Kutenkov A. Spatial-ecological divergence of the common frog (Rana temporaria L.) and the moor frog (Rana arvalis Nilss.) within their geographic ranges // Principy èkologii. 2017. № 1. P. 4‒51. DOI: 10.15393/j1.art.2017.5065

Issue № 1

Analytical review


Spatial-ecological divergence of the common frog (Rana temporaria L.) and the moor frog (Rana arvalis Nilss.) within their geographic ranges

Ph.D., Reservation "Kivatch", stapesy@mail.ru
Rana temporaria
Rana arvalis
winter shelters
breeding habitats
summer habitats
vertical distribution
habitable landscapes
Summary: R. temporaria and R. arvalis are widespread species, and their sympatry zone extends for some 4000 kilometers from west to east and in the widest part along meridian 33º east longitude about 2000 kilometers. The distinction in the physiology of hibernation and the nature of their winter shelters between R. temporaria and R. arvalis is principial. The common frog, as a kind of a potential water-hibernator, can remain in the terrestrial shelters where a local climate permits. Moor frog which is capable to cope with body freezing is a potential land hibernator, but certain situations make it spend winter under water. Both species can use bogs as wintering shelters. Strategy of R. temporaria spawning was formed in the conditions of small river basins not exposed to vast floods, with the accompanying streams-tributaries, inlets, flow-through lakes, oxbows, potholes, etc. Breeding behavior of frogs consists in forming a «mat» or «cushion» of the spawn clumps, which prevent their drifting towards deep water. In contrast, the «core» breeding habitat of R. arvalis is flat lowland areas with a non-static level of shallow water. Scattered oviposition of this frog significantly increases the chances of larvae to complete their development in isolated pools and ponds remaining after drying shallow water areas. The congestion of clutches can occur only at a high density of breeding specimens of the moor frog in some circumstances. In fact, this is only a reduction to the minimum the distance between males, but not a desire of animals to aggregate. In favorable conditions and living side by side, R. temporaria and R. arvalis have always some extent overlap of spawning habitats, and they can spawn close to each other. Common frog is a mobile and terrestrial species. Therefore it occupies the wide range of summer foraging habitats in the area having a certain landscape. Usually, this is a variety of forest habitats, scrubs and nearby meadow areas, rocky tundra, and marshy runoff beds covered with shrubbery or uneven scrub mires. Summer habitats of R. arvalis are primarily open or semi-open landscape faces, often soggy. In European mountains, R. temporaria is known to occur up to 2750 m above sea level, and in the south it does not come down to the lowlands. R. arvalis usually avoids areas located above 500–600 m, and they are rarely found at the height exceeding 900 m a. s. l. However, in the mountains of Southern Siberia populations of this species are common at altitudes up to 1800 m a. s. l., and the documented upper limit is 2400 m a. s. l. The causes of such differences in vertical distribution of the species are fundamental differences in the geomorphological properties of the landscape of mountainous regions of Europe and Asia. All european mountains (Pyrenees, Alps, Carpathians, Scandinavia and the mountains of the Balkan Peninsula), including the Urals, are mostly folded structures that arose in the process of deformation (folds and thrust faults) of the Earth crust in its horizontal motions. A less important role play blocky and arched uplifts, which are expressed only in the relief of midlands and lowlands of Central Europe (Slate Mountains, Schwarzwald, Bohemian Massif, etc.), stretching between the Central European plain and the systems of the Alps and Western Carpathians. Mountain systems are linearly elongated, dissected by weakly developed river valleys, which are narrow and steep-sided. These mountains do not have any large horizontal troughs, and the width of sloping valleys is less than a few kilometers everywhere. The bottoms of the valleys are seldom flat over large areas. Midlands and lowlands have the character of erosion dissected plateaus with narrow canyons. In contrast to the mountain areas in Europe, in Siberia we have to deal with the compound such as single mountain zone countries (Altai, Sayany, Baikal, Aldan). In the formation of the belt which stretches almost 3.500 kilometers arched and arched-block lifting plays the main role. In this case, the vast areas of the Earth crust experienced bending, followed by block division. Raised relict surfaces of alignment occupy areas on the flat hilly plains, uplands and tablelands, and form the bottom of the intermountain basins. Numerous valleys and hollows of South Siberian mountain belt sometimes so extensive, that separated highlands, ridges, river valleys with alluvium exist within them. Populations of R. temporaria in the European mountains are confined to the wooded valleys with rivers and streams running through them, to the shores of lakes in all altitudinal belts. In such circumstances, R. arvalis is deprived of its «core» landscape — flat marshy areas with stagnant water that impedes the penetration of this species to a mountainous terrain. In the mountains of Southern Siberia, uplifted hilly plains and plateaus are abundantly watered due to climatic conditions and water-physical properties of crumbly sediments. They are favorable for wide distribution of diverse bogs. Viable populations of R. arvalis inhabit all the altitudinal belts of mountain areas in Siberia where there are spacious and level surfaces: from waterlogged floodplains (Vitim river) to alpine tundra (Altai Mts.) The distribution of R. temporaria, the European species, everywhere in the area is associated with the landscapes, which are characterized by a pronounced micro- or mesorelief. The presence of small permanent and temporary streams and lakes with renewed water is necessary. Various small ponds usually present as well. General features of the relief does not permit significant overflows and floods of rivers and lake-river systems (areas affected by extensive flooding are avoided by the species categorically). Small marshes of eutrophic-mesotrophic series and the areas of wetlands are often present (in vast marshy terrains this frog does not occur). Spawning takes place in different riparian ponds, as well as in closed, relatively deep, temporary and permanent water bodies with similar hydrological regime, sometimes at a large distance from their hibernation sites. Obvious «outposts» of R. temporaria distribution are mountains. Of geographic range for some 15 thousand kilometers, if it is outlined on the extreme points of findings this species on the mainland, one half falls on the mountain massifs. As for the habitats of R. arvalis, the Europaean and Siberian species, a fundamentally different morphology of the landscape is inherent. This is a kind of flat and concave mesoforms and hydromorphic areas of the smooth relief with stagnant ponds. In any terrain, species demonstrates stronger preference for marshy landscape tracts, flooded areas of river valleys, reed quaking bogs, raw moorlands, and other wetlands. In different geographical conditions, such tract may be narrowed to the rim along the shores of lakes and rivers drying up in the steppe zone and in coastal meadows in northern Europe, or it dominates in the landscape either in Western Siberia or on the flat tundra in the North. In the landscape tracts inhabited by this species, tree or shrub vegetation rarely presents. The appearance of moor frogs in some «dry» forested habitats is probably caused by the presence of marshy areas nearby. Wintering, spawning and fattening of R. arvalis often occur in the same biotope and the year around. It is only where there is the interpenetration of «core» landscapes of each species (large areas of the northern and western parts of the East European Plain, and a part of the West Siberian Plain contiguous to the Urals), one can observe the examples of real sympatry of R. arvalis and R. temporaria.

