[The diversity, ecology and chorology of weevils of the Tatra Biosphere Reserve (Coleoptera: Curculionidea).]

Publication Type:Journal Article
:2005
Authors:S. Knutelski
Journal:Monografie Faunistyczne
Volume:23
Pagination:1-340
Type of Article:Article
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The present work is a recapitulation of studies which were conducted in the years 1984[long dash]1999. It summarizes all data on the diversity, ecology and chorology of the weevil species, which have so far been found in the area of the Tatra Mountains. The aim of the work was to prepare the revised faunistic list of Curculionoidea and to make an essay at the explanation of the genesis of the Tatra weevils on the basis of ecological, chorological and historical premises. The elaboration comprises the inventory of the weevil fauna, the regionalization of species on particular regions of the Tatra Mountains, and ecological and biogeographical characteristics, as well as the characteristics of the diversity, distribution. ecological requirements and geographical ranges of particular species. Special attention was paid to mountain species, the oldest and autochtonic element of the Tatra fauna. An additional task was to asses the value and usefulness of the weevil species for the purposes of valorization of the nature of mountain areas and monitoring changes in the natural environment of the Tatra Mts. The Tatra Mountains (UTM: DV 50-55, DV40-45; 49[degree] 10[minute] 42[second] 49[degree]20[minute] 05[second] N. 19[degree]45[minute] 36[second] and 20[degree]08[minute] 00[second] E) are situated at the border between Poland and Slovakia. The foot of the massif is determined by a contour line of 900 m a.s.l., and the highest peak is Gierlach (2655 m a.s.l.), lying on the Slovak part of the mountains. The total area of the Tatra Mts. covers about 785 km2, of which almost 175 km2 (about 25% of area) are within the territory of Poland, and 610 km2 (75%) in Slovakia. That part of the Tatra Mts., which belongs to Poland, is named the Polish Tatras (TP) and occupies mostly the northern slopes of these mountains. Their eastern part comprises western areas of the High Tatra (TW) and their western part, the eastern areas of the Western Tatra (TZ). The whole of the southern part of the investigated area, together with part of the northern slopes in the regions of Spisz and Orawa, belong to the Slovakian Tatras (TS). They include the whole of the Belianske Tatra Mountains (TB) and most part of the TW and TZ (Fig. 1). The main part of the Tatra Mountains is formed of a so-called crystalline core, composed of granite and older metamorphic rock (gneiss, amphibolite, shale etc). TW are built mainly of granite, while most part of the TZ ridge and their southern slopes, of rock (including also two belts of granite). The TB, forest belts of the TW and big fragments of the northern slopes and main ridge of the TZ have been modelled in sedimentary rock. mostly in limestone and dolomite. During the Pleistocene period, the Tatra Mts. were repeatedly glaciated (three times), and the last glacier finally retreated about 10,000 years ago. Since the retreat of the glacier the surface features of the Tatra Mts. have been shaped by gravitation , activity of running water, wind and snow, and by karstic processes in areas built of carbonate rocks. The climate of the studied area is under the influence of the air masses of different geographical origin, and its characteristic feature is great variability of weather conditions. The average annual temperature changes in the vertical profile of the massif from +4 to[long dash]4[degree]C, which results in an average gradient of 0.5[degree] per 100 m of altitude. Together with an increase in altitude, a decrease in the annual temperature amplitude is observed. There is also a correlation between elevation a.s.l. and absolute minimum and maximum temperatures. as well as a correlation between the intensity of direct solar radiation and slope exposure aspect. In the Tatra Mountains five vegetation belts have been : lower montane belt (from the foothills to 1250 m a.s.l.), upper montane belt (1250-1550 m a.s.l.), subalpine (dwarf pine belt) (1550-1800 m a.s.l..), alpine belt (from 1800 to 2250-2300 in a.s.l..) and subnival belt extending from 2250-2300 m a.s.l. to the highest peaks. These belts differ in the climatic features as well as in the predominating plant communities and ranges of plant and animal species. On account of the unique natural values on the world scale, the Tatra Mts. together with the established earlier national parks (Slovakian TANAP [long dash] Tatransky Narodny Park and Polish TPN [long dash] Tatrzanski Park Narodowy) were in 1992 included, by the UNESCO-MAB, in the list of the World Biosphere Reserves, as the Tatra Biosphere Reserve. The study was focused on the weevil species of seven families (Nemonychidae, Rhynchitidae, Attelabidae, Apionidae, Nanophyidae, Erirhinidae, Curculionidae) from the suprafamily of Curculionoidea occurring in the whole area of the Tatra Mountains, except for their inaccessible parts. Weevils were collected at 531 localities (Table 1) representing the 23 selected biotopes, belonging to 5 groups of plant communities: forest communities (beech forests, alder forests, fir-spruce forests, lower montane spruce forests, upper montane spruce forests), brushwood communities (roadside brushwood, ligneous vegetation along streams, xerothermic shrubs, wind-falls, willow-rowan thickets, dwarf pine thickets), tall herb communities (roadside tall herb communities, hydrophilous tall herb communities along streams and around springs, tall herb communities of the lower montane belt), meadow communities (meadow communities of the lower montane belt showing no symptoms of anthropopressure, humid meadows of the lower montane belt, mown meadows of the lower montane belt, grazed meadows of the lower montane belt, meadows of the upper montane belt, subalpine meadows, alpine meadows) and other biotopes (gullies, rock grasslands). These are the full representation of the Tatra natural environment. Weevils were being caught using the standard methods in the faunistic-ecological studies of herbivorous beetles. In addition, were documented all other observations concerning occurrence (e.g. the type of biotope and place of the collection of individuals), food plants and breeding periods of these Insects. Altogether 28,282 individuals were examined. The factographic material originated from the two main sources: 1) national and foreign collections of the Tatra weeevils, examined by the author [long dash] 3,940 specimens, 2) the authors' own material [long dash] 24,342 individuals (Table 2). The study on Curculionoidea of the Tatra Mts. started in 1857 and have been continued for 142 years with different intensity (Tables 3 and 4). The most effective research was conducted in the 1980s and 1990s, when most species were collected. Though in other periods the collections of weevils were smaller, each survey brought new records for the Tatra Mts. and knowledge of the weevil fauna was gradually being improved (Fig. 2). In the whole area of the Tatra Mts. weevils were collected by 90 persons with different intensity in particular time periods and regions (Table 3). The numbers of publications concerning the weevils of different investigated regions in the Tatras are uneven; most of