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

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

Kutenkov
   Anatoly
Ph.D., Reservation "Kivatch", 186200, Kondopoga reg., v. Waterfall Kivatch, stapesy@mail.ru
Keywords:
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

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