© Petrozavodsk State University

Reviewer: G. Lada
Reviewer: V. Ishchenko
Received on: 30 March 2016
Published on: 01 April 2017


Avramova O. S. Bulahov V. L. Bobylev Yu. P. Mating behavior and «pairing» systems of the lake frog and the moor frog, Gruppovoe povedenie zhivotnyh: Doklady uchastnikov II Vsesoyuzn. konf. po povedeniyu zhivotnyh. M.: Nauka, 1976. P. 6–8.

Ammon P. L. List of amphibians and reptiles of Tula district, Zhurn. Tul'skiy kray. Tula, 1928. P. 44–52.

Amphibians of Red data book of Bashkortostan. The Common frog, V. F. Habibullin. URL: http://redbook.ru/article80.html (data obrascheniya 22.03.2014).

Anufriev V. M. Bobrecov A. V. Amphibians and reptiles. Fauna of the Europaean North-East of Russia. T. IV. SPb.: Nauka, 1996. 131 p.

The Baykalo-Lensky Reserve. URL: http://irkipedia.ru/content/baykalo_lenskiy_zapovednik (data obrascheniya 12.12.2014).

Bannikov A. G. Ecological conditions of hibernation of the common frog in Moscow region, Sbornik nauchnyh studencheskih rabot MGU. Zoologiya. M., 1940. Vyp. XVI. P. 41–64.

Bannikov A. G. Denisova M. N. Studies on amphibians' biology. M.: Uchpedgiz, 1956. 168 p.

Barabash I. I. Review of a stationary distribution of vertebrates in the Kadadinsky experimental forest district of Penza province, Byulleten' Voronezhskogo obschestva estestvoispytateley. Voronezh, 1939. T. 3. Vyp. 2. P. 21–29.

Belik V. P. On amphibian fauna and ecology in the steppe part of the Don basin, Sovremennaya gerpetologiya. 2010. T. 10. Vyp. 3/4. P. 89–100.

Belimov G. T. Sedalischev V. T. To the biology of Rana arvalis in Yakutia, Ekologiya. 1979. No. 5. P. 92–95.

Belgium — Mining encyclopaedia. URL: http://www.mining-enc.ru/b/belgiya (data obrascheniya 19.04.2015).

Blinova T. K. Amphibians of the northern forest-steppe of Zaural'ye, Vid i ego produktivnost' v areale: Materialy 4-go Vsesoyuznogo soveschaniya. Ch. V. Voprosy gerpetologii. Sverdlovsk, 1984. P. 5–6.

Borkin L. Ya. Belimov G. T. Sedalischev V. T. On distribution of frogs of the genus Rana in Yakutia, Gerpetologicheskie issledovaniya v Sibiri i na Dal'nem Vostoke. L.: ZIN AN SSSR, 1981. P. 18–24.

Brysova L. P. Kozhevnikova R. K. Korotkov I. A. Krinickiy V. V. Physico-geographic conditions of the north-east Altai and tasks of its nature investigation, Trudy Altayskogo gosudarstvennogo zapovednika. Gorno-Altaysk, 1961. Vyp. 4. P. 3–32.

Vartapetov L. G. Anufriev V. M. Moor frog in the northern taiga of Pritazovie, Vid i ego produktivnost' v areale: Materialy 4-go Vsesoyuznogo soveschaniya. Ch. V. Voprosy gerpetologii. Sverdlovsk, 1984. P. 7–8.

Visimsky Reserve. URL: http://vitimskiy.ru/index.php/sovremennoe-sostoyanie-ekosistem (data obrascheniya 12.12.2014).

Vozniychuk O. P. Kuranova V. N. Amphibians and reptiles of Katun nature reserve and adjacent territories (the Central Altai), Sovremennaya gerpetologiya. 2008. T. 8. Vyp. 2. P. 101–117.

Voronezhsky Reserve. URL: http://zapovednik-vrn.ru/o-zapovednike1/priroda/ (data obrascheniya 17.02.2016).

Garanin V. I. Amphibians and reptiles of Volga-Kama district. M.: Nauka, 1983. 175 p.

Gvozdeckiy N. A. Golubchikov Yu. N. Mountains. M.: Mysl', 1987. 399 p.

Geography of Hungary. http://www.gecont.ru/articles/geo/hungury.htm (data obrascheniya 22.03.2014).

Geography of Voronezh province. URL: https://ru.wikipedia.org/wiki/Geografiya _Voronezhskoy_oblasti (data obrascheniya 17.02.2016).

Geography of the Nederlands. URL: http://www.gecont.ru/articles/geo/holland.htm (data obrascheniya 22.03.2014).

Geography of Romania. URL: http://www.gecont.ru/articles/geo/romania.htm (data obrascheniya 22.03.2014).

Geography of Slovenia. URL: http://www.gecont.ru/articles/geo/slovenia.htm (data obrascheniya 19.04.2015).

Geography of Croatia. URL: https://ru.wikipedia.org/wiki/Geografiya Horvatii (data obrascheniya 19.04.2015).

Glazov M. V. On the role of Rana terrestris in controlling the abundance of invertebrates in the biocenoses of oak groves, Byulleten' MOIP. Otd. biol. 1975. T. 80. No. 6. P. 59–66.

Gul' I. R. Amphibians and reptiles of the Chernogora ridge, Ukrainian Carpathians, Voprosy gerpetologii: Materialy I s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Puschino). M., 2001. P. 74–75.

Gumilevskiy B. A. On  some ecology-faunistical investigations in Valday Upland, Izvestiya Vsesoyuznogo Geograficheskogo obschestva. 1941. T. 73. Vyp. 1. P. 129–136.