them deal with the fauna of the Polish Tatra (Table 4). The investigated fauna of the Tatra Mountains contains 303 species (Tables 7, 8 and 9). Of these, the occurrence of 51 species needs confirmation or updating, on the basis of new materials (Table 5). Further 68 species that are marked in the catalogue with symbols "-" and "[tautomer] ", in the author's opinion, should not be included in the list of the Tatra fauna (Table 6). During the studies, there were 75 new species to the investigated area found; of these, the present paper for the first time presents the following species: Anthonomus humeralis, A. rufus, Cionus hortulanus, Dorytomus tremulae, D. dejeani, D. hirtipennis, Isochnus angustifrons, Ramphus pulicarius, Tachyerges pseudostigma, Tychius breviusculus, T. crassirostris, T. junceus, T, stephensi, Calosirus terminatus, Mogulones asperifoliarum, Thamiocolus viduatus, Trichosirocalus barnevillei, Pelenomus comari, Rhinoncus perpendicularis, Dodecastichus mastix, Otiorhynchus ligustici, Otiorhynchus denigrator, Barypeithes pellucidus, Sitona ambiguus, S. waterhousei, Hypera postica, Limobius borealis, Larinus brevis, Lixus iridis, Magdalis armigera, M. rufa and Hylobius pinastri. The numbers of new species in the faunas of certain regions of the Tatra Mts. are still higher (Chapter 5). The faunas of particular regions of the Tatra Mountains differ from each other and some even considerably, which is connected with the differentiation of the natural environment (geological substratum, slope aspect, climate, vegetation) of these mountains. Values of the similarity between these faunas are relatively small, and the most similar faunas form four distinct groups: Slovakian High Tatra, Slovakian Western Tatra, Belianske Tatra, Polish Western Tatra and Polish High Tatra. Differences in similarity values between the faunas of all weevils (Fig. 3) and the faunas of only mountain species of the particular Tatra regions (Fig. 4) indicate that the ranges of lowland and mountain species are much differentiated in the investigated area. These results also suggest that the mountain fauna is less susceptible to the diversification of the natural environment of the massif than the lowland species, though the occurrence of some mountain species is also limited in that area. In the whole area of the Tatra Mountains there occur only 65 species (21.4% of the investigated fauna), while the distribution of the remaining 238 species (78.6%) is limited to one or several regions. Species that are common for all the regions belong to Apionidae, Erirhinidae and Curculionidae families. Weevils from the family Nanophyidae have not been found in the Belianske Tatra and Slovakian High Tatra. The representatives of Rhynchitidae occur only in the Slovakian High Tatra and Polish Western Tatra, Attelabidae [long dash] only in the Slovakian Western Tatra and Polish Western Tatra, and Nemonychidae [long dash] only in the Polish Western Tatra (Tables 8 and 9). The Slovakian Tatra host more weevil species than the Polish Tatra, comprising 87% of the investigated fauna, though there are no representatives of the families Nemonychidae and Rhynchitidae among them. As compared with the fauna of the Polish Tatra the fauna of the Slovakian Tatra is richer by 66 species and more diverse in terms of the analyzed parameters (Tables 8, 9 and 10). which is probably the effect of: 1) larger area of the Slovakian part of the massif (about 3/4 of the whole area of the Tatra), 2) greater diversity of biotopes in the Slovakian Tatra (3 geobotanical regions) and their greater floristic richness (due to the greater patchiness of microhabitats) and 3) more favourable climatic conditions (considerable part of the slopes of the Slovakian Tatra are exposed to the south, south-east or south-west, while the Polish Tatra cover only L' of the area of the mountains, comprise only 2 geobotanical units and most of their slopes face the north). The distribution of weevils in the Tatra Mts. has been influenced by: aspect of slopes, patchiness of plant communities, floristic richness, and diversity of microhabitat conditions, while geological substratum is of less importance. These factors determine the pattern of the occurrence of many species, particularly those of the stenotopic character and limitate their ranges in the investigated area (Tables 8-11). Among the weevils of the Tatra Mountains the highest diversity of taxa is characteristic of Curculionidae (81% of the investigated fauna) and Apionidae (15% of the fauna), while species representing the remaining 5 families constitute only 4% of the fauna (Table 5 and 9). In mountain areas some of them have evolved into separate subspecies, varieties or races. These are: Notaris acridulus montanus, Graptus triguttatus vau, Cionus longicollis montanus, Donus comatus var. carpathicus and Otiorhynchus scaber var. oblongus i 0. rugosus krattereri. Morphological, biological, ecological and faunistic diversity of the examined species is described in detail in Tables 8 and 11. In the Tatra Mountains, spatial diversity in the distribution of particular species and groups of weevils, is also manifested in the horizontal aspect (Tables: 8, 11 and 12; Fig.: 5-7), though these differences are not so apparent and unequivocal as in case of the vertical aspect. They are particularly well marked in the faunas of different biotopes within the particular vegetation belts, showing the patchy character of the distribution of weevils in these mountains. The investigated biotopes are characterized by different groups of weevils, and the proportions of lowland species in these groups are more variable than the proportions of mountain species. That indicates the Tatra environment exerts a stronger influence on the lowland species than on the mountain species, though the occurrence of certain mountain species is also limited to certain biotopes. In the Tatra Mts. the most favourable conditions for the lowland fauna exist in: xerothermic shrubs, meadows of the lower montane belt and roadside brushwood, and for the mountain species: in rock grassland, willow-rowan thickets, dwarf pine thickets and alpine meadows. On the basis of species similarity index value, the weevil faunas of different biotopes may be classified into several groups (Fig. 6), which shows the possible migration routes of species between different habitats of the Tatra Mts. Rock grasslands, dwarf pine thickets, gullies and willow-rowan thickets host the most valuable faunas of weevils. The relatively high faunistic values (FV 4) are represented also by the weevil assemblages of hydrophilous tall herb communities along streams and around springs, subalpine and alpine and spruce forests of the upper montane belt (Table 11). In the light of these results these biotopes have been acknowledged as the most valuable in the Tatra Mts. Therefore, they need greater care than the habitats of secondary, anthropogenic origin, which are characterized by the least faunistic values of Curculionoidea. A diversity of both climatic and geomorphological conditions, as well as a diversity of vegetation result in both the vertical differentiation of weevils distribution in the Tatra Mountains and the exceptional patchiness of their occurrence, even within one vegetation belt (Tables 13 and 14, Figs 8-13). In the Tatra Mts., together with increasing altitude a.s.l., there are observed changes