., Ojeda, F., Ruano, A., Villate, I., González, pp. (In Russian). J., Cano, J.M., Garcia, E. 1998. Estimating the Borodin, L.P. 1963. Russian desman. (Русская population density of Galemys pyrenaicus in выхухоль). Saransk, Mordovian Book four Spanish rivers. – Journal of Zoology, 246, Publishing House. 301 pp. (In Russian). 454–457.

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Received November 22, 2019, revised December 6, 2019, accepted December 29, 2019