The Bumble bees and Cuckoo bees (Bombini, Apoidea) of the Western Bieszczady Mts.

The Bumble bees and Cuckoo bees (Bombini, Apoidea) of the Western Bieszczady Mts. and the Beskid Niski Mts. Andrzej Kosior, Jan Fijał, Wiesław Król, Piotr Płonka, Andrzej Kalemba, Joanna Korzeniak Institute of Nature Conservation, Polish Academy of Sciences Mickiewicza, - Kraków, Poland, e-mail: kosior@iop.krakow.pl Abstract The paper presents the results of a multi-year study on species composition and numbers of bumble bees and cuckoo bees Bombini, conducted in the period - in the Beskid Niski Mts., Western Bieszczady Mts., and a Transition Zone between them. In all, bumble-bee species and cuckoo bees representing. % and. %, respectively, of the total number of these species recorded in Poland. In terms of the Bombini species, the greatest number were found in the the Western Bieszczady Mts. (), with a lower number found in the Beskid Niski (), whereas the Transition Zone had the smallest number (). Six species were deemed new for the Beskid Niski Mts. and for the Western Bieszczady Mts. Four rare species of bumble bees have probably become extinct,including B. muscorum in the Beskid Niski Mts. as well as B. confusus, B. cryptarum, and B. humilis in the Western Bieszczady Mts., while other species are threatened with extinction. Key words Bombini, species composition and numbers, threats, feeding plants, Beskid Niski Mts., Western Bieszczady Mts., Polish Carpathians Introduction The earliest notes on Apoidea bees, including Bombini bumble-bees and cuckoo bees from the Polish Carpathians, date back to the second half of the th century (Siła- Nowicki ; Wierzejski, ; Śnieżek, ). The fairly scarce data pertains to distribution, biology, morphology and the frequency of occurrence of the majority of these species in the Tatra Mts., Pieniny Mts., Dynowskie foothills, and the eastern Beskid Mts. More detailed and complete early data on the biology, ecology and zoogeography of the Bombini species from various regions of the Polish Capathians, date back only to the first half of the th century (Śnieżek ; Noskiewicz ; Dylewska ). The increased interest in the bumble bees and cuckoo bees of the Polish Carpathians first occurred in the late s, and increased in subsequent decades up till the present day. The studies of this group of insects included either the whole area of the Carpathians (Ruszkowski et al. ; Biliński et al. ; Ruszkowski et al. ), or several intensively studied regions (Żak ; Dylewska and Zabłocki ; Banaszak ; Celary ; Kosior, ; Banaszak and Rasmont ; Dylewska et al. ; Celary et al. ; Kosior et al. ), and single physiographical regions, including the Tatra Mts. (Dylewska ; Kosior ; Kosior et al. ), Pieniny (Dylewska ; Kaźmierczak et al.; Kosior et al. ; Kosior et al. b), the Beskid Żywiecki Mts, including the Babia Góra (Dylewska ; Celary ) and Pilsko massifs (Witkowski and Kosior ), Beskid Mały Mts. (Witkowski et al.), Beskid Niski Mts. (Kosior et al. a; Celary et al. (msc)) and the Western Bieszczady Mts. (Banaszak ; Kosior,,,, (), ; Starzyk and Kosior ; Biliński ; Wiśniowski ; Celary et al. ; Kosior et al.;, Kosior and Płonka ). The topics of the above papers pertained chiefly to the biology, ecology and zoogeography of the Bombini, their role as pollinators, threats to the populations exposed to the impact of natural and anthropogenic factors, as well as to the active measures designed to protect these insects. Despite the ever-growing interest and intensive efforts in research on Bombini insects throughout the whole of the Polish Carpathians, none of the authors have ever compared the associations of the Bombini insects between the adjacent areas of the Western and Eastern Carpathians. For this reason, the authors of this paper have undertaken Nature Conservation :,.

U K R A I N E A. Kosior et al. comparative studies on species composition and numbers, as well as threats to the associations of bumble bees and cuckoo bees in the Beskid Niski Mts. and the Western Bieszczady Mts. Study area and methods The studies of associations of Bombini bees were carried out in the Western Bieszczady Mts. (Eastern Carpathians), and the Beskid Niski Mts. (Western Carpathians), within an area restricted by two mountain passes: Dukielska and Łupkowska, referred to in this paper as the Transition Zone (Fig. ). The division into physiographical regions follows Kondracki (), whilst designating the Transition Zone, which does not appear in this source, resulted from geobotanical research indicating that the vegetation of this region includes both East- and West- Carpathian elements (Pałczyński, Michalik, Zemanek ). Details of the physiography, climate and vegetation cover of the study area have not been supplied in this paper, because they were presented in the earlier papers of a number of authors, either for the Beskid Niski Mts. or their part: Sulma ; Tacik et al. ; Hryncewicz ; Święs ; Paczos ; Staszkiewicz ; Medwecka-Kornaś ; Hess et al. ; Michalik ; Brzeźniak et al. ; Szafrański DV DV DV EV EV EV EV EV EV Szymbark Folusz Grybów Kornuty EV EV EV EV EV Biała B Ropa E S K I D Wisłoka Krempna EV EV EV EV EV Wysowa Krynica EV EV EV EV Dukla Jasiołka N I S K EV EV EV EV Wisłok I EV EV EV Sanok EV Osława EV Czaszyn Rzepedź EV EV San FV Solina FV FV FV Ustrzyki Dln. FV FV B I E EV EV Nw. Łupków S Z C Z A D EV Cisna FV FV San FV Dwernik EV FV Smerek FV FV FV S L O V A K I A km Y Z A C H O D N I E Ustrzyki Grn. FV U K R A Tarnica FV I N E Fig.. Selected regions of the Polish Carpathians, where comparative studies of Bombini were conducted; namely: the Beskid Niski Mts. (Western Carpathians), the Western Bieszczady Mts. (Eastern Carpathians) and their Transition Zone. Explanations: boundary of investigated area, boundary of the Beskid Niski Mts., boundary of the Western Bieszczady Mts., transition zone between the two regions. confusus Fig.. Locations of Bombus confusus Schenck,, in the ; Dubiel et al., for the Western Bieszczady Mts. or their parts: Pałczyński ; Zarzycki ; Jasiewicz ; Zarzycki ; Michna and Paczos ; Paczos ; Kosior ; Michalik and Szary ; and for the Transition Zone : Pałczyński ; Grodzińska, Pancer- Kotejowa ; Zarzycki ; Michna and Paczos ; Paczos ; Karczmarz ; Kosior and Michalik. The study is based on data collected irregularly from the s to s and on regular fauna inventories taken in the years from -, when the research work proceeded in month-long intervals in various seasons. The great majoriy of the observations of Bombini insects were made in the summer months (June-August). In the zones Nature Conservation :,.