in the fauna and a decrease in the number of species; that correlation is statistically significant (Fig. 11). As compared with the mountain weevils, the declining trend of the lowland species is stronger (Fig. 12). There are observed certain fluctuations in the species richness within particular montane belts (Fig. 10), which is probably the effect of the environment differentiation and different area of the compared belts. Generally, at higher elevation in the lower montane belt and in the upper montane belt, the number of species continuously diminishes. Only in the lower part of of the lower montane belt it initially grows, and in the subalpine belt and in the lower part of the alpine belt, fluctuates slightly. In the subnival belt the number of weevils gradually decreases, and at an altitude of 2630 m a.s.l. the investigated fauna reaches the maximum upper limit of its vertical range (Table 14, Figs 10 and 11). Together with the growth of altitude a.s.l., the numbers of mountain and lowland species show a similar declining trend. However, the faunistic richness of each of these two groups of weevils changes along the altitudinal gradient in a different way (Figs 10, 12 and 13), which indicates that the mountain species are better adapted to the severe conditions of the high mountain environment than the lowland ones. The species richness differs in particular vegetation belts (Figs 8 and 10). All the distinguished altidudinal belts of the Tatra Mts. are inhabited by: Otiorhynchus arcticus, O. coecus, O. nodosus and O. tenebricosus. Extensive vertical ranges are also characteristic of the other 13 species, which have not been collected only in the subnival belt. Most of them are mountain weevil species. Vertical ranges of the remaining 223 species are clearly limited to certain vegetation belts. The results indicate that altitude exerts a stronger effect on the lowland species than on the mountain ones. In the Tatra Mts. the zonal distribution of weevils manifests itself also in the qualitative composition of their assemblages. One may observes a gradual of rapid exchange of species of different adaptations and habitat selectivity along the altitudinal gradient, which results in the structural changes in the whole zoocenoses. The populations of mountain species show many specific adaptations (ecological, behavioral and morphological) to low temperatures, shorter vegetation period, strong winds and long duration of snow cover. The results obtained show that the vertical distribution of the weevils in the Tatra Mts. depends first of all on physical conditions of the environment, and primarily on changes in climatic factors altitudinal gradient. Plants are an equally important factor, as shown by certain weevils connected strongly with forest and brushwood habitats (Table 8), whose ranges are limited to the two mountain forest belts of the Tatras (Table 13). The investigated Tatra weevils feed on plants belonging to 47 families (Fig. 14), 162 genera and 91 species, which shows on the one hand, considerable diversity of the food base of weevils in the Tatra Mountains, and on the other hand, different food specializations of the Tatra weevils and a wide range of food taken by some species. Most weevil species (65) were collected on plants from the family Leguminosae, but these were exclusively lowland weevils. On other plants there live less species (Table 8, Fig. 14), though some of them constitute a basic food for the investigated fauna. These are the representatives of the families: Rosaceae (on which there feed 46 weevil species), Salicaceae (44). Compositae (43), Pinaceae, Betulaceae and Polygonaceae (po 31 ), Cruciferae (19), Geraniaceae and Umbelliferae (13 species each). Both lowland and mountain weevils live on these plants. Probably, these plants played a key part in the development of the present weevil fauna of the Tatra Mts. The results presented in Figures 14 and 15 show differences in food specialization and competition for certain plants between the lowland and mountain weevils. It seems that the competition is strongest for those plants (Ranunculaceae, Caryophylaceae, Campanulaceae, Violaceae and Liliaceae), on which there were collected similar numbers of species representing both faunistic groups of Curculionoidea. On plants, on which feed more lowland or more mountain weevils, the competition for food is probably smaller. Next, on plants on which there were found species representing only one of these groups, there is no competition between the lowland and mountain faunas. On the species of the families: Leguminosae, Euphorbiaceae, Lythraceae, Primulaceae, Alismataceae, Equisetacae, Papaveraceae, Guttiferae, Oenotheraceae, Halorrhagidaceae, Rhamnaceae, Empetraceae, Oleaceae, Potamogetonaceae and Iridaceae collected were only lowland species, and on the representatives of: Tiliaceae, Oxalidaceae, Convolvulaceae, Gentianaceae and Dipsacaceae, only mountain species (Fig. 14). Representatives of the weevil mountain fauna were found on vegetation representing 30 families of plants (Table 8, Fig. 14). Most of these weevils feed on species from the families Rosaceae and Compositae (22 weevil species each), as well as Polygonaceae (13) and Pinaceae (11). Other plants attracted less mountain weevils. The plants mentioned seem to constitute a basic food for the majority of mountain weevils in the investigated area. They probably played a key part in colonizing the Tatra Mts. by the investigated Curculionidea. Though in terms of species diversity trees yield precedence to herbaceous plants, they are an important food base for many weevils. Among them, particularly important is Picea abies, on which 28 weevil species were collected. Many other arboreous and herbaceous plants are equally important for them (Table 8). Most mountain weevil species occur in the Tatras on: Petasites spp., Alchemilla spp., Cirsium spp., Picea abies, Geranium spp., Rumex spp., Chaerophyllum hirsutum, Rubus idaeus, Geum spp., Salix spp., Betula spp. and Alnus spp. On: Polygonum bistorta, Rumex alpinus, Vaccinium myrtillus, Caltha palustris, Salvia spp., Doronicum austriacum, Geum montanum, Cerastium latifolium, Stellaria nemorum, Tanacetum spp., Veratrum lobelianum, Filipendula ulmaria, Tussilago spp., Dryas octopetala, Oxalis acetosella, Gentiana asclepiadea, Knautia spp., Convolvulus spp., Lathrea squamaria, Calamagrostis spp. and Dactylis glomerata there were found only mountain weevils. The largest trophic groups of weevils in the Tatra Mountains is composed of oligophagous species (159). The monophagous group is least numerous (46 species), while the polyphagous group comprises 98 species (Fig. 15). Quantitative proportions of these groups in the fauna of lowland species are similar, while the mountain weevils are dominated by polyphagous species. These results show that the occurrence of the majority of lowland weevils in the Tatra Mts. is connected with the occurrence of food plants; the distribution of a small number of species only is limited by other factors. In case of the mountain fauna, the distribution of most species is not so strictly connected with food plants, and their occurrence is limited mostly by abiotic factors. The distribution of only few mountain species is limited by the presence of determined food plants (Table 8, Fig. 15). That relatively small dependence of the mountain species on plants and the