Ground beetles of the Ukraine (Coleoptera, Carabidae)

Abstract A review of the ground beetles of the Ukrainian fauna is given. Almost 750 species from 117 genera of Carabidae are known to occur in the Ukraine. Approximately 450 species of ground beetles are registered in the Carpathian region. No less than 300 species of ground beetles are found in the forest zone. Approximately 400 species of Carabidae present in the forest-steppe zone are relatively similar in species composition to those in the forest territories. Some 450 species of Carabidae are inhabitants of the steppe zone. Representatives of many other regions of heterogeneous biotopes such as forest, semi desert, intrazonal, etc. can be found in the steppe areas. The fauna of Carabidae (ca. 100 species) of the lowlands of southern Ukraine (sandy biotopes), situated mostly in the Kherson region, is very peculiar. The fauna of the Crimean mountains contains about 300 species. Conservation measures for the Carabidae are discussed.

At present, there are more than 1000 literature sources that concern the general biology, systematic and ecology of ground beetle species, recorded from the territory of present-day Ukraine. The checklist of Carabidae of Russia and adjacent territories (Kryzhanovskij et al. 1995) is the largest summary on the species diversity of ground beetles in the Ukraine. In this book, data on the East Carpathian, Crimea and other regions of the Ukraine are presented. Furthermore, a more recent survey in the first volume of the Catalogue of Palearctic Coleoptera (2003) lists nearly 720 species of Carabidae that are indicated for the whole territory of the Ukraine. However, in spite of the fact that these publications span different geographical zones of the Ukraine, the distribution of ground beetles within the country remains poorly studied. Besides, there are nearly thirty species of Carabidae registered in the Ukraine that are not included in the Catalogue of Palearctic Coleoptera, 2003 (marked in this article by*).
The aim of the present paper is to summarise all available data from literature sources and collections and to provide an overview of the present-day species composition and distribution of ground beetles in the Ukraine.

Material and methods
The complete list of Carabidae of the Ukraine (Appendix 1) was compiled on the basis of a critical literature review and collections in several biological institutions in Kiev, Moscow, St.-Petersburg, Budapest, Vienna and Prague, including my own large collection. The tiger beetles (Coleoptera, Cicindelidae), as a separate family (Putchkov, Cassola, 2005) is not included in this article. The classification of Carabidae follows Kryzhanovskij et al. (1995) with some additional revision (Catalogue of Palearctic Coleoptera, 2003). The analysis of the distribution of Carabidae in the Ukraine is given on the basis of the whole territory of the country; however special attention was paid to 13 separate specific regions, districts and provinces (Fig. 1).

