THE ROLE OF INSECTS IN POLLINATION AND DISSEMINATION OF SOME PLANT SPECIES IN HIGH-MOUNTAINS..

On the example of 5 species of herbaceous flowering plants of the Ukrainian Carpathians it is shown that they are in close relationships with a number of insects, some of which are involved into pollination and hence in the formation of mature seeds. It was found out that the species composition of insects does not depend on plant species composition, but on the ecological conditions of their habitat. The highest number of insects is connected with Dipsacaceae (30 species) and Asteraceae plants (29 species), less with Apiaceae (21 species), and the least with Boraginaceae (13 species). A significant role of phytophagous insects in the reproduction of plant populations was established. For example, Senecio carpaticus is one of species that depends significantly on the amount of received damage from the leaf beetles (Chrysomelidae) in the higher parts of the Ukrainian Carpathians. By the example of a number of Formicidae species, the role of myrmecochory in plants seeds dissemination is shown. It was confirmed that successful reproduction of Pulmonaria filarszkyana mainly depends on the presence of ants which disperse the species’ seeds over large area. Formica cunicularia, F. lemani, F. cinerea, Myrmica ruginodis, Tetramorium caespitum dominate among the ant species mentioned above.


INTRODUCTION
It is known that heterotrophic organisms have both positive (pollination and exchange of genetic information between the populations) and negative effect on the reproduction of plants (damaging vegetative and generative parts of the plants). The study on the role of heterotrophic organisms in plant pollination, seeds dispersal and damage to the reproductive organs of plants is a matter of particular interest.
The aim of the study was to establish the quantitative and qualitative composition of insects that are involved in the reproduction of plants, to determine the characteristics and importance of identified animals in their reproduction, and to clarify the role of ants in the dissemination of seeds of some high-mountain plant species in the Ukrainian Carpathians. Biol Long-term research of entomophilous plants and their pollinators' relationships on the example of some families of angiosperms showed the important role of flower morphology known as "the length of corolla tube" [3,4]. The family Compositae is differentiated into species by inflorescence colour and length of flowers' corolla tube [3]. Species with white and yellow or yellow inflorescence have short corolla tube; insects with short proboscis (Diptera) dominate among the pollinators. Species with purple-violet inflorescence have long corolla tube; insects with long proboscis, primarily bees, dominate among the pollinators. The relevance of our work is about continuing research in this field through the example of consortial relationships of teasel plant family (Dipsacaceae) and other phylogenetically unrelated high-mountain plant families of the Ukrainian Carpathians [7,8,[11][12][13].
Myrmecochory is one of the most interesting examples of the relationships between plants and insects. While seeds of plants have insignificant role in trophics of ants, such an aspect of herbaceous plants usage by ants resulted in important co-evolutionary effect, i.e. the origin of myrmecochory [1,15]. Myrmecochory is characterized by the availa bility of specialties both in seeds morphology and instincts of ants, which are seed collectors mostly. In total the relationships between ants and plants are quite peculiar and aren't limited by collecting seeds or fruits only. Some plant species are "cultivated" by ants for aphids, which live on the plants. In some cases ants consume nectar and can pollinate flowers, but there is a lack of available information about the role of Formicidae in pollination for nowadays. At least it is known that species of Formica and Leptothorax genera pollinate some species of such families as Brassicaceae Burnett, Boraginaceae Juss. and Apiaceae Lindl. in alpine zone. The plant species have favorable morphological and biological features to be pollinated by ants, e.g. small simple flowers, scarce amount of nectar to compensate energy costs of flying pollinators etc. [6]. The investigations show that Formica lemani Bondroit, 1917 and F. picea Nylander, 1846 are active pollinators of Vaccinium myrtillus L., V. vitis-idaea L., and V. uliginosum L. [2].
To achieve the aim the following objectives were set: to identify the groups of insects that have consortial relationships with reproductive sphere of plants, to specify the activity of the studied animals (myrmecochory, entomophily, etc.), to evaluate the impact of animals on generative sphere of plants, and to establish specific insect taxa that have relationship to the studied plant species.
The resulting information will fill a significant gap in the variety of plants' pollinators in the Ukrainian Carpathians, since the data contained in the literature on pollinators of the families Compositae and Dipsacaceae are mainly related to one species of plants [9], and references to pollinators of most species of plants are fragmentary.

MATERIALS AND METHODS
Case study was conducted on 5 species of plants that grow in different habitats at the altitude of 1350-1700 m a.s.l. and belong to the different families. In particular they are The collection of insect pollinators was made by means of entomological net according to the methodology proposed by K. Fasulati [5]. Mirmecochore relationships of the plants were studied by the standard methods that intend to locate the seeds on 100 sq cm area with the following observation of their dispersal by ants [10].
Ratio between the specimen number of the certain insect species and the total number of counted individuals was determined using the system proposed by W. Tischler [14].

RESULTS AND DISCUSSION
Phoric relationships conducted mainly with the activity of insect pollinators are at the forefront among all the consortial relationships as they are tightly combined with the trophic ones. We found out 30 species of insects for the plants of family Dipsacaceae, 29 ones for the family Asteraceae, 21 species for Apiaceae and 13 insect species for Boraginaceae altogether (see Table).  All the insect visitors of these plant species are presented by anthophilous (60 %) and phytophagous species (40 %); the latter ones are mostly represented by leaf beetles of the genus Chrysochloa. The species depends a lot on the amount of received damage from the Chrysochloa beetles (See Figure). The generative sphere of plants of the family Apiaceae, which are exemplified by Astrantia major were visited by 21 species of insects. The dominant species are Eristalis tenax, E. rupium, Cheilosia impressa and others. Recedative and minor species are Bombus pyrenaeus, B. campestris, Sphaerophora scripta, Eupeodes corolla and others, which occurred with the frequency of 0.03-0.07.
During the research it was found out that A. montana consorts include 13 species, 85 % of which are anthophilous. They are as follows: Bombus lucorum, B. pratorum, and hoverflies of the genus Eristalis. 15 % of the species are presented by butterflies, namely Gonepteryx rhamni and Erebia medusa.
Lower number of insect species (11 ones) is typical for Pulmonaria filarszkyana due to the long flower corolla tube that is inaccessible for many insect species. Among the species observed there are typically polytrophic insects as Sphecodes gibbus, Heriades truncorum, Bombus lucorum, B. pratorum and the species-specific insects as Andrena combinata and A. dorsata with a visit frequency of 0.04-0.06. It was found out also that renewal of P. filarszkyana is tightly connected not only with typical insect pollinators (reproductive process), but also with the significant activity of ants, that disperse seeds for a considerable distances from the maternal plant. Formica cunicularia Latreille, 1798, F. lemani, F. cinerea Mayr, 1853, Myrmica ruginodis Nylander, 1846, Tetramorium caespitum L., 1758 dominate among the recognized species. Due to the activity of ants (seeds dispersal) some loci of P. filarszkyana in the typical phytocoenoses of the Ukrainian Carpathians at the different altitude levels are established.
Obligate and facultative myrmecochors can be distinguished among the myrmecochorous plants in general. Their seeds usually have attractive greasy corpuscules or elaiosomes, which are consumed by ants. The seeds are covered by fuzz and fringe to liken them to insects and their larvae what is very attractive for ants. Myrmecochorous species are characterized by early flowering and fruitification. Their morphology has a row of adaptations for self-seeding as well. The elaiosomes of majority of seeds and fruits gathered in ant-hills after having been consumed by ants are removed from there being able to grow. So, that is the reason of different plant composition on the ant-hills of Formicidae and around them from the rest of the territory without the ant-hills and favorable soil conditions, accordingly.
The disruption of the relationships between plants and insects can, on the one hand, results in the extinction of populations of entomophilous plant species and in the loss of the food base for leaf beetles, for example, on the other hand. The bumblebees are one the most essential for the viability of plant populations of the Ukrainian Carpathian high-mountains among the other pollinators. We discovered that the pollinators' composition does not depend a lot on plant species, but depend on the ecological conditions of their habitat. A stable relationship between flowering plants and insect pollinators (they are mainly Hymenoptera) ensures successful reproduction, sufficient exchange of genetic material between populations of plants to form a mature seed.
Thus, after finding out the ways of pollination and seed dispersal of plants in the Ukrainian Carpathians it is now possible to explain the processes and mechanisms of plant population survival in the high-mountain regions and predict the dynamics of their populations.

CONCLUSIONS
It was found out that such high-mountain plant species of the Ukrainian Carpathians as Knautia dipsitifolium, Arnica montana, Senecio carpaticus, Pulmonaria filarszkyana and Astrantia major are tightly connected with a number of insect pollinators. The most numerous ones among them are representatives of Bombyliidae and Syrphidae, and the less numerous species are insects from the families of Andrenidae and Halictidae. The highest number of pollinator species is recorded for K. dipsitifolium (30 ones), and the lowest one -for P. filarszkyana (9 species).
The most significant influence of phytofagous insects over the vegetative sphere of the plants occurs for S. carpaticus where 40 % of the total amount of consorts belongs to the leaf beetles (Chrysochloa). The important role of such species of ants as Formica cuni cularia, F. lemani, F. cinerea, Myrmica ruginodis, Tetramorium caespitum in the disse mination of P. filarszkyana seeds is established. This fact points at the lungwort as a typical myrmecochorous species in high-mountains of the Ukrainian Carpathians.