large proportion of hortophagous species in their fauna suggest that some of these weevils may have colonized the Tatra Mts. before trees and shrubs appeared in the area. Analyzing the characteristics of biodiversity (Table 8), one may find that in mountain areas the highest ecological tolerance and considerable colonization potential are shown by the following species: Anthonomus rubi, Barynotus obscurus, Dodecastichus inflatus, Otiorhynchus nodosus, O. tenebricosus, O. coecus, O. lepidopterus, O. proximus, O. scaber, O. equestris, O. obtusus, O. rugosus krattereri, O. pauxillus, Phyllobius arborator, Ph. glaucus, Polydrusus amoenus and Sciaphilus asperatus. The investigated fauna is a kind of conglomerate of species, differing in ecological requirements as well as in the area and shape of ranges (Chapter 5). Particular chorotypes have been distinguished on the basis of the presently known natural ranges of species, excluding areas where they have been introduced. The Holarctic range group includes one Holarctic element with 14 species belonging to 3 families (Tables 15 and 16). Their occurrence in the Tatra Mountains is confirmed by the common origin of part of the investigated fauna from the temperate areas of Eurasia and cold areas of North America. It is probably due to the different histories of the two continents, and especially different environmental conditions prevailing in the fauna refuges during the largest glaciations, that the percentage of Holarctic species (4.5% of the investigated fauna) is small in the Tatra Mts. To the Palaearctic range group there were assigned 229 species (75.5% fauna) belonging to 7 families of weevils. Because of differences in the shape and area of their Palaearctic ranges, they were reckoned among 23 separate geoelements (Tables 15 and 16). The differentiated zoogeographical structure of the present weevil fauna of the Tatra Mts. points to different origin of their representatives. Most Curculionoidea (81 species; 27% of the investigated fauna) came to the investigated area from the European refuges. A large group (48 species; 16%) is composed of weevils with wide Euro-Siberian ranges, including also the Tatra Mts. It seems that many of them spread also from European refuges reaching as far as Siberia. The present state of knowledge does not allow one to mention all species, but among them there are certainly representatives of the genera Otiorhynchus, Polydrusus and Trachyphloeus. The migration of some Euro-Siberian species probably has also an adverse direction, which concerns among others: Involvulus cupreus, Lepyrus volgensis and representatives of the genera Magdalis and Pissodes. The results (Tables 15 and 16) show also the members of these two chorotypes, and particularly European species played an important part in the formation of the present fauna of the Tatra weevils. The remaining range elements are less numerously represented in these mountains and some of them only by one species each. The proportions of species representing particular families of weevils in the distinguished chorotypes (Table 16) suggest that after the period of glaciations species belonging to Curculionidae may have appeared earliest in the Tatra Mts. Later on, the representatives of Apionidae and other families came to this area. Most of the lowland species (they belong to 21 distinguished chorotypes) occurring in the Tatra Mts. are characterized by extensive horizontal ranges, while their vertical ranges are relatively poorly differentiated (Chapter 5, Table 15). On the other hand, the mountain species have also small and often fragmented horizontal ranges. The vertical ranges of many of these species are much differentiated. Because of differences in horizontal ranges these weevils have been assigned to the boreal-mountain and mountain element. The boreal-mountain chorotype comprises 10 species (3.3% of investigated fauna) which are characterized by disjunctive ranges, oriented S-N. Among the mountain element there reckoned 49 species (16.2% of the fauna), distributed mainly in the mountain and sub-mountain areas; some of them occur locally also in uplands and lowlands (Table 15). On the basis of the adopted criteria of vertical ranges classification there were distinguished the following elements: lowland, including 174 species, collected mostly in the lower montane belt; lowland-mountain, 69 species; all-mountain, including Donus comatus, Liophloeus lentus, Otiorhynchus clavipes, O. coecus, O. equestris, O. kollari, O. lepidopterus, O. nodosus, O. proximus, O. rugosus krattereri, Polydrusus amoenus and Scleropterus serratus; mountain (forest), with Anthonomus conspersus, Ceutorhynchus pandellei, Dodecastichus injlatus, D. mastix, D. obsoletus, D. pulverulentus, Donus intermedius, D. ovalis, D. oxalis, D. palumbarius, D. velutinus, Dorytomus carpathicus, Graptus weberi, Leiosoma cribrum, Liparzrs glabrirostris, Lepyrus volgensis, Miarus abnormis, M. monticola, Mogulones angulicollis, Notaris aterrirnus, Onvxacalles croalicus, O. pyrenaeus, Otiorhynchus austriacus, O. bisulcatus, O. corvus, O. morio, O. multipunctatus, O. pauxillus, O. reichei, O. scaber, Phyllobius alpinus, Ph. scutellaris, Pissodes scabricollis, Plinthus sturmii, P. tischeri, Polydrusus fulvicornis, P. impar, P. pallidus, P. paradoxes, P. pilosus, Rhinomias forticornis and Rutidosoma fallax; subalpine element (Donus maculatus, Otiorhynchus denigrator and Trachystyphlus beigerae) and alpine element (Otiorhynchus arcticus and O. obtusus. Species representing the two last elements, are determined also as the high mountain elements. Together with an increasing distance from the Tatra Mountains, the number of mountain species decreases in all directions, however much more species are lost in meridional directions than in parallel ones (Fig. 16). These may indicate the directions of migrations of the postglacial mountain fauna of the Tatra Mts. From the point of view of the palaeogeography, the Tatras rather young mountains. Therefore, the investigated fauna has had not enough time to evolve spectacularly and is relatively young, mostly postglacial, as evidenced by the absence of endemic genera; the only Tatra endemite is Trachystyphlus beigerae, and there are only few Carpathian endemic species. The Tatra weevil fauna has developed over a long time period and is a product of many migration waves from different directions, repeated expansions and following contractions of the formerly occupied ranges. It is the complex history of the Tatra nature in the past geological epochs, particularly during climatic changes in the Pleistocene period, which has exerted the strongest influence on the structure of the present fauna. The results of the present study show that species from warmer interglacial periods have not survived in the Tatra Mts., and the oldest elements of the present fauna are represented by glacial relicts related to the last glaciations. The retreat of the glaciers, conducted to the disruption of ranges of the ecologically specialized alpine species, as well as arctic-alpine. or boreal-alpine ones. Freed from the glacier, the area of the Tatra Mts. was first colonized by arctic-alpine, boreal-mountain, alpine and mountain species. Only later on, there came species from the European lowland, far Siberia and other areas of the Palearctics and Holarctics. At last, also southern species appeared in the Tatra Mts. from sun-heated areas, though not numerously. The present studies allow one to draw the following conclusions: 1. The Tatra Mountains may have been colonized easiest and quickest by species that were simultaneously eurytopic, parthenogenetic, polyphagous and lying eggs to the ground: colonization was much more difficult for stenotopic, bisexual and monophagous species. 