Darvinsky Reserve. URL: http://www.darvinskiy.ru/ecosistem.php (data obrascheniya 21.12.2014).

Dinesman L. G. Amphibians and reptiles of the south-east of Turgay table-country and Northern Priaralje, Trudy IG AN SSSR. Vyp. 54. M., 1953. P. 383–422.

Danube. URL: https://ru.wikipedia.org/wiki/Dunay (data obrascheniya 19.04.2015).

Eliseeva V. I. Fauna of the lower terrestrial vertebrates in Central Black earth Reserve, Trudy Central'no-Chernozemnogo zapovednika. Vyp. 10. M.: Lesnaya promyshlennost', 1967. P. 83–87.

Es'kov K. Yu. Striking paleontology. M.: ENAS, 2008. 312 p.

Azas Reserve. URL: http://green-azas.ru/zapovednik-azas/fiziko-geograficheskaya-kharakteristika.html (data obrascheniya 12.12.2014).

The Belogorie Reserve. URL: http://www.zapovednik-belogorye.ru (data obrascheniya 09.02.2016).

The Kuznetsky Alatau Reserve. Kemerovo, 1999. 255 p.

Zapovednik «Kuzneckiy Alatau». Fauna. URL: http://www.kuz-alatau.ru/priroda/fauna/#midl (data obrascheniya 12.12.2014).

The Tungussky Reserve. URL: http://www.tunzap.ru (data obrascheniya 12.12.2014).

Ivanter E. V. Survey data on ecology of the common frog in the North, Trudy zapovednika «Kivach». Petrozavodsk, 1969. Vyp. 1. P. 136–145.

Inozemcev A. A. Significance of insectivore birds in forest ecosystems. L.: LGU, 1978. 264 p.

Iskakova K. Amphibians of Kazakhstan. Alma-Ata, 1959. 92 p.

Ischenko V. G. Long-term studies of demography of amphibian populations: contemporary problems and methods, Voprosy gerpetologii: Materialy III s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Puschino-na-Oke, 9–13 oktyabrya 2006 g.). SPb., 2008. P. 151–169.

Ischenko V. G. Ledencov A. V. Influence of environmental conditions on the age structure dynamics of the moor frog, Vliyanie usloviy sredy na dinamiku struktury i chislennosti populyaciy zhivotnyh. Sverdlovsk: UNC AN SSSR, 1987. P. 40–51.

Kaleckaya M. L. Fauna of amphibians and reptiles of Darvinsky Reserve and their variations influenced by Rybinskoe reservoir, Rybinskoe vodohranilische. Ch. 1. M., 1953. P. 171–186.

The Katun Nature Biosphere reserve. http://www.turistka.ru/altai/info.php?ob=1202 (data obrascheniya 14.10.2015).

Kac N. Ya. Mires of the world. M.: Nauka, 1971. 296 p.

Kireev D. M. Ecological and geographical terms in forestry. Novosibirsk: Nauka, 1984. 182 p.

Kondrashev S. L. Gnyubkin V. F. Dimentman A. M. Orlov O. Yu. Role of visual stimuli in the breeding behaviour of Rana temporaria, Bufo bufo and Bufo viridis, Zoologicheskiy zhurnal. 1976. T. 55. Vyp. 7. P. 1027–1037.

Korneeva T. M. Bykov A. V. Rechan S. P. Terrestrial vertebrates of the lower part of Onega river. M.: Nauka, 1984. 89 p.

Korosov A. V. Ecology of the Vipera berus in the North. Petrozavodsk, 2010. 264 p.

Korosov A. V. Fomichev S. N. Key factors of amphibians distribution on the islands of the Kizhi archipelago, 10 let ekologicheskomu monitoringu muzeya-zapovednika «Kizhi». Petrozavodsk, 2005. P. 120–126.

Korosov A. V. Fomichev S. N. Spatial distribution of brown frogs during breeding period, Ekologiya. Eksperimental'naya genetika i fiziologiya. Trudy KNC RAN. Vyp. 11. Petrozavodsk, 2007. P. 85–92.

Korotkov Yu. M. Korotkova E. B. New data on ecology of Rana terrestris in the Tuva, Ekologiya. 1976. Vyp. 3. P. 102–103.

Kochetkov S. N. Shubina Yu. E. Distribution and ecology of the common frog in Lipetsk oblast’, Voprosy gerpetologii: Materialy IV s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Kazan', 12–17 oktyabrya 2009 g.). SPb., 2011. P. 113–118.

Krasavcev B. A. Data on ecology of the moor frog, Voprosy ekologii i biocenologii. L.: LGU, 1939. Vyp. 4. P. 253–267.

Krivko A. M. On the distribution and ecology of amphibians of Northern Prikaspiy, Biologicheskie nauki. Alma-Ata, 1976. Vyp. 3. P. 46–48.

Kuz'min S. L. Amphibians and reptiles of the north-west of Moscow, Zemnovodnye i presmykayuschiesya Moskovskoy oblasti. M.: MOIP, 1989. P. 48–60.

Kuz'min S. L. Amphibians of the former Soviet Union. M.: Tovarischestvo nauchnyh izdaniy KMK, 2012. 370 p.

Kuranova V. N. Population dynamics in anuran amphibians in the south-west of Western Siberia, Voprosy gerpetologii: Materialy I s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Puschino). M., 2001. P. 147–149.

Kuranova V. N. Kashtanova M. V. Amphibians and reptiles of the Ilmensky Reserve and Miass suburb area (Chelyabinsk province), Voprosy gerpetologii: Materialy I s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Puschino). M., 2001. P. 149–152.

Kutenkov A. P. Breeding and some ecological patterns of the moor frog in middle Karelia, Vid i ego produktivnost' v areale: Materialy 4-go Vsesoyuznogo soveschaniya. Ch. V. Voprosy gerpetologii. Sverdlovsk, 1984. P. 25.

Kutenkov A. P. Breeding of anuran amphibians in southern Karelia and impact of human activity, Antropogennye vozdeystviya na prirodu zapovednikov. M.: CNIL Glavohoty RSFSR, 1990. P. 38–50.