studied, observations also covered from to as many as full vegetation seasons (April/May September/October). In the years:, -, and all three zones were checked over the same period. In total, the observations were carried out in the Beskid Niski Mts. over days, in the Transition Zone for days and in the Western Bieszczady Mts. for days. The studies covered the following types of habitats: deciduous, coniferous, and mixed forests, forest glades, alder shrub and ossiers, riparian tall-herb communities, swards on rocks, polonina-type mountain meadows, heathlands, bilberry communities, meadows of various kinds, roadsides (along forests, fields, rural, or upgraded roads), paved roads, railway embankments, fallow-lands, ruderal sites, and cultivated fields (clover, alfalfa, lupine, rape, as well as root or cereal crops). The observations of bumble bees and cuckoo bees carried out in summer, and in some years also throughout the whole of the vegetation season, allowed for compilation of fairly comprehensive lists of flowering plants, visited by these insects in the three selected regions of the Polish Carpathians. The data on Bombini species composition and numbers, their food plants, occupied habitats, appearance or disappearance in/from particular locations in selected regions, was entered into a computer database. The collected specimens of bumble bees and cuckoo bees were mounted and species identified. The whole collection was transferred to the collections of the Institute of Systematics and Evolution of Animals, PAS, Krakow. The distribution of particular species of Bombini in specific research localities, in selected regions of the Polish Carpathians were presented in the UTM geographical grid. Similarly, the combined numbers of Bombini species were presented in research locations situated within the same UTM squares. The systematics of Bombini is given according to Reinig () and von Hagen, Aichhorn (). The categories of the extinction risk for the Bombini species in Poland were adopted in line with the IUCN guidelines (IUCN ), and presented after Banaszak (); namely: VU vulnerable, DD data-deficient. The Bombini species covered by legal measures of protection, were listed according to the most recent Ordinance by the Minister of the Environment [Dz. U. (Journal of Laws) b]. The authors adopted a convention of referring to species and subspecies of Bombini combined as simply species. The term association was used to indicate a group of Bombini species associated with a specific type of vegetation habitat. The systematics and names of food plant species follow Mirek et al. (). Results In the three selected regions of the Polish Carpathians, a total number of species of Bombini were recorded (Table ), including species of bumble bees and species of cuckoo bees. By region, the numbers of Bombini species were: in the Beskid Niski Mts. -, in the Transition Zone - only species, whereas in the Western Bieszczady Mts. there were as many as. It was found that as many as species were common to either or studied regions (Figs -, -, -, -, -, -,, and -), and only species were recorded exclusively in the Beskid Niski Mts. (Figs, and ), and species, exclusively in the Western Bieszczady Mts. (Figs,,,,,,, and ; see Table ). Because the bumble bees and cuckoo bees were studied in various time slots covering the whole season where these insects appear, it was possible to compare the changes in the species compositions of Bombini within particular regions (see Table ). A number of species new to particular areas were found: for the Beskid Niski Mts. [B. cryptarum (Fig. ), B. s. proteus and B. s. soroeensis (Figs -), Ps. norvegicus (Fig. ), Ps. rupestris (Fig. ), and Ps. vestalis (Fig. )] and for the Western Bieszczady Mts. [B. distinguendus (Fig. ), B. magnus (Fig. ), B. pyrenaeus (Fig. ), B. semenoviellus (Fig. ) and Ps. flavidus alpium (Fig. )]. Next, the studies conducted at the turn of the st century failed to confirm the occurrence of several, bumble bee species already very rare at that time: B. muscorum (Fig. ) in the Beskid Niski Mts. and B. confusus (Fig. ), B. cryptarum (Fig. ), and B. humilis (Fig. ) in the Western Bieszczady Mts. It is fairly probable that these species became extinct in the regions under study. In the year in the Transition Zone, several species of Bombini were not found, which were however, recorded to various degrees in other habitats, e.g.: B. hortorum (Fig. ), B. hypnorum (Fig. ), B. pratorum (Fig. ), B. sylvarum (Fig. ), Ps. bohemicus (Fig. ), Ps. campestris (Fig. ), and Ps. sylvestris (Fig. ). This result was evidently affected by the fact that the year, was the last year of studies in the Transition Zone, while in the adjacent regions the studies continued for another two years (see Table ). Among the Bombini species presented in Table, occurring only in the Western Bieszczady Mts., the following species, are noteworthy: B. pyrenaeus (alpine species, Fig. ), B. w. mastrucatus (mountain species, Fig. ), B. jonellus (glacial relict, north-mountain species, Fig. ), and Ps. flavidus alpium (also a north-mountain species, Fig. ), followed by B. semenoviellus (eastern species, Fig. ), as well as B. magnus (Fig. ) and Ps. Nature Conservation :,.