2. The process of colonization of the area of the Tatra Mts. probably followed the model: favorable abiotic conditions [forward arrow] preferred plant community [forward arrow], presence of host plant species (at least one) [forward arrow] attempt at the colonization of the area by a species [forward arrow] adaptation to new environmental conditions [forward arrow] possibility of the development of a given population and influence of other organisms [forward arrow] colonization of a new area (one may speak about the full colonization when in a given area there develop successive generations of a species). 3. Colonization of the studied area by weevils went on as follows: 1) species of subnival and alpine belts, subalpine species, 2) forest and brushwood species of the upper montane belt, 3) forest and tall-herb species of the lower montane belt, 4) species of open areas of the lower montane belt, originated as a result of human activity. 4. Weevils feeding on pioneer grasses and tall-herbs may probably have spread quicker towards the Tatra Mts. than species which are trophically related to trees; however, it was not a rule because the process of colonization was governed by other environmental factors. The main features of the investigated fauna were established in the Holocene period, and its present structure is an effect of a long-term influence of primarily abiotic factors and next, biotic ones. Colonization of the studied massif by present weevil species was certainly connected with the all-European postglacial recolonization by plants and animals. From the southern glacial refuges they probably reached the Tatra Mts. at different times. The populations of similar ecological characteristics and distribution belonged undoubtedly to the same migration wave, while species which differed markedly from each other in this respect and much distant from each other in time and space, undoubtedly used different migration routes. It seems that after the period of glaciations, the Tatra Mts. were colonized by mountain weevils simultaneously from many directions. However, there were two main migration routes: alpine, from SW and W parts of Europe along the western margins of the Carpathians, and Balkan-Carpathians, from SE and E regions of the continent, at first along the Southern Carpathians, and next along both slopes of the Eastern Carpathians. Probably, the initial stage these migrations went on mainly along the southern slopes of the Carpathian arch, and only later on, a route on the northern slopes of the massif was initiated. Other important route was also the Transcarpathian migrational track (S-N), leading along the depressions and river valleys of the Carpathians. Less important for the development of the present mountain fauna of the Tatra Mts. may have been migrations from SW direction, originated in the Pyrenees, and from S direction, with the centre of dispersion in Apennine Peninsula. A relatively small proportion of the Caucasian and Siberian species in the Tatra mountain fauna seems to indicate that the influence of the Caucasian fauna on the Carpathian fauna, and the Carpathian fauna on the Siberian fauna, was rather small. Probably a considerable distance and serious geographical and ecological barriers, originated after the period of Pleistocene glaciations, and perhaps also other causes in the earlier geological periods, limited migrations between these regions. It seems that parthenogenetic species from the genus Otiorhynchus, occurring in the Tatra Mts., originated from the southern and south-western regions of Europe. Probably from the south-European and Asiatic refuges there may have come to the Tatra Mts. also some species from the genera: Betulapion, Archarius, Dorytomus, Ellescus, Isochnus, Phyllobius, Polydrusus, Chlorophanus and Magdalis, and only from the southern regions of Europe, representatives of the genera Strophosoma and Trachyphloeus. Some mountain species, such as Mogulones angulicollis, Rutidosoma fallax, or species representing the genus Dodecastichus, for example, reached the Tatra Mts. probably even from the areas belonging to the Mediterranean Sea basin. Of the Asiatic origin there are undoubtedly the representatives of the families: Nemonychidae, Rhynchitidae and Attelabidae and some species from the genera: Apion, Perapion, Ceratapion, Omphalapion, Eutrichapion, Hemitrichapion, Holotrichapion, Oxystoma, Catapion, Ischnopterapion, Squamapion, Caanapion, Stenopterapion, Protapion, Pseudoprotapion, Miarus, Cionus, Ceutorhynchus, Glocianus, Trichosirocalus, Mogulones, Amalus, Microplontus, Thamiocolus, Tapeinotus, Pelenomus, Ramphus, Tychius, Brachysomus, Eusomus, Orobitis, Sitona and Flypera, and also the representatives of the subfamily Lixinae. Migrations of these weevils may have been accompanied by some Holarctic species from the genera: Grypus, Acalyptus, Tachyerges, Auleutes, Cryptorhynchus, Limnobaris, Lepyrus and Pissodes, which may have spread through Bering Strait when the land passage existed between Asia and North America during the period of great Pleistocene migrations. Notaris aterrimus, Miarus abnormis and M. monticola, species connected with meadows, colonized the investigated area mostly from refuges in Balkan Peninsula and Turkey. That expansion was particularly quick in the eastern regions. The postglacial expansion of alder and beech from the north-Balkan refuges to the north was similarly quick. Therefore, one may assume the similarity of the migration routes of some species connected with alder forests and beech forests in the Tatra Mts. To these species belong: Temnocerus tomentosus, Curculio betulae, Anoplus roboris, Rhynchaenus fagi, Tachyerges pseudostigma, T. stigma, Cryptorhynchus lapathi, Acalles camelus, Onyxacalles croaticus, Polydrusus cervinus, P. amoenus, Rhinomias forticornis, Chlorophanus viridis, Phyllobius glaucus, Ph. maculicornis, Polydrusus pilosus, P. fulvicornis, P. picus and P. moths. Forest species were colonizing Central Europe, and the Tatra Mountains, from two directions: Iberian Peninsula and Carpathian-Caucasian refuges. The model of colonization of the brushwood species is completely different from that of the meadow and forest species. This model assumes that Europe had been colonized from three glacial refuges: Iberian, Appeninian and Balkan. The similar model of colonization was probably characteristic of Phyllobius glaucus, Polydrusus impar, P. pilosus and Pissodes piceae, foraging also on fir, whose genomes in some southern refuges indicate expansion to the north. These weevil species are, however, polyphagous, hence they may have participated also in other migration waves. Migrations had probably also the opposite direction and some species, as e.g. Otiorhynchus nodosus and O. equestris may have migrate from the Balkans and the Alps to as far as Siberia where they have occurred until now. For most of the mountain element the periods of climatic changes were probably less important though some species, such as: Otiorhynchus scaber, Rutidosoma. fallax, Liparus