Kutenkov A. P. Dynamics of liver, fat bodies and gonads dimensions in the common frog and the moor frog, Ekologiya nazemnyh pozvonochnyh. Petrozavodsk, 1991. P. 14–24.

Kutenkov A. P. Ecology of Rana temporaria in the North-West of Russia. Petrozavodsk, 2009. 140 p.

Kutenkov A. P. Korosov A. V. Biogeography of amphibians of the Kola-Karelian region, Biogeografiya Karelii. Trudy KNC RAN. Ser. B. Vyp. 2. Petrozavodsk, 2001. P. 103–109.

Kutenkov A. P. Panarin A. E. Shklyarevich F. N. Breeding ecology of anuran amphibians of Karelia and Kola Peninsula, Nazemnye pozvonochnye zhivotnye v zapovednikah severa evropeyskoy chasti RSFSR. M.: CNIL Glavohoty RSFSR, 1990. P. 54–70.

Kushniruk V. A. On the hibernation of some amphibians in the western provinces of Ukraine, Voprosy gerpetologii: Avtoref. dokl. 3 Vsesoyuznoy gerpetol. konf. L.: LGU, 1964. P. 37–38.

Kushniruk V. A. On the biology of the moor frog in the west of Ukraine, Fauna Moldavii i ee ohrana: Materialy dokladov Pervoy respublikanskoy konf. Kishinev, 1970. P. 141.

Lada G. A. Ecological and faunistic analysis of amphibians in central Chernosem'ye. SPb., 1993. 22 p.

Leont'eva O. A. Some aspects of ecology of brown frogs in the northern border of their geographical area, Vzaimodeystvie organizmov v tundrovyh ekosistemah: Tez. dokl. Vsesoyuznoy sovesch. Syktyvkar, 1989. P. 146.

Leont'eva O. A. Pereshkol'nik S. L. Indicator role of herpetofauna in ecosystems with different anthropogenic transformation, Zhivotnyy mir centra lesnoy zony evropeyskoy chasti SSSR. Kalinin, 1982. P. 49–62.

Lobanov V. A. Distribution of the moor frog in Bolshezemelskaya tundra, Voprosy gerpetologii: Avtoref. dokl. 4 Vsesoyuznoy gerpetol. konf. L.: Nauka, 1977. P. 134–135.

The moor frog. Herpetofauna of Tyumen province. URL: http://herptyumen.narod.ru/amphibia/rana_arvalis.html (data obrascheniya 26.10.2015).Lyagushka travyanaya [The common frog], V. A. Krivosheev, Krasnaya kniga Ul'yanovskoy oblasti (griby, zhivotnye). T. 1. Ul'yanovsk, 2004. URL: http://geohobby.ru/enc/02-3-1-002.html (data obrascheniya 24.09.2014).

The common frog, O. A. Ermakov, Krasnaya kniga Penzenskoy obl. T. 2. Penza, 2005. P. 105.

The common frog, A. I. Fayzulin, A. G. Bakiev, Krasnaya kniga Samarskoy obl. T. II. Redkie vidy zhivotnyh. P. 237. URL: https://issuu.com/dd_nn/docs/krasnaq_kniga_samarskoj _oblasti._tom_2 (data obrascheniya 14.09.2014).

The common frog. Herpetofauna of Tyumen province. URL: http://herptyumen.narod.ru/amphibia/rana_temporaria.html (data obrascheniya 26.10.2015).

Lyapkov S. M. Long-term number dynamics of brown frog populations in Moskowskaya province: natural fluctuations or the response to the increased anthropogenic impact?, Voprosy gerpetologii: Materialy I s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Puschino). M., 2001. P. 171–173.

Maksimov A. A. Merzlyakova E. P. Characteristic of overflows in the floodplain of the Ob river in 1968–1977, Sukcessii zhivotnogo naseleniya v biocenozah poymy reki Obi. Novosibirsk, 1981. P. 165–174.

Maleev V. G. to distribution of amphibians and reptiles in the territory of Ust-Ordinsky autonomous region (upper Priangar'ye), Baykal'skiy zoologicheskiy zhurnal. 2009. No. 1. P. 48–49.

The Meshchera lowland. URL: https://ru.wikipedia.org/wiki/Mescherskaya_nizmennost' (data obrascheniya 17.02.2016).

Nikitenko M. F. Amphibians of the Soviet Bukovina, Zhivotnyy mir Sovetskoy Bukoviny. Chernovcy, 1959. P. 160–205.

Oksky Reserve. URL: http://oksky-reserve.ru/index/geography/ (data obrascheniya 21.12.2014).

The rivers Oka and Don Plain. URL: http://ru.wikipedia.org/wiki/Oksko-Donskaya_ravnina (data obrascheniya 17.02.2016).

The moor frog, Krasnaya kniga Respubliki Buryatiya. URL: http://www.minpriroda-rb.ru/redbook/2004/animals/amphibious_index.php?ELEMENT_ID=2449 (data obrascheniya 24.09.2014a).

The moor frog, O. N. Chernyshova, L. N. Erdakov, V. N. Kuranova, M. V. Pestov), Informacionnye materialy k gerpetofaune Sibiri. URL: http://www.balatsky.de/NSO/Amphibii/Rana_arvalis.htm (data obrascheniya 09.09.2014b).

Panchenko I. M. Reproduction efficiency of Rana arvalis in the south-eastern Meshchera, Ekologiya. 1980. No. 6. P. 95–98.

Pereshkol'nik S. L. Leont'eva O. A. Long-term observations on the variation of the herpetofauna in Prioksko-terrasny State Reserve, Zemnovodnye i presmykayuschiesya Moskovskoy oblasti. M.: MOIP, 1989. P. 84–96.

Pikulik M. M. The amphibians of Belarus. Minsk, 1985. 192 p.

Preobrazhenskaya E. S. Baykalova A. S. Abandance and habitat distribution of amphibians beyond the ponds, Zhivotnyy mir yuzhnoy taygi. Problemy i metody issledovaniya. M.: Nauka, 1984. P. 83–90.

Nature objects of Aktiubinsk province. URL: http://kzgov.docdat.com/docs/76/index-3326806.html (data obrascheniya 05.09.2014).