A. Kosior et al. Table. Bombini species of the Beskid Niski Mts., Transition Zone, and the Western Bieszczady Mts. Species Western Bieszczady Beskid Niski Mts. Transition Zone Mts. - - - - - Categories of risk of extinction Bombus confusus Schenck, * - - - - +? VU Bombus cryptarum (Fabr., )* - + - - +? DD Bombus distinguendus Mor. * + + - - - + VU Bombus hortorum (Linnaeus, )* + + + - + + Bombus humilis Illiger, * + + - - +? VU Bombus hypnorum (Linnaeus, )* + + + - + + Bombus jonellus (Kirby, )*! - - - - + + VU Bombus lapidarius (Linnaeus, ) + + + + + + Bombus lucorum (Linnaeus, )* + + + + + + Bombus magnus Vogt, * - - - - - + DD Bombus muscorum (Fabricius, )* +? - - - - Bombus pascuorum (Scopoli, )* + + + + + + Bombus pratorum (Linnaeus, )* + + + - + + Bombus pyrenaeus Pérez, * - - - - - + DD Bombus ruderarius (Müller, )* + + + + + + Bombus ruderatus (Fabricius, )* + + - - + + VU Bombus semenoviellus Skorikov, * - - - - - + Bombus s. proteus (Gerstaecker, )* - + - - - - VU Bombus s. soroeensis (Fabicius, )* - + - - + + VU Bombus subterraneus (Linnaeus, )* + + + - + - VU Bombus sylvarum (Linnaeus, )* + + + - + + Bombus terrestris (Linnaeus, ) + + + + + + Bombus veteranus (Fabricius, )* + + + - + + VU Bombus w. mastrucatus Gerstaecker, * - - - - + + DD Psithyrus barbutellus (Kirby, ) + + + - + + Psithyrus bohemicus (Seidl, ) + + + - + + Psithyrus campestris (Panzer, ) + + + - + + Psithyrus flavidus alpium (Richards, ) - - - - - + Psithyrus norvegicus Sparre-Schneider, - + - - + + DD Psithyrus quadricolor Lepeletier, - - - - + + DD Psithyrus rupestris (Fabricius, ) - + - - + + Psithyrus sylvestris Lepeletier, + + + - + + Psithyrus vestalis (Fourcroy, ) - + + - + - Combined number of species Explanations: * - species under strict protection acc. to the Ordinance of the Minister of the Environment [Dz. U (Journal of Laws) ( b)]; Banaszak (), Banaszak, Rasmont ();! in the Carpathians, B. jonellus is a glacial relict (May ); categories of risk of extinction in Poland given after Banaszak (): VU vulnerable in Poland, DD data-deficient in Poland;? probably extinct in a given region. quadricolor (Fig. ), recorded only in the polonina high mountain meadows. The number of Bombini species in all research sites situated within the same UTM squares in three areas of the Carpathians, are shown in Fig.. The numbers of these insect species in different squares of the study area, varied from - to -. The highest numbers were recorded in the Western Bieszczady Mts, namely in the Bieszczadzki National Park (-), with high numbers (-) in its buffer zone, and Nature Conservation :,.

cryptarum Fig.. Locations of Bombus cryptarum (Fabricius, ) in the distinguendus Fig.. Locations of Bombus distinguendus Morawitz,, in the hortorum Fig.. Locations of Bombus hortorum (Linnaeus, ) in the hortorum Fig.. Locations of Bombus humilis Illiger,, in the selected study areas. hypnorum Fig.. Locations of Bombus hypnorum (Linnaeus, ) in the in farmlands; while in the north western parts of the area the numbers were lower (- species), whereas the lowest ranges of these numbers were recorded in the eastern zones (- and -). In the Beskid Niski Mts., high numbers of jonellus Fig.. Locations of Bombus jonellus (Kirby, ) in the the Bombini species were found in the Magurski National Park, its buffer zone, and in the vicinity of the Szymbark locality (- in each of the areas). The mid-western areas of the region were evidently less abundant in bumble-bee Nature Conservation :,.

A. Kosior et al. lapidarius Fig.. Locations of Bombus lapidarius (Linnaeus, ) in the lucorum Fig.. Locations of Bombus lucorum (Linnaeus, ) in the Fig.. Locations of Bombus magnus Vogt,, in the selected study areas. Fig.. Locations of Bombus muscorum (Fabricius, ) in the pascuorum Fig.. Locations of Bombus pascuorum (Scopoli, ) in the and cuckoo bee species (-), while the lowest numbers of Bombini species were recorded in the area of the Transition Zone. In this zone, only farmlands and forest glades were pratorum Fig.. Locations of Bombus pratorum (Linnaeus, ) in the somewhat richer in terms of the numbers of species (-). The remaining areas of the Transition Zone contained either low or the lowest numbers of Bombini species (- and - Nature Conservation :,.

Fig.. Locations of Bombus pyrenaeus Pérez,, in the ruderarius Fig.. Locations of Bombus ruderarius (Müller, ) in the ruderatus Fig.. Locations of Bombus ruderatus (Fabricius, ) in the semenoviellus Fig.. Locations of Bombus semenoviellus Skorikov,, in the semenoviellus Fig.. Locations of Bombus soroeensis proteus (Gerstaecker, ) in the Fig.. Locations of Bombus soroeensis soroeensis (Fabricius, ) in the ), which confirms how poor this region of the Carpathians is in term of species abundance, compared with the two adjacent regions (see Fig. ). According to the last Ordinance by the Minister of the Environment [Dz. U. (Journal of Laws) b], among all species of bumble bees occurring in Poland, only two are Nature Conservation :,.