glabrirostris, and other species migrated also far beyond the mountain areas. Among the studied fauna, Otiorhynchus scaber and probably O. lepidopterus, and also other parthenogenetic weevil species, such as 0. pauxillus, Barynotus obscurus and B. moerens originated from the area of the Alps. It seems that the first three species and Pissodes scabricollis came to the Tatra Mts. together with spruce forests, and the next two species, and Polydrusus paradoxus - with tall-herb communities. Liophloeus tessulatus reached the area of the Tatra Mts. probably also from the Alps, or even from the Pyrenees. Otiorhynchus morio has certainly originated from the Pyrenee Mountains. Strophosoma melanogrammum, Sciaphilus asperatus and Trachyphloeus bifoveolatus, parthenogenetic species in most part of their ranges colonized the Tatra Mts. probably from the south-western regions of Europe. After the retreat of Pleistocene glaciations the first two species migrated to the east with some brushwood and forest communities, and T. bifoveolatus, with herbaceous plants. Polydrusus mollis, Tropiphorus elevatus and T. terricola came undoubtedly from the Balkans. From Balkan Peninsula to the Tatra Mts. migrated probably also some bisexual mountain species, such as: Notaris aterrimus, Miarus abnormis, M. monticola, Onyxacalles croaticus, Dodecastichus pulverulentus, D. mastix, D. inflatus, D. obsoletus, Otiorhynchus bisulcatus, O. rugosus, O. reichei, O. denigrator, O. austriacus, O. kollari, Phyllobius scutellaris, Rhinomias forticornis, Donus comatus,.D. maculates. D. ovalis, D. oxalis, D. palumbarius and Plinthus sturmii. Otiorhynchus rugosus came to the investigated area probably from one of the Carpathian refuges. The other representatives of the present mountain fauna of the Tatra migrated only between the mountains of Europe. Of these, Anthonomus conspersus, Ceutorhynchus pande/lei, Mogulones angulicollis, Rutidosoma fallax, Onyxacalles pyrenaeus, Graptus weberi, Otiorhynchus tenebricosus, O. multipunctatus, O. coecus, Phyllobius alpinus, Liophloeus lentus, Polydrusus impar, P. pallidus, Donus intermedius, Liparus glabrirostris and Leiosoma cribrum reached farthest. However, the present state of knowledge does not allow one to indicate the routes of the postglacial colonization by these species. Otiorhynchus ligustici, which is parthenogenetic in the Tatra Mts., probably has come from the Eastern Carpathians and perhaps also from the Balkans. It seems that Hylobius pinastri and Dorytomus nordenskioldi colonized the Tatra Mts. from the northern regions of Europe, and their migration to the the south occurred together with the migration wave of the boreal vegetation. Eusomus ovulum, Sitona languidus and Mogulones austriacus and perhaps also Tychius crassirostris, termophilous species, reached the investigated massif from Pontic refuges, and Larinus brevis from Ponto-Mediterranean regions. The colonization of the Tatra Mts. by these xerothermic weevils is chronologically one of the latest. However, not all Tatra weevils extended so spectacularly their ranges during the Pleistocene and Holocene periods. Many European forest species remained in the Carpathian and sub-Carpathian localities, e.g. Carpathain-Sudetic (Plinthus tischeri) and Carpathian (Dorytomus carpathicus, Otiorhynchus corvus, O. proximus, Donus velutinus and Trachystyphlus heigerae), as evidenced by the fact that these species are absent in the fossil coleopterofana in lowland localities of Poland and other countries. Colonization of the Tatra Mts. by Otiorhynchus proximus started probably from the south-Carpathian refuges where there occur its bisexual populations. Trachystyphlus beigerae, the only one endemite in the investigated fauna, has undoubtedly originated from one of many sister lines of the alpine complex of T. alpinum; it probably had survived the period of Pleistocene in one of the Carpathian refuges. The first traces of some species occurring currently in the studied mountains are dated in Europe at Eocene period (55-36 min years ago). These are mainly forest species from the families Rhynchitidae and Attelabidae and some species of several subfamilies of Curculionidae (Table 17). That fauna probably differed considerably from the present one and resembled the fauna which is currently known from the mountain areas of China. In the Oligocene period (36-25 m a thousand years ago) the European fauna was already very rich and diverse in terms of both taxonomy and ecology, and included among others the following genera, represented in the present fauna of the Tatra Mts. by: Involvulus, Neocoenorrhinus, Apion, Nanophyes, Notaris, Curculio, Cionus, Dorytomus, Tychius, Sibinia, Bagous, Baris, Ceutorhynchus, Coeliodes, Cryptorhynchus, Acalles, Otiorhynchus, Phyllobius, Polydrusus, Sitona, Donus, Hypera, Larinus, Lixus, Liparus, Magdalis, Plinthus, Hylobius and Pissodes (Table 17). Had in those times existed any old preglacial mountain faunas in the area of the Tatra Mts., they must have receded or been inundated by warm seas during the Upper Eocene and Oligocene periods. Currently, there are no premises indicating what kind of faunas they could be. It seems that the colonization of mountain areas by the representatives of the present European fauna may have occurred at the end of the Palaeogene and in the lower Neogene. In that period, as the result of the folding of the Flysch Carpathians and uplift movements in the Central Carpathians, the sea had definitively retreated from the Tatra Mts. After the alpine orogenesis, the first mountain range ready for colonization was the Pyrenees, next the Carpathians, while the Alps and Caucasus were colonized not earlier than in the Quaternary. At the end of the Pliocene, the fauna of the Tatra weevils may have already been diverse, as indicated by the rich fossil materials of weevils, dated at the Upper Miocene, found in the different parts of Europe. Those materials comprised specimens of many genera represented in the investigated fauna, e.g.: Apion, Curculio, Cionus, Tychius, Bagous, Cryptorhynchus, Otiorhynchus, Sitona, Donus, Hypera, Larinus, Lixus, Plinthus, Hylobius and Pissodes (Table 17). However, it is worthy of mention that the European Tertiary floras and consequently, probably also the assemblages of weevils, differed in composition. At the end of the Neogene period (1.8 m1n years ago), the climatic and vegetation belts were already developed and the typical mountain fauna may have already occurred. Probably in those times also the high mountain fauna had developed. Many premises suggest that the contemporary fauna of the Tatra Mts., except for a few genera, resembled the Holocene fauna. However, not all representatives of that fauna have survived to our times due to subsequent, drastic environmental changes, which occurred in the Pleistocene. In the Pleistocene period (1.8 min [long dash] 10-25 thousand years ago), as the result of a general decrease in temperature in the Carpathians and in other mountains of Europe (e.g.: Alps, Pyrenees, Sudetes, Balkans), and the slow accumulation of mountain glaciers, the preglacial fauna must have underwent almost total extermination or it withdrew to the south- and east-European refuges, mainly to peninsulas: Iberian, Appeninian, Balkan and Crimean and to some part of the Caucasus and the area of the Caspian Sea. Its place was taken by cold-loving