Ptushenko E. S. Terrestrial vertebrates of the Kursk region. Amphibians and reptiles, Byulleten' MOIP. Otd. biol. 1934. T. 33. Vyp. 1. P. 35–51.

Puzachenko Yu. G. Kuz'min S. L. Sandlerskiy R. B. Quantitative estimation of area parameters (on the example of genus Rana ), Zhurnal obschey biologii. 2011. T. 72. No. 5. P. 339–354.

Ravkin Yu. S. Abundance and distribution of the amphibian in the forest zone of the Western and Middle Siberia, Ekologiya. 1976. No. 5. P. 53–61.

Mordovia Republic. Geographic review. URL: www.geografia.ru/mordovia.html (data obrascheniya 04.09.2014).

Ruchin A. B. Ecology of amphibians and reptiles of Mordovia. The common frog Rana temporaria, Trudy Mordovskogo gosudarstvennogo prirodnogo zapovednika. Saransk, 2015. Vyp. 14. P. 344–358.

Ryzhevich K. K. Landscape and geographical variation in the proportion of Rana temporaria and Rana arvalis numbers in Belarus, Voprosy gerpetologii: Materialy I s'ezda Gerpetologicheskogo obschestva im. A. M. Nikol'skogo (Puschino). M., 2001. P. 254–255.

Ryzhov M. K. Ruchin A. B. Biology of Rana arvalis in Mordovia. Distribution, abundance and habitats, Biologicheskie nauki Kazahstana. 2007. No. 3. P. 33–39.

Severcov A. S. Lyapkov S. M. Surova G. S. Comparative analysis of ecological niches of common frog and moor frog, Zhurnal obschey biologii. 1998. T. 59. No. 3. P. 279–301.

Sedalischev V. T. Belimov G. T. Bekeneva G. N. Some morphophysiological adaptations of Rana arvalis in southern Yakutia, Voprosy gerpetologii: Avtoref. dokl. 5 Vsesoyuznoy gerpetol. konf. L.: Nauka, 1981. P. 122–123.

Sergeev A. M. Vetsheva A. G. On the drought impact on the number dynamics of the common frog in European part of USSR, Zoologicheskiy zhurnal. 1942. T. 21. Vyp. 5. P. 202–206.

Starikov V. P. Matkovskiy A. V. Spread and some aspects of the common frog ecology in the north of Western Siberia, Vestnik Krasnoyarskogo gosudarstvennogo agrarnogo universiteta. 2011. No. 12. P. 125–128.

Tisa. URL: http://dic.academic.ru/dic.nsf/enc_geo/4887/%D0%A2%D0%B8%D1%81%D0%B0 (data obrascheniya 19.04.2015).

Toporkova L. Ya. Varfolomeev V. V. On distribution of amphibians and reptiles on Basegi mountain ridge, Vid i ego produktivnost' v areale: Materialy 4-go Vsesoyuz. sovesch. Ch. V. Voprosy gerpetologii. Sverdlovsk, 1984. P. 44–45.

Toporkova L. Ya. Zubareva E. L. On the ecology of the common frog in the Polar Ural, Trudy Instituta biologii UF AN SSSR. 1965. Vyp. 38. P. 189–193.

The common frog, A. V. Davygora, Krasnaya kniga Orenburgskoy obl. Zhivotnye i rasteniya.  Orenburg, 1998. URL: http://oopt.aari.ru/rbdata/1100/bio/8480 (data obrascheniya 24.09.2014).

The common frog, M. V. Didorchuk, Krasnaya kniga Ryazanskoy oblasti. Redkie i nahodyaschiesya pod ugrozoy ischeznoveniya vidy zhivotnyh. Ryazan', 2001. P. 138.

The common frog, P. V. Repitunov, A. I. Masalykin , Krasnaya kniga Voronezhskoy obl. T. 2. Zhivotnye. Voronezh, 2011. P. 258.

The common frog, V. G. Ischenko, Krasnaya kniga Kurganskoy oblasti. Zemnovodnye. Kurgan, 2012a. P. 94.

The common frog, G. A. Lada, A. P. Sokolov, Krasnaya kniga Tambovskoy oblasti: Zhivotnye. Tambov, 2012b. P. 208.

The common frog, V. L. Vershinin, M. V. Chibiryak , Ural'skaya ekologicheskaya enciklopediya. URL: http://ecoinf.uran.ru/00003437.html (data obrascheniya 31.03.2014).

Ukrainian Carpathians — the geography. URL: http://uchebnikionline.com/geografia/geografiya_-_oliynik_yab/ukrayinski_karpati.htm (data obrascheniya 17.10.2015).

Ulyanovsc province. Geographic review. URL: http://www.geografia.ru/ulyan.html (data obrascheniya 17.02.2016).

Fomichev S. N. Ecology of island populations of brown frogs in Karelia. Petrozavodsk, 2004. 188 p.

Central Black earth Reserve. URL: http://zapoved-kursk.ru (data obrascheniya 05.09.2014).

Shaposhnikov L. V. Golovin O. Sorokin M. Tarakanov A. Fauna of the Kalinin province. Kalinin, 1959. 460 p.

Sharleman E. V. Note about fauna of amphibians and reptiles in the Kiev suroundings, Materialy k poznaniyu fauny yugo-zapadnoy Rossii. Kiev, 1917. P. 1–17.

Shvarc S. S. Ischenko V. G. Ways of adaptations of terrestrial vertebrates to the Subarctic living conditions. Sverdlovsk, 1971. 60 p.

Shvecov Yu. G. Amphibians and reptiles of northern Pribaykalie, Voprosy gerpetologii: Avtoref. dokl. 4 Vsesoyuznoy gerpetol. konf. L.: Nauka, 1977. P. 232–235.

Shlyahtin G. V. Tabachishin V. G. Zav'yalov E. V. Tabachishina I. E. Fauna of the Saratov province. Vol. 4. Amphibians and reptiles. Saratov, 2005. 116 p.

Epova L. A. Kuranova V. N. Babina S. G. Species diversity, spatial distribution, and abundance of amphibians and reptiles in Kuznetsk Alatau natural reserve in altitude gradient (south-east of Western Siberia), Vestnik Tomskogo gosudarstvennogo universiteta. Biologiya. 2013. No. 4 (24). P. 77–97.