A. Kosior et al. subterraneus Fig.. Locations of Bombus subterraneus (Linnaeus, ) in the sylarum Fig.. Locations of Bombus sylvarum (Linnaeus, ) in the terestris Fig.. Locations of Bombus terrestris (Linnaeus, ) in the veteranus Fig.. Locations of Bombus veteranus (Fabricius, ) in the wurflenii Fig.. Locations of Bombus wurflenii mastrucatus Gerstaecker,, in the not strictly protected: B. lapidarius (Fig. ) and B. terrestris (Fig. ). Thirteen species of bumble bees and two species of cuckoo bees were placed on the Red List of species threatened with extinction (Banaszak ) and assigned Psithyrus barbutellus Fig.. Locations of Psithyrus barbutellus (Kirby, ) in the to groups with the relevant level of threat (Table ). Noteworthy among the vulnerable species are particularly valuable and rare species: B. jonellus a glacial relict (Fig. ), and two subspecies of Bombus soroeensis: B. s. soroeensis Nature Conservation :,.

Psithyrus bohemicus Fig.. Locations of Psithyrus bohemicus (Seidl, ) in the Psithyrus campestris Fig.. Locations of Psithyrus campestris (Panzer, ) in the Psithyrus flavidus alpium Fig.. Locations of Psithyrus flavidus alpium (Richards, ) in the Psithyrus norvegicus Fig.. Locations of Psithyrus norvegicus Sparre-Schneider,, in the Psithyrus quadricolor Fig.. Locations of Psithyrus quadricolor Lepeletier,, in the Psithyrus rupestris Fig.. Locations of Psithyrus rupestris (Fabricius, ) in the and B. s. proteus (Figs -), whose distribution areas cross each other in Poland (Dylewska ). The existence of colonies of Bombini showed relationships both with species composition and abundance of nectar plants. In the Beskid Niski Mts., the insects visited plant species from families, in the Transition Zone plant species, also from families, whereas in the Western Bieszczady Mts. Nature Conservation :,.

A. Kosior et al. Psithyrus sylvestris Fig.. Locations of Psithyrus sylvestris Lepeletier,, in the Psithyrus vestalis Fig.. Locations of Psithyrus vestalis (Fourcroy, ) in the BN Total number of species: SP BZ Fig.. Combined number of the Bombini species in the research locations in the the number was plant species from families. In total, species of vascular plants visited by Bombini were recorded. Particular favourites visited in all three regions were the flowering plants of the following families: aster family, Asteraceae (- visiting Bombini species), mint family Lamiaceae (- Bombini species), and papilionaceous plants Fabaceae (- Bombini species). Particular species of bumble bees visited from to species of flowering plants, whereas in the case of the cuckoo bee species they visited from to species of plants. The plant species most visited by Bombini insects include: Centaurea phrygia ( species of Bombini), Centaurea jacea ( species), Carduus personata ( species), Cirsium palustre ( species), Trifolium pratense, Knautia dipsacifolia and Carduus acanthoides (each visited by species), as well as Knautia arvensis and Cirsium vulgare (each visited by species). > The knowledge of the flowering plants visited by Bombini in the three studied regions of the Carpathians, allowed us to differentiate these areas from the viewpoint of selected categories of plant species, and the bumble bee and cuckoo bee species pollinating them. The categories of species both plants and the Bombini, take into account various aspects, linked to the specific nature of some species, i.e. with their geographical range, vertical distribution, and their protection status (Table ). Among the three regions of the Carpathians studied, the most abundant in almost all categories of feeding plants and their pollinators were the Western Bieszczady Mts. As regards the plant species, one all-carpthian endemite and subendemites were recorded in the Western Bieszczady Mts, along with one species endemic to the Carpathians and Sudety Mts. Among the East-Carpathian species, the following occurred: Centaurea kotschyana and Cirsium waldsteinii, two species associated with poloninas and subalpine tall-herb communities; Dianthus compactus and Scorzonera rosea, two other polonina-related species, although often entering the meadows at lower elevations; and Lapsana intermedia, a rather infrequent plant from shrub and ruderal communities. In one of the two remaining regions; namely the Transition Zone, was one all-carpathian endemite found. Among East-Carpathian plants the most visited was Cirsium waldsteinii, a species fairly common in the Bieszczady Mts. which attracted as many as Bombini species, including rare species: B. jonellus, B. pyrenaeus, and B. w. mastrucatus. Centaurea kotschyana, visited by species, including B. jonellus and B. w. mastrucatus was slightly less significant as a food plant. None of these plants was, however, provided a principal food base for B. jonellus, visiting chiefly Solidago virgaurea, Chamaenerion angustifolium, Knautia dipsacifolia, and Thymus pulegioides, nor for Nature Conservation :,.