assemblages, represented currently in the Tatra Mts. by arctic and boreal species, coming from the northern part of the Holarctics and higher parts of mountains. It seems that of the elements of the Tertiary fauna, only Trachvstyphlus beigerae and Otiorhynchus obtusus, Carpathian species, resistant to low temperatures, may have survived during the unfavorable period in some places in the Tatra Mts. Specific climatic conditions, changing in glacial and interglacial, and stadial and interstadial (particularly changes in humidity, and to a lesser degree in temperature) periods, triggered the phenomenon of migrations. During the Pleistocene period, the biggest area were colonized mainly by boreal-mountain elements, such as Otiorhynchus nodosus, O. equestris, O. arcticus, O. lepidopterus, Polydrusus pilosus, P. fulvicornis, P. amoenus, Lepyrus volgensis and Scleropterus serratus, as well as some mountain species: Rhinomias forticornis and Notaris aterrimus. It was probably during the Krakow glaciation when Lepyrus volgensis, the only representative of the Arctic fauna, which in the Tatra Mts. has survived until now, reached the south of Europe from the arctic area. It undoubtedly met with the periglacial high mountain fauna, represented by Otiorhynchus arcticus and O. nodosus, and perhaps also with O. obtusus and Trachystyphlus beigerae. In those times in the Carpathians there occurred also: Otiorhynchus multipunctatus, O. cornicinus, O. coecus, O. kollari, O. equestris, 0. rugosus and O. corvus. in the interglacial periods of warmer climate, environmental conditions prevailing in the Tatra Mountains, were probably more favorable for weevils than the present ones and more thermophilous and dendrophilous species occurred there. It seems that in those times the mountains may have already been colonized by some of the genera represented in the present fauna: Otiorhynchus, Phyllobius, Polydrusus, Rhinocylus, Anthonomus, Hylobius, Magdalis and Pissodes. Towards the close of interglacials, the climate oscillated and climatic conditions had been worsening anew. A gradual decline of forests created conditions for the expansion of herbaceous plants, dominated by grasses mugwarts and sedges, and at some localities there occurred also Caryophyllaceae, Chenopodiaceae, Cruciferae, Compositae and Ericaceae. Those conditions may have been conducive to the occurrence of some representatives of the weevil genera: Micrelus, Notaris, Limnobaris, Donus and species from the tribe Ceutorhynchini, represented in the present fauna of the Tatra Mts. On the other hand, during the glacial periods those weevils withdrew to the lower situations of the Carpathians. At the end of the glacial period (13[long dash]11 thousand years ago) the Tatra Mountains were still dominated by tundra; there were no trees and lower parts of the mountains were occupied by non-forest communities with the representatives of: Graminae, Cyperaceae, Artemisia spp., Chenopodiaceae, Gentiana spp., Saxifraga spp., Helianthemum spp., Selaginella spp. and Polemonium spp., which together with trees and shrubs that came later, entered in the composition of the forest tundra. On the other hand, the arctic-mountain tundra extended at higher situations. It seems that in those times both belts mentioned may have been colonized by: Otiorhynchus arcticus, O. obtusus, O. nodosus, O. coecus, Lepyrus volgensis and Trachystyphlus beigerae, species most resistant to extreme thermal conditions. During the last interstadial (allerod, 11.8-11 thousand years ago) the main plant communities of the Tatra Mts. were pine forest and dwarf pine thickets, predominating in the area above the upper forest limit. The weevils already living in that area has been extended their vertical ranges, and others, such as: Otiorhynchus multipunctatus, O. kollari, O. equestris, O. rugosus krattereri and O. corvus started to colonize the Tatra Mts. During the younger Dryas (about 11 thousand years ago) a cold and humid period came. Many weevil species must have retreated from the Tatra Mts. and later on, returned by to the area. During the period of vistulian, environmental conditions in some parts of the Carpathians and their foreland were favorable to the migrations of hygrophilous, and simultaneously cold-loving fauna, represented currently in the Tatra Mts. by some species from the genera: Notaris, Dorytomus, Otiorhynchus, Isochnus, Donus, Ceutorhynchus and Ellescus bipunctatus. During the interglacial periods of warmer climate, mountain areas were colonized or recolonized by the forest and brushwood coleopterofauna, and also by water and riverside, or even marshy faunas. At the beginning of the Holocene, in the preboreal (10.25-9.1 thousand years ago) and boreal (9.1-7.7 thousand years ago) periods, the mountain glaciers of the Pyrenees, Alps and Carpathians, and of other mountains of Europe shrank or completely retreated from certain places. Freed from the glaciers, the area of the Tatra Mts. was first colonized by high mountain (alpine), arctic-alpine, mountain and boreal-mountain species. They were followed by species from the European lowland, far Siberia and other areas of Palearctics and Holarctics. At last, there appeared in the Tatra Mts., though not numerously, southern species and species from sun-heated areas. A separate group of species consisted of the already mentioned preglacial species, which either recolonized the Tatra Mts. or had survived the most difficult periods somewhere on the southern slopes of these mountains. In that period, the expansion of coniferous stands was accompanied by the expansion of mountain and boreal weevil species from the glacial refuges. Some of them have now disjunctive ranges and represent the Euro-Siberian zoogeographical element. Currently, these in species are characteristic of the upper montane belt and some patches of the lower montane (belt where spruce forests occur (Table 13). At that time, the Tatra Mountains remained under the big influence of the old autochthonic mountain fauna, and particularly the Carpathian fauna, with: Trachystyphlus beigerae, Otiorhynchus obtusus, O. corvus, O. proximus and Donus velutinus. However, there already existed suitable conditions for colonization of that area by: Otiorhynchus coecus, O. rugosus krattereri. Otiorhynchus equestris, O. kollari, O. lepidopterus and by other species, such as representatives of the genus Donus. Together with the expansion of spruce forests these mountains were gradually being colonized by other species, particularly those connected with spruce biotopes, such as: Anthonomus phyllocola, A. pinivorax, Rhyncolus ater, Onyxacalles pyrenaeus, O. tenebricosus, O. scaber, Phyllobius arborator, Ph. glaucus, Polydrusus impar, P. pallidus, Magdalis violaceus, M. barbicornis, M. duplirata, M. nitida, M. phlegmatica, Trachodes hispidus, Hylobius excavatus, H. abietis, H. pinastri, Pissodes pini, P. harcyniae, P. piceae and P. scabricollis. The contemporary re birch groves and birch-pine forests may have been colonized by: Temnocerus tomentosus, Deporaus betulae, Betulapion simile, Coeliodes rubicundus, Rhamphus pulicarius, Rhynchaenus jota, Tachyerges pseudostigma, T. stigma, Polydrusus pilosus, P. fulvicornis and P. amoenus. In the similar period there came also: Cimberis attelaboides, Magdalis rufa, Pissodes castaneus and P. piniphilus, species living on pines. Then, in the Tatra Mts., the first peat bogs