Yakovlev V. A. Distribution and habitat allocation of amphibians and reptiles in Altai Reserve, Voprosy gerpetologii: Avtoref. dokl. 4 Vsesoyuznoy gerpetol. konf. L.: Nauka, 1977. P. 241–242.

Yakovlev V. A. On reproduction and development of the moor frog, Rana arvalis, in  Altai nature reserve, Ekologiya i sistematika amfibiy i reptiliy. Trudy ZIN AN SSSR. 1979. T. 89. P. 109–117.

Yakovlev V. A. On vertical distribution and reproduction of Rana terrestris in Altai Reserve, Ekologiya. 1980. No. 4. P. 89–90.

Yakovlev V. A. Data on reproducnion of Rana arvalis under conditions of highmountains, Ekologiya. 1981. No. 1. P. 97–101.

Andrén C., Nilson G. Effects of acidification on Swedish brown frogs, Memoranda soc. fauna et flora fennica. 1988. Vol. 64. No. 3. P. 139–141.

Bea A., Rodriquez-Teijerio J. D., Jover L. L. Relations between meteorological variables and the initiation of the spawning period in populations of Rana temporaria L. in the Atlantic region of the Basque Country (northern Spain), Amphibia – Reptilia. 1986. Vol. 7. No. 1. P. 23–31.

Berger L. Gady I płazy (Amphibia et Reptilia), Fauna słodkowodna Polski. No. 4. Warszawa; Poznań, 1975. 109 s.

Cogălniceanu D., Szekely P., Samoilă C, Iosif R., Tudor M., Plăiaşu R., Stănescu F., Rozylowicz L. Diversity and distribution of amphibians in Romania, ZooKeys. 2013. No. 296. P. 35–57. DOI: 10.3897/zookeys.296.4872.

Costanzo J., de Amaral C., Rosendale A., Lee R. Hibernation physiology, freezing adaptation and extreme freeze tolerance in a northern population of the wood frog., Journ. of exp. boil. 2013. Vol. 216. P. 3461–3473.

Covaciu-Marcov S, D., Cicort-Lucaciu A., Gaceu O., Sas I., Ferenţi S., Bogdan H. The herpetofauna of the south-western part of Mehedinţi County, Romania, North-western journ. of zool. (Romania). 2009a. Vol. 5. No. 1. P. 142–164. URL: http://www.lacerta.de/AS/Bibliografie/BIB_4967.pdf (data obrascheniya 23.08. 2015).

Covaciu-Marcov S, D., Dincă I., Dimancea N. The herpetofauna of the hydrographical basin of the Moca stream from Valea lui Mihai town, Bihor County, Romania, Biharean biologist. 2009b. Vol. 3. No. 2. P. 125–131. URL: http://biozoojournals.ro/bihbiol/cont/v3n2/bb.031118.Covaciu.pdf (data obrascheniya 30.04. 2015).

Delft J. van, Creemers R. Distribution, status and conservation of the moor frog (Rana arvalis) in the Netherlands, Zeitschrift für feldherpetologie. 2008. Suppl. 13. P. 255–268. URL: http://www.ravon.nl/Portals/0/Pdf/21-van%20Delft%20%20Cremers%20DEF%20DEF.pdf (data obrascheniya 15.01.2013).

Demeter L., Csergő A-M., Sándor A. D., Imets I., Vizauer C. T. Natural treasures of the Csík Basin (Depresiunea Cicului) and Csík Mountains (Munţii Ciucului), Mountain hay meadows: hotspots of biodiversity and traditional culture. 2011. URL: http://www.mountainhaymeadows.eu/online_publication/03-natural-treasures-of-the-csik-basin-and-csik-mountains.html (data obrascheniya 30.04. 2015).

Demeter L., Kelemen A. Data on the distribution and population size of amphibians in the Csík Mountains (Munţii Ciucului), Eastern Carpathians, Romania, Mountain hay meadows: hotspots of biodiversity and traditional culture. 2011. URL: http://www.mountainhaymeadows.eu/online_publication/06-data-on-the-distribution-and-population-size-of-amphibians-in-the-csik-mountains.html (data obrascheniya 30.04. 2015).

Dolmen D. Norwegian amphibians and reptiles; current situation 1985, Studies in Herpetology (Roček Z. ed.). Prague, 1986. P. 743–746.

Dolmen D. Distribution, habitat, ecology and status of the moor frog (Rana arvalis) in Norway, Zeitschrift für feldherpetologie. 2008. Suppl. 13. P. 167–178.

Elmberg J. Ǻkergrodan. En artöversikt samt nya rön om dess utbredning i Nord-och Mellansverige, Fauna och flora. 1978. Vol. 73. No. 2. R. 69–78.

Elmberg J. Apparent lack of territoriality during the breeding season in a boreal population of common frogs Rana temporaria, Herpetol. journ. 1986. Vol. 1. No. 2. P. 81–85.

Elmberg J. Factors affecting male yearly mating success in the common frogs Rana temporaria, Behav. ecol. sociobiol. 1991. Vol. 28. P. 125–131.

Elmberg J. Grod- och kräldjurens utbreding i Norrland, Natur i Norr (Umeá). 1995. Ǻrgáng 14, h. 2. P. 57–82.

Elmberg J. Ecology and life history of the moor frog (Rana arvalis) in boreal Sweeden, Zeitschrift für feldherpetologie. 2008. Suppl. 13. P. 179–194.

Elmberg J., Lundberg P. Intraspecific variation in calling, time allocation and energy reserves in breeding male common frogs Rana temporaria, Annales zool. fennici. 1991. Vol. 28. No. 1. P. 23–29.

Gasc J, P., Cabela A., Crnobrnja-Isailovic J, Dolmen D., Grossenbacher K., Haffner P., Lescure J., Martens H., Martinez R., Maurin H., Oliviera M., Sofianidou T., Veith M., Zuiderwijk A. (eds). Atlas of amphibians and reptiles in Europe. Paris, 1997. 516 p.

Gislén T., Kauri H. Zoogeography of the Swedish amphibians and reptiles, with notes on their growth and ecology, Acta vertebrat. 1959. Vol. 1. No. 3. P. 191–397.

Glandt D. Rana arvalis Nilsson, 1842 — Moorfrosch, Handbuch der reptilian und amphibien Europas/ Band 5/III A: froschlurche (Anura) III A (Ranidae I). AULA–Verlag, 2014. S. 12–113.