Table. Comparison between three regions in the study concerning the distribution and protection status of the Bombini species and their food plants. Beskid Niski Mts. Transition Zone Western Bieszczady Mts. Categories of species Bombini Plants Bombini Plants Bombini Plants Geographical distribution range Carpathian endemite Carpathian subendemite Carpathian- Sudety East-Carpathian Eastern non-mountain Western Altitutdinal range Alpine Mountain or montane North-mountain Protection status Strictly protected Partly protected Explanations: plants endemites, subendemites, species endemic to the Carpathians and the Sudety Mts., East-Carpathian species, and mountain species are given after Zemanek, Winnicki (), and Piękoś Mirkowa, Mirek (); species under strict or partial protection acc. to the Ordinance of the Minister of the Environment ( a); Bombini insects western species (Dylewska ); eastern species (Reinig ); alpine and mountain species (Banaszak ); northern mountain (Celary et al. ); Bombini species under strict protection in Poland are given acc. to the Ordinance of the Minister of the Environment [Dz. U. (Journal of Laws) b]. B. w. mastrucatus, which visited more than different species of flowering plants. With respect to the visited xerothermophilous plant species, the three study areas are very similar. Because of the lack of habitats typical of xerothhermic swards, the thermophilous species occur chiefly in secondary habitats, such as stony roadsides, sides of railway embankments or dry slopes along roads. Usually the localities of these plants are situated within river valleys. Some of the species recorded in the Beskid Niski Mts. (Ballota nigra, Genista germanica, Jovibarba sobolifera, Stachys germanica) probably no longer occur in the Western Bieszczady Mts. In the natural, semi-natural, and anthropogenic communities of the three studied regions of the Polish Carpathians, the Bombini visited and pollinated not only the abovementioned groups of plants but also a number of medicinal plant species: herbs, shrubs, or trees (,, and species respectively), cultivated species (,, and respectively) as well as weeds (,, and species respectively). Discussion Among the Bombini associations from the three studied regions, the following species were found only in the Western Bieszczady Mts. (see Table ): B. pyrenaeus an alpine species (Fig. ), B. w. mastrucatus a mountain species (Fig. ), B. jonellus (Fig. ) and Ps. flavidus alpium (Fig. ) two northern-mountain species, as well as B. magnus (Fig. ) and Ps. quadricolor (Fig. ) found on poloninas, and finally B. semenoviellus (Fig. ) an eastern species. The differences occurring in the western part of the Beskid Niski Mts. are further signified by the fact that only two species of bumble bees were recorded there; namely: B. muscorum (Fig. ) and B. s. proteus which is a western element (Fig. ). As expected, the area of the Transition Zone, contained species of bumble bees and cuckoo bees in common with either of the neighbouring regions. These similarities and differences in the distribution of some groups of the entomofauna in this part of the Carpathians were also confirmed by Kosior (), Michalik (), Kosior et al. (a), and Celary et al. (,, msc). Nature Conservation :,.

A. Kosior et al. The nature of differences in the composition of Bombini insects occurring in the three regions under study (the occurrence of mountain species, eastern and western species) within the area of the Western Bieszczady Mts., and two areas of the Beskid Niski Mts., illustrates very well the phytogeographical differences of the Bieszczady Mts. and the transitional nature of the remaining two regions, shown in classical floristic accounts of the area (Jasiewicz, Zemanek ). The East-Carpathian species visited by Bombini insects were recorded exclusively within the Western Bieszczady Mts., with the only exception being Glechoma hirsuta, a non-mountain species classified as an eastern element, which was observed in all three studied regions. In spite of marked differences in the frequency of field observations between the Beskid Niski Mts., the Transition Zone, and the Western Bieszczady Mts., all have been almost completely penetrated over the course of many years of studies, as confirmed by the successful findings of even very rare species of bumble bees and cuckoo bees. Then, the series of field studies in two periods, i.e. in the second half of the th century and at the turn of this century, has enabled both the recognition of new and possibly extinct species of Bombini in this portion of the Polish Carpathians. The species composition of the Bombini insects in the studied regions, does not differ significantly vis-à-vis the fauna of this group described throughout the whole of the Polish Carpathians [Wierzejski, ; Śnieżek,, ; Dylewska,,, ; Żak ; Banaszak, ; Kosior, (msc), Kosior et al. ; Celary, ; Ruszkowski et al., ; Rasmont ; Witkowski and Kosior ; Dylewska et al. ; Witkowski et al. ; Kosior et al.,, a; Wiśniowski ; and Celary et al.,, msc.]. The differences pertain only to several species, namely: B. mesomelas and B. pomorum, which have never been recorded in the three studied regions of the Carpathians, and B. semenoviellus and Ps. flavidus alpium, which are species new to the Polish Carpathians. Additionally, B. semenoviellus is also a species new to southern Poland, while Ps. flavidus alpium has established only its second location in Poland here. Also noteworthy is B. pyrenaeus, an alpine species which turned out to be new to the Polish Eastern Carpathians. Acknowledgments The authors wish to express their cordial thanks to Professor H. Piękoś-Mirkowa, Ph.D. for her valuable comments on the substantive content, regarding the diffentiation of the food plant species of Bombini insects, from the viewpoint of their geographical distribution and altitudinal zones, and to Professor T. Pawlikowski, Ph.D. Assoc. Professor M. Biliński, Ph.D, J. Galas Ph.D. (Ms.), J. Galas, Ph.D., J. Partyka, Ph.D., and Mr. J. Skibiński for their dedicated help in field research over several years. REFERENCES BANASZAK J.. Przyczynek do znajomości fauny trzmieli (Bombus Latr.) Bieszczadów Przegl. Zool., :. BANASZAK J.. Materiały do znajomości fauny trzmieli (Bombus Latr.) i trzmielców (Psithyrus Lep.) Bieszczadów. Przegl. Zool., : - (in Polish with an English summary). BANASZAK J.. Materiały do znajomości pszczół (Hymenoptera, Apoidea) fauny Polski. Bad. Fizjogr. n. Polską Zach., C-Zoologia: -. BANASZAK J.. Czerwona Lista Zwierząt Pszczoły Apoidea. In: Z. Głowaciński (Ed.). Czerwona Lista Zwierząt Ginących i Zagrożonych w Polsce, IOP PAN, Kraków: -. BANASZAK J., RASMONT P.. Occurrence and distribution of the subgenus Bombus Latreille sensu stricto in Poland (Hymenoptera, Apoidea). Pol. Pis. Entom. : - (in English). BILIŃSKI M., SOWA S., KOSIOR A.. Density of pollinating insects on red clover in Poland. In.: J. Banaszak (Ed.). Natural resources of wild bees in Poland, Pedag. Univ., Bydgoszcz: -. BILIŃSKI M.. Gniazda trzmieli w naturze (Bumblebee nests occurring in nature). In: T. Cierzniak (Ed.). Postępy apidologii w Polsce.Wyższa Szkoła Pedagog., Bydgoszcz: - (in Polish with an English summary). BRZEŹNIAK E., CZEMERDA A., PACZOS S.. Opady atmosferyczne na obszarze Beskidu Niskiego. Probl. Zagosp. Ziem Górsk. : - (in Polish with an English summary). CELARY W.. Review of the cuckoo bees of the genus Psithyrus Lep. (Hymenoptera, Apidae) in Poland. Rocz. Muz. Górnośl. w Bytomiu : -. (in English). CELARY W.. Hymenoptera Aculeata (excluding Formicoidea) of Babia Góra Mt. and adjacent area. Acta Zool. Cracov.,,: -. (in English). CELARY W., FIJAŁ J., KOSIOR A.. Bombus pyrenaeus Perez (Hymenoptera, Apoidea, Apidae) in Poland. Acta Zool. Cracov., : - (in English). CELARY W., FIJAŁ J., KOSIOR A.. Psithyrus flavidus (Eversmann) (Hymenoptera: Apoidea: Apidae) in Poland. Acta Zool. Cracov.,: - (in English). CELARY W., FIJAŁ J., KOSIOR A. (msc). Bombini (Hymenoptera, Apoidea, Apidae) of the Beskid Niski Mts. Instytut Systematyki i Ewolucji Zwierząt PAN, Sławkowska Kraków. Nature Conservation :,.