originated on e.g. the meadow of Molkowka, Rowieri Waksmundzka and Hala Ornaczanska and others which could be occupied by species from the genera: Notaris, Limnobaris and Bagous. During the Atlantic period (7.7-5.1 thousand years ago), which was characterized by a warm and humid climate, even warmer than at the present time, a considerable number of species characteristic of forest and tall-herb habitats, as well as many termophilous species, particularly Ponto-Pannonic and Mediterranean, invaded the Tatra Mts. (Table 15). On the other hand, cold-loving weevils must have moved to the highest parts of the mountains. It seems that in those times the Tatra Mountains were colonized by most species occurring there now and that the contemporary fauna was undoubtedly much richer and more diverse than the present one. In the subboreal period (5.1-2.3 thousand years ago) a climate got cooler and more humid, which resulted in the lowering of the upper forest limit and a change in the predominating forest communities. Probably in that period, together with beech and alder forests, there came the following species: Curculio betulae, Anophus roboris, Rhynchaenus fagi, Rh. lonicerae, Acalles camelus, Onyxacalles croaticus, Dodecastichus inflatus, Phyllobius maculicornis, Ph. oblongus, Polydrusus picus, P. mollis, P. cervinus and Magdalis armigera. At the end of that phase (3.6-3 thousand years ago) all vegetation belts, distinguished currently, had already been formed. The subsequent natural changes in their ranges have not affected significantly the present fauna of weevils. In the subatlantic period (2.3-0 thousand years ago), in the natural environment of the Tatra Mountains no spectacular changes, such as those in former periods, did occur and the contemporary fauna resembled the present one. Historical factors and effects of human activity, particularly during the past two centuries, were also of a big importance for the development of the present fauna. A response to changes in forest communities was the total extermination or considerable reduction in the populations of some species connected mainly with beech forests, e.g. Rhynchaenus fagi, Polydrusus pterygomalis and P. mollis, as well as colonization of the clear cut areas by, above all, Otiorhynchus coecus, O. tenebricosus, O. nodosus and O. scaber, weevils usually connected with spruce forests. Deforestation of large area and creation of ruderal habitats, which had not occurred earlier, created suitable conditions for colonization of the Tatra Mountains by many species preferring non-forest habitats, such as: Apion cruentatum, A. frumentarium, Catapion seniculus, Ceratapion onopordi, Ceutorhynchus erysimi, C. typhae, C. obstrictus, C. pallidactylus, Hypera diversipunctata, H. plantaginis, Nedyus quadrimaculatus, Oxystoma cerdo, Perapion curtirostre, P. marchicum, P. violaceum, Phyllobius oblongus, Ph. vespertinus, Protapion apricans, Rhinoncus pericarpius, Sitona hispidulus, S. humeralis, S. puncticollis, S. suturalis, Squamapion atomarium, Tychius lineatulus, T. meliloti, T. picirostris, T. stephensi and Z. geranii. In the 19th century and at the beginning of the 20th century these weevils were still not found in the Tatra Mts. The creation of open areas within mountain forests make also the expansion of Donus comatus, D. ovalis, D. velutinus, Otiorhynchus equestris, O. kollari and O. rugosus krattereri easier. These species, from the formerly occupied tall herb communities, along streams and around springs, gullies, wind-falls, and alpine meadows among the dwarf mountain pine thickets, invaded and dominated the newly created habitats, through which they could spread farther. The effects of human activity (anthropogenic factor) on the Tatra nature are also the present symptoms of the synanthropization of the investigated fauna. This phenomenon may be attributed to the synanthropization of certain habitats and the appearance of plants, considered currently as synanthropic ones. The appearance of, among others, Melilotus albus in the Tatra Mts. made easier the colonization of the area by the following weevils: Holotrichapion pullum, Stenopterapion meliloti, Tychius breviusculus, Tychius crassirostris, T. junceus, T. meliloti, Sitona cylindricollis, Hypera postica, H. suspiciosa and H. venusta. Most of these species occur now only in the lower montane belt, in the neighborhood of anthropogenic habitats (e.g. roadsides, surroundings of tourist shelters and other constructions, etc.). The present fauna of Curculionoidea of the Tatra Mountains is, generally, a stable fauna. About 3/4 of the species are permanently connected with the area and in the nearest future should also be stable; only 1 /4 of that fauna need monitoring (Tables 3 and 4). The most distinctive feature of this fauna is its high mountain character. As compared with the weevil faunas of the other Carpathian ranges and other areas of Poland, the Tatra Mountains host most of all (59) mountain species (Fig. 16). The Tatra Mountains are the largest centre of the mountain weevil fauna in the Western Carpathians and at least for that reason the range is of great importance for conservation of the present biodiversity. This importance stems from: 1) the presence of many high mountain species and forms and 2) the zonal distribution of zoocenozes, corresponding more or less to the vegetation belts. This feature distinguishes the investigated fauna from the weevil faunas of other Carpathian ranges. Among other geographic regions, the individual character of the Tatra Mts. is underlined by the presence of: Trachystyphlus beigerae [long dash] Tatra endemite and: Dorytomus carpathicus, Otiorhynchus corvus, O. obtusus, O. proximus and Donus velutinus [long dash] Carpathian endemites or subendemites. Only the weevils of the Babia Gora Mountain also show certain high mountain features. Due to the specific mountain conditions of the Tatra Mts., the occurrence of many weevil species is very local and often limited to a single valley, gully, or meadow. An important place in the Tatra fauna is occupied also by forest weevils, related mostly to tree and shrub species. These are mostly relics of the old Carpathian forest, e.g. representatives of the genera: Hylobius, Pissodes, Rhyncolus, Rhinomias, Otiorhynchus, Phyllobius, Polydrusus and Strophosoma (Table 8). In that fauna not less important are relict species related to tall herb communities along streams and around well-heads, wind falls, and open forest and brushwood stands or with originated in a natural way, small patches of tall herb meadows. This fauna is represented mainly by the representatives of the genus Donus and other genera, such as: Graptus, Liparus, Plinthus, Scleropterus, Trachystyphlus and Tropiphorus, and also by some species of the genera Ceutorhynchus and Otiorhynchus (Table 8). Most of them occur also in many meadows that are currently not grazed and overgrown by perennial plants, or in specific ecotone communities at the border between meadows and forests, dominated by herbaceous vegetation. This work does not bring studies on the diversity of the Tatra weevils to an end. It only summarizes the scientific output in that field, indicating problems to be explored in the future.

Notes:

Roznorodnosc, ekologia i chorologia ryjkowcow rezerwatu biosfery "Tatry" (Coleoptera: Curculionidea).

Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith