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Collage by M. Dianov
(photo by V. Tanasijchuk)

Geographic distribution is one of the major characteristics of any animal taxon, be it a species, a genus, or a family. This assumption is undoubtedly true also for beetles. The distribution range (i.e. the area or a totality of areas where the particular taxon occurs) of the entire order Coleoptera is enormous and is probably of record size as compared to ranges of other terrestrial animals. Beetles inhabit nearly every part of land available. They are only absent on ice sheets of the Antarctic Continent and Greenland, on some Arctic islands and in the zones of eternal snows in the alpine areas. For instance in the Himalayas they even occur at an altitude of 6000 m and in the Pamirs at an altitude of more than 4500 m. Beetles inhabit nearly every inland water body. The extensive distribution of Coleoptera is demonstrated in the collage by our Webmaster Michael Dianov.

Distribution patterns of animals on Earth are studied by zoogeography. There are different viewpoints of division of zoogeography into different disciplines. We believe that for entomologists using zoogeography as a tool in a study of an insect taxon it is sufficient to distinguish three disciplines for three phases of a typical study of a taxon's geographic distribution pattern.

The first phase is usually a study of local faunae, which includes compiling lists of species for separate areas. This issue is dealt with in the faunal study -- a special domain of zoogeography. After sufficient information on local faunae is accumulated the researcher usually begins defining distribution ranges of separate taxa, first distribution ranges of species, then of groups of species, subgenera, genera and taxa of higher ranks. A study of distribution ranges and classification of their types is called chorology or arealogy. Gathering of data on faunae and distribution ranges permits us to enter the final phase of applied zoogeographic research - division of the area studied (or the entire Earth) into zoogeographic regions in accordance with the character of their faunae and boundaries of distribution ranges of taxa of different ranks. The discipline dealing with this kind of study is called regional or analytical zoogeography or zoogeographic zoning.

Abundance of species of Coleoptera, diversity of their mode of life, ancient origin of many taxa, not infrequently dating back to the Triassic, and intensive adaptive radiation in a number of families determined the outstanding diversity of types of distribution ranges of beetles. This considerably hinders establishing their major geographic distribution patterns. Species distribution ranges are so very diverse that it is extremely difficult to classify them by type. Therefore, coleopterists prefer analysing distribution ranges on the level of subgenera, genera, tribes, subfamilies and families.

Beetles have long been successfully used for zoogeographic purposes. A.P.Semenov-Tian-Shansky developed a detailed scheme of zoogeographic subdivisions of the Palaearctic Region on the basis of Coleoptera distribution (1936), and proposed an interesting hypothesis for the origin of Alpine faunae (1937); in his works on systematics he considered many issues of historical zoogeography (Semenov-Tian-Shansky, 1898, 1900, 1934; Semenov-Tian-Shansky and Shchegoleva-Barovskaya, 1935). Noteworthy among zoogeographic publications of the past decades are works based on coleoptorological material by O.L.Kryzhanovsky (1965, 1971, 1976, 1980, 1983, 1987), I.K.Lopatin (1977, 1980), E.G.Matis (19..), G.M.Abdurakhmanov (1984, 1987), and others.

Among foreign entomogeographers we will mention only a few, who were at the same time eminent systematists of Coleoptera. The French entomologist Jeannel in his numerous works on systematics and faunal studies gave much attention to zoogeographic issues. He published a number of works devoted to zoogeography (Jeannel, 1942, 1961, etc.). Lindroth (Sweden) studied mostly entomofauna of Northern Europe and North America. His zoogeographic studies nearly approached the level of exact sciences. His major works concern faunae of Fennoscandia, Newfoundland, Canada and Alaska. He also published works on formation of faunae of the Azores, New Zealand, etc. (Lindroth, 1945-1949, 1955, 1957, 1961-1969, 1963, 1976, 1979, etc.). The American entomologist Gressitt conducted, based mostly on Cerambycidae and Chrysomelidae material, important studies on entomofauna of eastern and South-Eastern Asia and Papua Region and produced a concise but informative review on general zoogeography of insects (Gressitt, 1965-1967, 1974, etc.). The Australian entomologist Darlington based mainly upon material on distribution of ground beetles published a number of excellent zoogeographic works mostly on the Southern Hemishpere (Darlington, 1943, 1949, 1952-1971, 1957, 1961, 1965). Interesting generalizations of types of distribution ranges for higher taxa of beetles (tribes, subfamilies, families) were made by Crowson, 1981 (Great Britain) in the zoogeographic chapter of his fundamental work on life history of beetles.