Glasnovič P., Krystufek B., Sovinc A., Bojović M., Porej D. Protected area gap analysis (final report). 2009. 327 p. URL: http://www.discoverdinarides.com/files/file/gap-analysis-final-report-1363265933.pdf

Gleed-Owen C. P. Subfossil records of Rana cf. lessonae, Rana arvalis and Rana cf. dalmatina from Middle Saxon (c. 600-950 AD) deposits in eastern England: evidence for native status, Amphibia – Reptilia. 2000. Vol. 21. P. 57–65.

Haapanen A. Site tenacity of the common frog (Rana temporaria L.) and the moor frog (R. arvalis Nilss.), Annales zool. fennici. 1970. Vol. 7. No. 1. P. 61–66.

Hangartner S., Laurila A., Räsänen K. Adaptive divergence of the moor frog (Rana arvalis) along an acidification gradient, BMC Evolutionary Biology. 2011. Vol. 11. P. 336. URL: http://www.biomedcentral.com/1471-2148/11/366 (data obrascheniya 13.03.2015).

Hartel T., Öllerer K., Demeter L., Nemes Sz., Moga C., Cogălniceanu D. Using a habitat based approach in mapping amphibian distribution in Romania: study case of the Saxon cultural landscape in the Târnava Mare basin. URL: www.mihaieminescutrust.org/files/studies/Hartel.amphibian.distribution.pdf (data obrascheniya 23.09.2014).

Heimer W. Amphibienvorkommen im Ostteil des landkreises Darmstadt-Dieburg, Hess. faun. briefe. 1981. Vol. 1. No. 2. S. 20–23.

Heráň I. Výsledky výzkumu prostorové aktivity skokana hnédégo, Rana temporaria L., 1758, v Krknosském národnim parku, Sbornik národneho muzea v Praze. 1982. T. 38 B. No. 4. S. 239–263.

Ishchenko V. Rana arvalis Nilsson, 1842, Atlas of amphibians and reptiles in Europe (Gasc J, P. ed.). Paris, 1997. P. 128–129.

Jablonski D., Jandzik D., Gvoždik V. New records and zoogeographic classification of amphibians and reptiles from Bosnia and Herzegovina, North-western journ. of zool. (Romania). 2012. Vol. 8. No. 2. P. 324–337. URL: http://biozoojournals.3x.ro/nwjz/index.html

Jędrzejewska B., Brzeziński M., Jędrzejewski W. Seasonal dynamics and breeding of amphibians in prestine forests (Białowieża National Park, E Poland) in dry years, Folia zool. 2003. Vol. 52. No. 1. P. 77–86.

Juszczyk W. Płazy i gady kraiowe. Warszawa, Cześć 2, 1987. 384 s.

Koskela P., Pasanen S. The wintering of the common frog, Rana temporaria L., in northern Finland, Aquilo, ser. zool. 1974. Vol. 15. P. 1–17.

Kutenkov A. P., Panarin A. E. Ecology and status of populations of the common frog (Rana temporaria) and the moor frog (Rana arvalis) in northwestern Russia with notes on their distribution in Fennoscandia, Amphibian populations in the Commonwealth of Independence States: current status and declines. Moscow: Pensoft, 1995. P. 64–70.

Laan R., Verboom B. Effects of pool size and isolation on amphibian communities, Biol. conserv. 1990. Vol. 54. P. 251–262.

Loman J. Macro- and microhabitat distribution in Rana arvalis and R. temporaria (Amphibia, Anura, Ranidae) during summer, Journ. of herpetol. 1978. Vol. 12. No. 1. P. 29–33.

Loman J. Annual and daily locomotor activity of the frogs Rana arvalis and R. temporaria, Brit. journ. of herpetol. 1980. Vol. 6. No. 3. P. 83–85.

Loman J. Density and survival of Rana arvalis and Rana temporaria, Alytes. 1984. Vol. 3. No. 4. P. 125–134.

Loman J., Andersson G. Monitoring brown frogs Rana arvalis and Rana temporaria in 120 south Swedish ponds 1989–2005. Mixed trends in different habitats, Biological conserv. 2007. Vol. 135. P. 46–56.

Loman J., Lardner B. Does pond quality limit frogs Rana arvalis and R. temporaria in agricultural landscapes? A field experiment, Journ. of applied ecol. 2006. P. 690–700.

Miaud C., Guyétant R., Humbert A. Age structure in montane population of common frog (Rana temporaria), 8th Ordinary general meeting Societas Eur. Herpetol. Bonn, 1995. P. 85–86.

Montanarella L., Jones R. J. A., Hiederer R. The distribution of peatland in Europe, Mires and Peat. 2006. Vol. 1. Article 01, 10 pp. URL: http://www.mires-and-peat.net.

Okulova N. M. Amphibians in northern European Russia, Amphibian populations in the Commonwealth of Independent States: current status and declines. Moscow: Pensoft, 1995. P. 59–63.

Opatrný E. Beitrag zur erkenntnis der verbreitung der amphibienfauna in der Tschechoslowakei, Acta univer. palack. olom. 1978. T. 59. S. 205–220.

Pasanen S., Karhaää M. Can boreal common frog (Rana temporaria L.) survive in frost?, Annales zool. fennici. 1997. Vol. 34. No. 4. P. 247–250.

Pasanen S., Sorjonen J. Partial terrestrial wintering in a northern common frog population (Rana temporaria L.), Annales zool. fennici. 1994. Vol. 31. No. 2. P. 275–278.

Pascual X., Montori A. Contribucion al estudo de Rana temporaria L. (Amphibia, Ranidae) en Santa Fe del Monsteny (Barcelona). I. Descripcion de la zona y estima de la poblacion, Miscel. zool. 1981. No. 7. P. 109–115.

Petrov B. 4. Amphibians and reptiles of Bulgaria: fauna, vertical distribution, zoogeography, and conservation, Biogeography and ecology of Bulgaria (V. Fet and A. Popov eds.). Springer, 2007. P. 85–107. URL: http://www.nmnhs.com/downloads/pdfs/petrov-boyan/biogeograpgy-ecology-bulgaria-2007-85-107.pdf.

Petrov B., Tzankov N., Strijbosch H., Popgeorgiev G., Beshkov V. The herpetofauna (Amphibia and Reptilia) of the Western Rhodopes mountain (Bulgaria and Greece), Beron P. (ed.). Biodiversity of Bulgaria. 3. Biodiversity of Western Rhodopes (Bulgaria and Greece). Sofia, 2006. P. 863–912. URL: http://www.lacerta.de/AS/Bibliografie/BIB_1439.pdf

Puky M., Shád P. Distribution and status of the moor frog (Rana arvalis) in Hungary, Zeitschrift für feldherpetologie. 2008. Suppl. 13. P. 309–316.

Puky M., Shád P., Szövéniyi G. Magyarország herpetológiai atlasza. Herpetological atlas of Hungary. Budapest, 2005. 207 p.

Räsänen K., Laurila A., Merilä J. Carry-over effects of embryonic acid conditions on development and growth of Rana temporaria tadpoles, Freshwater biol. 2002. Vol. 47. P. 19–30.

Rittenhouse T., Semlitsch R. Distribution of amphibians in terrestrial habitat surrounding wetlands, Wetlands, 2007. Vol. 27. No. 1. P. 153–161. URL: http://www.researchgate.net/ publication/226216986_Distribution_of_amphibians_in_terrestrial_habitat_surrounding_wetlands.

Roček Z., Šandera M. Distribution of Rana arvalis in Europe: a historical perspective, Zeitschrift für feldherpetologie. 2008. Suppl. 13. P. 135–150.

Sas I., Covaciu-Marcov S, D., Demeter L., Cicort-Lucaciu A, S., Strugariu A. Distribution and status of the moor frog (Rana arvalis) in Romania, Zeitschrift für feldherpetologie. 2008. Suppl. 13. P. 337–354.

Sas I., Covaciu-Marcov S, D., Kovács E, H., Radu N, R., Tóth A., Popa A. The populations of Rana arvalis Nills. 1842 from the Ier Valley (The Western Plain, Romania): present and future, North-western journ. of zool. (Romania). 2006. Vol. 2. No. 1. P. 1–16. http://01.nwjz.2.1.Sas.pdf

Savage R. M. The breeding behaviour of the common frog Rana temporaria Linn., and Bufo bufo Linn., Proc. zool. soc. London, 1934. Vol. 6. P. 55–70.

Serra-Cobo J., Lacroix G., White S. Comparison between the ecology of the new europaean frog Rana pyrenaica and that of four pyrenean amphibians, Journ. of zool. 1998. Vol. 246. No. 2. P. 147–154.

Schmid W. D. Survival of frogs in low temperature, Science. 1982. Vol. 215. No. 4533. P. 697–698.

Schweiger M. Rana arvalis near Vienna (video). URL: http://www.youtube.com/watch?v=bXJMK3ocgFY (data obrascheniya 19.11.2014).

Strijbosch H. Habitat selection of amphibians during their aquatic phase, Oikos. 1979. Vol. 33. No. 3. P. 363–372.

Storey K. B. Persistence of freeze tolerance in terrestrially hibernating frogs after spring emergence, Copeia. 1987. No. 3. R. 720–726.

Stugren B. Geographic variation and distribution of the moor frog, Rana arvalis Nills., Annales zool. fennici. 1966. Vol. 3. No. 1. P. 29–39.

Sztatecsny M., Hödl W. Can protected mountain areas serve as refuges for declining amphibians? Potential threats of climate change and amphibian chytridiomycosis in alpine amphibian population, Ecology of mountains. 2009. Vol. 1. No. 2. P. 19–24.

Sztatecsny M., Preininger D., Freudmann A., Matthias-Claudio L., Maier F., Hödl W. Don’t get the blues: conspicuous nuptial coloration of male moor frogs (Rana arvalis) support visual mate recognition during scramble competition in large breeding aggregations, Behav. ecol. sociobiol. 2012. Vol. 66. P. 1587–1593. DOI: 10.1007/s00265-012-1412-6.

Terhivuo J. Provisional atlas and population status of the Finnish amphibian and reptile species with reference to their ranges in northern Europe, Annales zool. fennici. 1981. Vol. 18. No. 3. P. 139–164.

Thiesmeier B. Untersuchungen zur Jahresaktivitat der erdkrote (Bufo bufo Linnaeus, 1758) und des grasfrosches (Rana temporaria Linnaeus, 1758) im Niederbergishen land, Decheniana. 1992. Vol. 145. S. 146–164.

Vences M., Grossenbacher K., Puente M., Palanca A., Vieites D. The Cambales fairy tale: elevational limits of Rana temporaria (Amphibia, Ranidae) and other Europaean amphibians revisited, Folia zool. Vol. 2003. 52. No. 3. P. 189–202. URL: http://www.ivb.cz/folia/52/2/189-202.pdf.

Vences M., Pique N., Lopez A., Puente M., Miramontes C., Vieites D. R. Summer habitat population estimate and body size variation in a high altitude population of Rana temporaria, Amphibia – Reptilia. 1999. Vol. 20. No. 4. P. 426–435.

Vogrin N. The status of Amphibians in Slovenia, Froglog. The newsletter of IUCN DAPTF. 1997. No. 20. P. 1–2.

Vos C. C., Chardon J. P. Effects of habitat fragmentation and road density on the distribution pattern of the moor frog Rana arvalis, Journ. of applied ecol. 1998. Vol. 35. P. 44–56.

Vos C. C., Goedhart P., Lammertsma D., Spitzen-Van der Sluijs A. Matrix permeability of agricultural landscapes: an analysis of movements of the common frog (Rana temporaria), Herpetol. j. 2007. Vol. 17. P. 174–182.

Vukov T., Kalezić M., Tomović L., Krizmanić I., Jović D., Džukić G. Amphibians in Serbia — distribution and diversity patterns, Bull. of the Natural Museum (Serbia). 2013. No. 6. P. 90–112. DOI: 10.5937/bnhmb1306090V

Wind-Larsen H., Jørgensen C. B. Hormonal control of seasonal growth in a temperate zone toad Bufo bufo, Acta zool. 1987. Vol. 68. No. 1. P. 49–56.

Displays: 5047; Downloads: 709;