DUBIEL E., STACHURSKA A., GAWROŃSKI S.. Nieleśne zbiorowiska roślinne Magurskiego Parku Narodowego (Beskid Niski). Prace Bot. UJ : -. DYLEWSKA M.. Zarys rozsiedlenia gatunków z rodzaju Bombus Latr. na obszarze Polski (The distribution of the species of genus Bombus Latr. in Poland). Acta Zool. Cracov., : - (in English). DYLEWSKA M.. Fauna trzmieli (Bombus Latr.) i trzmielców (Psithyrus Lep.) Tatr Polskich (The Bombus Latr. and Psithyrus Lep. fauna of the Polish part of the Tatra Mountains). Acta Zool. Cracov., : - (in English). DYLEWSKA M.. The Apoidea of the Pieniny National Park. Part I. Megachilidae and Apidae (partim) [Apoidea Pienińskiego Parku Narodowego. Część I. Megachilidae and Apidae (partim)]. Acta Zool. Cracov., : - (in English). DYLEWSKA M.. The Apoidea of the Babia Góra Mountain (Apoidea Babiej Góry). Acta Zool. Cracov., : - (in English). DYLEWSKA M.. Trzmiele i trzmielce parków narodowych tatrzańskiego i Babiogórskiego oraz czynna ochrona tych owadów. Monitoring pszczół. TPN, Zakopane (in Polish). DYLEWSKA M., GĄSIENICA-CHMIEL M., KOSIOR A., SUMERA B., SZAFRANIEC S., WERSTAK K., WIŚNIOWSKI B.. Skład gatunkowy i liczebność trzmieli i trzmielców (Bombinae, Apoidea, Hymenoptera) na łąkach w wybranych parkach narodowych oraz kwiecistość łąk w tych parkach w roku [Species composition and numbers of the Bombus Latr. and Psithyrus Lep. on selected meadows in particular selected Parks in Poland and the floral abundance of these meadows in ]. Prądnik. Prace Muz. Szafera, -: - (in Polish with an English summary). DYLEWSKA M., ZABŁOCKI J.. Nowe i mało znane Apoidea (Hymenoptera) z obszaru Polski [New and little known Apoidea (Hymenoptera) from Poland]. Acta Zool. Cracov.,, : - (in Polish with an English summary). GRODZIŃSKA K., PANCER-KOTEJOWA E.. Zbiorowiska leśne pasma Bukowicy w Beskidzie Niskim. Fragm. Flor. Geobot., : - (in Polish with an English summary). HAGEN VON E., AICHHORN A. A. Hummeln: bestimmen, ansiedeln, vermehren, schützen; -. Fauna Verlag, Nottuln, Deutschland (in Germany). HESS M., NIEDŹWIEDŹ T., OBRĘBSKA-STARKLOWA B.. Stosunki termiczne Beskidu Niskiego. Zesz. Nauk. UJ, Prace Geogr. : - (in Polish with an English summary). HRYNCEWICZ Z.. Łąki i pastwiska Beskidu Niskiego pod względem geobotanicznym i gospodarczym. Zesz. Problem. Postępów Nauk Roln. : -. IUCN. Categories and Criteria (version.). Red list of threatened species. IUCN, Gland, Switzerland. JASIEWICZ A.. Rośliny naczyniowe Bieszczadów Zachodnich (The vascular plants of the Western Bieszczady Mts. East Carpathians). Monogr. Bot. : -. KARCZMARZ K.. Mszaki Pasma Bukowicy w Beskidzie Niskim - Bryophyta of the Bukowica Range in the Low Beskid (Polish Western Carpathians). Fragm. Flor. Geobot., : - (in Polish with an English summary). KAŹMIERCZAK R., KAŹMIERCZAK T., KOSIOR A.. Kwiecistość łąk Pienińskiego Parku Narodowego i jej związek z fauną trzmielowatych (Bombini) i gąsienicznikowatych (Ichneumonidae). Ochr. Przyr. : -. KONDRACKI J.. Geografia regionalna Polski. PWN, Warszawa: - (in Polish). KOSIOR A.. Trzmiele (Bombus Latr.) i trzmielce (Psythirus Lep.) Bieszczadzkiego Parku Narodowego oraz terenów użytkowanych gospodarczo (Bereska, Myczków, Terka) w Bieszczadach. Przegl. Zool., : - (in Polish with an English summary). KOSIOR A.. Znaczenie, zagrożenie i ochrona trzmieli (Bombus Latr.) na obszarze zlewni Bereźnicy w Bieszczadach Zachodnich. Stud. Nat. A : - (in Polish with an English an summary). KOSIOR A.. Rola trzmieli (Bombus Latr.) w biocenozach Bieszczadów Zachodnich (The role of bumblebees (Bombus Latr.) in the biocenoses of the West Bieszczady Mountain range). Ochr. Przyr. : - (in Polish with an English summary). KOSIOR A.. Wpływ działalności gospodarczej na populacje trzmieli Bombus Latr. w Bieszczadach Zachodnich (Impact of economic activity upon bumblebee Bombus Latr. population in the West Bieszczady Mts.). Ochr. Przyr. : - (in Polish with an English summary). KOSIOR A.. Trzmiele Bombus Latr. wybranych polan reglowych Tatrzańskiego Parku Narodowego (The bumblebees Bombus Latr. of the selected glades of the Tatra National Park). Stud. Naturae, ser. A : -. KOSIOR A.. The influence of anthropogenic factors on the decline of bumblebees (Bombus Latr.) in Poland. In: J. Banaszak (Ed.). Natural resources of wild bees in Poland. Pedagog. Univ., Bydgoszcz:- (in English). KOSIOR A.. Changes in the fauna of bumble-bees (Bombus Latr.) and cuckoo - bees (Psithyrus Lep.) of selected regions in Southern Poland. In: J. Banaszak (Ed.).; Changes in Fauna of Wild Bees in Europe, Pedagog. Univ., Bydgoszcz: - (in English). KOSIOR A. (msc). Trzmiele (Bombus Latr.) i trzmielce (Psythirus Lep.) Jaworzyny Krynickiej, Instytut Ochrony Przyrody PAN, Mickiewicza, Kraków (in Polish). KOSIOR A., PŁONKA P., WITKOWSKI Z.. Zgrupowania trzmielowatych (Bombini, Apidae) w wybranych zbiorowiskach roślinnych Pienin. Ochr. Przyr. : -. Nature Conservation :,.