Entomologists who have not accumulated sufficient faunal data frequently try to use a ready system of entomogeographic zoning of the land of Earth. However choosing one of the available systems it is necessary firstly to consider the principles of separation of large zoogeographic regions and zoogeographic zoning in general. This issue is one of the major ones in modern zoogeography because these principles have not been sufficiently elaborated yet and remain controversial in many respects. Many authors believe that simple zonal landscape division should supplant zoogeographic zonation schemes based on a detailed study of distribution ranges of taxa and faunal assemblages and on defining of historically determined differences between faunae. It is also a common viewpoint that the study of zootae (i.e. of animal population in landscapes and landscape zones) is the major and even the only goal of modern zoogeography. An optimal solution is probably to be found in the middle. It is possible to combine the two extreme directions because the zonal boundaries are such for the majority of organisms, and at the same time the vagueness of boundaries allows us to combine borders of different types of distribution ranges situated close to the zonal boundary.

Distribution of animals is determined by many factors, to which different groups of animals and plants respond differently. However the nature of Earth is a single entity and all animal and plant species inhabiting Earth are associated with each other: species inhabiting the same area are more closely associated with each another, whereas they are also bound by more distant mediated relations with all other species. Climate is the major factor responsible for geographic differentiation. It determines zonal distribution of living organisms, which is most dramatically demonstrated by vegetation cover. Nearly all other organisms, although to different degrees, depend on the character of vegetation. These relationships determine priority of biogeographic zoning over partitive general zoogeographic divisions, such as entomological, coleopterological, or even more partitive, for instance carabidogeographic (i.e. for ground beetles).

Another important point is that there cannot be a single biogeographic division in principle, as well as there cannot be partitive divisions, as for instance coleopterological. Division is determined by the principles underlying it; and the principles may be different. Search for universal principles is not the only trend in zoogeography.

The most simple and natural zoning for systematists is zoning by taxonomically important indices, i.e. distribution ranges of species, genera, endemism of different ranks, combinations of distribution ranges and "condensing" of their boundaries. Other approaches may be also used, based upon counts of numbers of species, analysis of their distribution by type of habitat (biotope), etc.

The zonal approach in its extreme form ignores species composition in the particular area. There are for instance steppe and prairie zones in Eurasia and North America, however, these zones are sharply different by species and generic composition because the biotas evolved nearly independently although from similar sources. The faunal approach, in its extreme, neglects zones and unites territories by composition alone and attaches primary importance to particular features of distribution of the particular group. Therefore, in such divisions a discovery of a new species or of a wider distribution range of the particular species frequently leads to modification of the scheme.

The general geographic approach permits regarding distribution of separate groups against the general nature background; it permits understanding distribution of insufficiently known groups by fragmentary data and predict with a certain degree of reliability the entire true distribution range by a few casual discoveries.

Combining the advantages of faunal and biogeographic zoning schemes is probably the major task for the future development of zoogeography. This goal can be achieved only based upon distribution of particular groups and therefore on many partitive zoning schemes, which will be subsequently permanently combined. In case of a partitive zoning, apart from the traditional approach "from below" (from species ranges to districts and provinces), it is also possible to use the approach "from above": from small parts of the general biogeographic zonation by combining them and establishing a particular hierarchy fit for the particular group of organisms.

The traditional (historically initial) way of zoogeographic zoning is dramatically demonstrated in the scheme of division of the Palaearctic by A.P. Semenov-Tian-Shansky (1936), in which very many boundaries coincide with the zonal ones. In this scheme the combinations also have a pronounced zonal character (e.g. Palaearchearctic comprises combined nemoral and subtropic zones of the eastern subsector of the Palaearctic). Further elaboration of this direction on the scale of the entire Earth was undertaken by coleopterist O.L.Kryzhanovsky in his last unpublished work.

A.F.Emeljanov (1974) proposed an approach from the opposite side ("from above"), i.e. a scheme of division of the Holarctic, which in his opinion combines the major principles of zonal and historical approaches. It can be used for clarification of distribution of insufficiently studied groups and primary classification of the material. This scheme is already used as a basis of partitive zoogeographic zoning.

A.F. Emeljanov, A.L. Lobanov
November, 1999


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