Results and discussion
Ground beetles (Carabidae) are one of the largest beetle families in the territory of the Ukraine. There are nearly 750 species from 117 genera present. Such rich biodiversity is due to the large area of the country on one hand, and the heterogeneity of natural conditions of the separate geographical regions on the other hand.
The ground beetle fauna of TL and CM are most diverse in the Ukraine (ca 330 and 400 species from 75 genera were found here, respectively) ( Table 1). Eight endemic taxa are registered in the East Carpathians: Leistus baenningeri Roubal, 1926, L. ucrainicus Lazorko, 1954, Nebria heegeri Dejean, 1826, Duvalius transcarpathicus Shilenkov et Rizun, 1989, D. ruthenus ruthenus Reitter, 1878, D. corpulentus Weise, 1825, Trechus pseudomontanellus Rizun, 1994, Carabus zawadskyi serriatissimus Reiter, 1896, C. fabricii ucrainicus Lazorko, 1951. More than 20 taxa are subendemic for this region (mostly from the genera Carabus, Nebria, Trechus and Pterostichus). In addition, more than 80 species that are known from the East Carpathians are absent from other geographic regions of the Ukraine. For approximately 50 taxa the Carpathians appear to be the eastern border of their ranges. These are some species belonging to the genera Nebria, Carabus, Pterostichus, Tachyura, Trechus, and separate species of Bembidion.   Most of these species inhabit subalpine and alpine biotopes. Some typical Middle-European species occur in the different types of mountain forests, where they comprise one of the major components of carabid diversity in the Carpathians. At the same time, the fauna of ground beetles in the Carpathians includes also many widespread species that inhabit other forest and forest-steppe areas of the Ukraine. The fauna of forest areas of the Ukraine is represented by no less than 300 species from 70 genera and is characterized by considerable similarity in species composition in all separate regions and zones (Table 1). It appears also similar to those from other northern areas of eastern Europe. Endemics are absent from forest area of the Ukraine, although about 10 species are present only in forest areas of the country. There are some boreal taxa of carabids that are registered only in the north-western part of the Ukraine (some species of Carabus, Miscodera*, Agonum, Trechus, Pterostichus). The territory of RF is slightly richest in terms of species diversity of Carabidae then LF of the forest zone (Table 1).
Ground beetles of the present-day forest-steppe zone are represented by nearly 400 species from 75 genera ( Table 1). The fauna of this zone is not typically transitional from forest to steppe. Obviously, the species composition of ground beetles in the forest-steppe can be characterized as quite distinct, although with some similarities to the forest zone. The variety and number of typical forest species (especially hygrophilous and mesophilous ones) exceeds the number of steppe-specific inhabitants. The territory of WRS is the richest (365 species from 73 genera) in terms of species diversity of Carabidae as compared to that of ERS (310 species from 65 genera) (Table 1). Forest species are more common in the western region between the Dniester and South Bug rivers. Four species -Carabus sibiricus rybinskii Reitter, 1896, Laemostenus tichyi Kult, 1946 (both are endemics of the Ukraine), Poecilus szepligetii Csiki, 1908 and Aptinus bombarda Illiger, 1800 occur only in the western area. Moreover, some ground beetles (Carabus excellens Fabricius, 1798, C. marginalis Fabricius, 1794, C. scabriusculus Olivier, 1795, some Calathus) are more abundant in the forest-steppe zone than in northern or southern regions. However there are many forest species of Carabidae that occur in WRS that are absent from the ERS. The composition Carabidae in the ELS is characterized by an increasing number of mesoxerophilous species, which are more common in the steppes than in the forest-steppe (Harpalus, Cymindis) ( Table 1).
The ground beetle fauna of the steppe area of the Ukraine is the richest in species diversity and is characterized by the presence of approximately 450 species from 76 genera ( Table 1). The taxonomic structure of ground beetles of the steppe is very diverse due to the heterogenous origin of steppe Carabidae fauna. The occurrence of many extrazonal (forest or semi-desert) and intrazonal (littoral, halophilous) species in the steppe region zone makes it difficult to characterize the general composition of the carabid fauna. As a whole, about 100 taxa of ground beetles in the fauna of the Ukraine occur exclusively in the steppe area (especially the genera Scarites, Apotomus, Zuphium, many Harpalini, Zabrini and some species of Poecilus, Chlaenius and Brachinus). However, the majority of ground beetles in this area is formed by typical steppe or Mediterranean taxa (mostly from the tribes Harpalini, Zabrini, Lebiini). Some forest and forest-steppe species (tribes Nebriini, Carabini, Platinini) are more common in the northern subzone (NRS and NLS) of the steppe zone. In addition, anthropogenic factors have supported predominance of some widespread mesophilous species in this subzone.
The ground beetle fauna of the steppe area of NLS is similar as a whole to that of the NRS; however it is characterized at the same time by the predominance of typical steppe taxa. Only the district of the Donetskyi heights (ridge) is characterized by more mesophilous elements including polytopic and forest species (some Carabus, Pterostichus and Agonum). At the same time, the occurrence of typical steppe taxa, including separate inhabitants of Caucasian and Kazakhstanian steppes (Poecilus anodon Chaudoir, 1868, P. lyroderes Chaudoir, 1846, Curtonotus propinquus Menetries, 1832, and some Cymindis species) could be observed in this region. Possibly, earlier, the Donetskyi ridge was characterised by a ground beetle fauna transitional between forest-steppe and steppe zones. Present-day diversity of Carabidae of this region is relatively closer to that of typical steppe fauna.
The southern steppe subzone (SRS and SLS) is characterized by the prevalence of xerophilous and mesoxerophilous species from the tribes Harpalini and Lebiini, while relatively mesophilous taxa occur more exceptionally in river valleys, ravines or in agricultural biotopes. The occurrence of some mesohygrophilous species in the steppe is usually related to irrigation.
The majority of littoral and halophilous species (tribes Clivinini, Bembidiini, Tachyini, Pogonini and Stenolophina) occur in river valleys, coastal beaches of gulfs, lakes, estuaries and other water basins. The ground beetle fauna of seashores and estuaries is characterized by the prevailence of many species that are absent from other regions of the Ukraine (some Dyschirius, Tachys, Bembidion, Acupalpus, Trichocellus etc.).
The ground beetle fauna of the Crimean Peninsula is one of the most specific in the Ukraine (about 390 species from 74 genera). There are some typical inhabitants of steppe and halophitic biotopes of the plains of Crimea (near 370 species): Calosoma Carabus, Poecilus, Amara, many Harpalini and Cymindina (Table 1). On the Kerch Peninsula some relatively forest mesophilous species occur: Carabus cancellatus Illiger, 1798, Leistus ferrugineus Linnaeus, 1758, Pterostichus niger Schaller, 1783 and Pt. anthracinus Illiger, 1798. This confirms the presence of arboreal areas in the ancient past.
Special attention should be paid to the fauna of anthropogenic landscapes of the Ukraine. In agricultural habitats, the species composition of ground beetles is relatively uniform throughout the different geographical regions. Agrocenoses are generally poor in species richness consisting of some 70-100 widely distributed common species, but the abundance of some of these is much higher than in natural biotopes. The core faunal composition consists of approximately 20 widespread (mainly polytopic) species from the genera Amara, Bembidion, Harpalus, Poecilus and Pterostichus. The fauna of urban territories (for example cities) is rather impoverished as a rule and consists of some 10-15 polytopic species.

Conclusions
I conclude that the overall species composition of the ground beetle fauna of the Ukraine is well studied. Therefore, finding new taxa in any part of the Ukraine is not likely to happen in the near future. Regions where new taxa for the Ukraine could be found are mainly boundary regions of the country (the Carpathians, Crimea, north and east regions), as well as in errors of some findings in the past (for example, misidentifications of some taxa or species with wrong labels). Poorly answered questions that remain include the origins of carabid fauna of the Ukraine. Moreover it is necessary to study the preimaginal stages of ground beetles. For example the larvae of only 360 species from 86 genera are described, representing only 45% of carabid species from the Ukraine. Further studies on the bionomics of single species require urgent attention. This applies in particular to ecologically related, non-competitive species occurring in common biotopes, e.g. many littoral species of Bembidion, Dyschirius, Acupalpus or some steppe species of the Harpalini tribe.