A. Kosior et al. KOSIOR A. (). Trzmielowate (Bombini, Apidae) i motyle dzienne (Rhopalocera) wybranych rejonów Bieszczadów. Rocz. Bieszczadzkie : - (in Polish with an English summary) KOSIOR A., KRÓL W., PŁONKA P. a. Trzmielowate (Bombini, Apoidea) Magurskiego Parku Narodowego i jego otuliny. Parki Nar. i Rez. Przyr., : - (in Polish with an English summary). KOSIOR A., KRÓL W., PŁONKA P. b. Changes in bumblebees and cuckoo-bees (Bombini, Apoidea) in the Pieniny National Park and its buffer zone (Southern Poland). Nature Conservation : -. KOSIOR A.. Trzmielowate (Bombini, Apidae), motyle dzienne (Rhopalocera) i kraśniki (Zygaenidae) Bieszczadzkiego Parku Narodowego. Rocz. Bieszczadzkie : - (in Polish with an English summary). KOSIOR A., FIJAŁ J., HOLLY M., SOLARZ W.. Long-term population decline of the Heath Bumblebee Bombus jonellus (K.) (Hymenoptera, Apidae) in Bieszczady National Park. Rocz. Bieszczadzkie : - (in English). KOSIOR A., PŁONKA P.. Trzmiele i trzmielce (Bombini, Apidae) oraz motyle dzienne (Rhopalocera) na wybranych stanowiskach Bieszczadzkiego Parku Narodowego. Rocz. Bieszczadzkie : -. KOSIOR A., KUCIEL H., WALUSIAK E.. Tojad mocny (Aconitum firmum) (Rchb.) Neilr. i oblatujące go trzmielowate (Bombini, Apidae) w Tatrzańskim Parku Narodowym. Chrońmy Przyr. Ojcz., : - (in Polish with an English Summary). KOSIOR A., POŁCZYŃSKA-KONIOR G., FIJAŁ J.. Wpływ budowy i spiętrzenia zbiorników wodnych Czorsztyn-Niedzica i Sromowce Wyżne na owady trzmielowate (Bombini). W: XII Sympozjum Sekcji Hymenopterologicznej Polskiego Towarzystwa Entomologicznego. Streszczenia referatów; -. Ojców, - maja r. MAY J.. Čmeláci v ČSR, jejich bionomie, chov a hospodářský význam. Čs. Acad. Zeměděl. Věd., Praha, Čechoslovakia (in Czech). MEDWECKA-KORNAŚ A.. Szata roślinna dorzecza Białej Dunajcowej. Studia Ośr. Dok. Fizjogr. : -. MICHALIK S.. Beskid Niski. Przyr. Pol. : - (in Polish). MICHALIK S., SZARY A.. Zbiorowiska leśne Bieszczadzkiego Parku Narodowego. Monogr. Bieszcz. : -. MICHNA E., PACZOS S.. Zarys klimatu Bieszczadów Zachodnich. Ossolineum, Prace Wydz. Mat.-Fiz.-Chem. LTN: -. MIREK Z., PIĘKOŚ-MIRKOWA H., ZAJĄC A., ZAJĄC M.. Vascular plants of Poland. A checklist. Krytyczna lista roślin naczyniowych Polski. Pol. Bot. Stud. : - (In English and Polish). NOSKIEWICZ J.. Przyczynek do znajomości fauny żądłówek Tatr Polskich. Kosmos, -: -. Nakł. Pol. Tow. Przyrodn. Lwów. PACZOS S.. Z badań nad opadami atmosferycznymi w województwie rzeszowskim. Fol. Soc. Sc. Lubl., Sect. Geogr.,: -. PAŁCZYŃSKI A.. Łąki i pastwiska w Bieszczadach Zachodnich. Rocz. Nauk Roln. D : -. PIĘKOŚ-MIRKOWA H., MIREK Z.. Endemic taxa of vascular plants in the Polish Carpathians. Acta Soc. Bot. Pol.,: -. REINIG W. F.. Synopsis der in Europa nachgewiesenen Hummel - und Schmarotzerhummelarten. Spixiana:,, -. München, Deutschland (in Germany). Rozporządzenie Ministra Środowiska z dn. lipca r. w sprawie gatunków dziko występujących roślin objętych ochroną: ochrona ścisła - Załącznik Nr oraz ochrona częściowa - Załącznik Nr. Dz. U. a, Nr, poz.. Rozporządzenie Ministra Środowiska z dn. września r. w sprawie gatunków dziko występujących zwierząt objętych ochroną, z wyszczególnieniem gatunków wymagających ochrony czynnej: ochrona ścisła - Załącznik Nr oraz ochrona częściowa - Załącznik Nr. Dz. U. b, Nr, poz.. RUSZKOWSKI A., BILIŃSKI M., SOWA S., KOSIOR A.. Localization of bumblebee species in Poland. In.: J. Banaszak (Ed.). Natural resources of wild bees in Poland; -, Pedag. Univ., Bydgoszcz. RUSZKOWSKI A., KOSIOR A., BILIŃSKI M., KACZMARSKA K.. Trzmiele Południowej Polski. Pszczeln. Zesz. Nauk. : -. SIŁA-NOWICKI M.. Przyczynek do owadniczej fauny Galicyi. Druk. Uniw., Kraków. STARZYK J. R., KOSIOR A.. Wpływ turystyki pieszej na entomofaunę Bieszczadzkiego Parku Narodowego i terenów przyległych. Parki Nar. i Rez. Przyr., : -. STASZKIEWICZ J.. Zbiorowiska leśne okolic Szymbarku (Beskid Niski). Dok. Geogr. : -. SULMA T.. Kornuty rezerwat na Łemkowszczyźnie. Ochr. Przyr. : -. SZAFRAŃSKI J.. Magurski Park Narodowy. Przyr. Pol. : -. ŚNIEŻEK J.. O krajowych gatunkach trzmieli (About the native bumblebee species). Spraw. Kom. Fizj. : - (in Polish). ŚNIEŻEK J.. O krajowych gatunkach trzmielców (Psithyrus). Spraw. Kom. Fizj. : -. ŚNIEŻEK J.. Błonkówki pszczołowate (Apidae) zebrane w Galicji. Spraw. Kom. Fizj. : -. ŚWIĘS F.. Geobotaniczna charakterystyka lasów w dorzeczu górnego biegu Białej Dunajcowej w Beskidzie Niskim. Cz. I. Lasy olchowe, jesionowo-jaworowe i grabowe. Ann. UMCS, C : -. Nature Conservation :,.

TACIK K., ZAJĄCÓWNA M., ZARZYCKI K.. Z zagadnień geobotanicznych Beskidu Niskiego. Acta Soc. Bot. Pol., : - (in Polish with an English summary). WIERZEJSKI A.. Przyczynek do fauny owadów błonkoskrzydłych (Hymenoptera) (Contribution to the fauna of Hymenoptera). Spraw. Kom. Fizjogr. : - (in Polish). WIERZEJSKI A.. Dodatek do fauny błonkówek (Hymenoptera) (Addition to the Hymenoptera fauna). Spraw. Kom. Fizjogr. : - (in Polish). WIŚNIOWSKI B.. Błonkówki (Hymenoptera) polskich Bieszczadów ze szczególnym uwzględnieniem Bieszczadzkiego Parku Narodowego Monogr. Bieszcz. : - (in Polish with an English summary). WITKOWSKI Z., ADAMSKI P., JENNER B., KOSIŃSKA R., KOSIOR A., KRÓL W., MICHALIK S., NOWAK J., PŁONKA P., SOLARZ W., ZAJĄC T.. Bioróżnorodność ekosystemów lądowych na obszarze przyszłego zbiornika wodnego Świnna Poręba: stan obecny i perspektywy zmian. In.: A. Paulo (Ed.). IX Konf. Sozolog. PTG - Sozologia na obszarze antropopresji na przykładzie zbiornika Świnna Poręba. AGH Kraków: -. WITKOWSKI Z., KOSIOR A.. Wpływ narciarstwa na wybrane grupy chrząszczy (Coleoptera), pszczołowatych (Apoidea) i motyli (Lepidoptera) partii szczytowych Pilska. In: A. Łajczak, S. Michalik, Z. Witkowski (Eds). Wpływ narciarstwa i turystyki pieszej na przyrodę masywu Pilska. Studia Naturae : -. ZARZYCKI K.. Lasy Bieszczadów Zachodnich (The forests of the Western Bieszczady Mts. Polish Eastern Carpathians). Acta Agr. et Silv., S. leśna : -. ZARZYCKI K.. Ogólna charakterystyka Bieszczadów Zachodnich i ich roślinności (Allgemeine Charakteristik des Bieszczady-Gebirges und dessen Pflanzenkleides). Fragm. Faun., : -. ZEMANEK B.. The phytogeographical character of the northwestern part of the Eastern Carpathians (SE Poland). Veroff. Geobot. Inst. ETH, Stiftung Rubel, Zurich : -. ZEMANEK B., WINNICKI T.. Rośliny naczyniowe Bieszczadzkiego Parku Narodowego [Vascular plants of the Bieszczady National Park]. Monogr. Bieszcz. : - (in Polish with an English summary). ŻAK B.. Trzmiele (Bombus Latr.) i trzmielce (Psithyrus Lep.) Pogórza Wielickiego oraz przyległych partii Beskidu Średniego [Bumblebees (Bombus Latr.) and cuckoos (Psithyrus Lep.) of the Wieliczka Foothills and of the adjacent parts of the Middle Beskid]. Zesz. Nauk. UJ, : - (in Polish with an English summary). Nature Conservation :,.