ISSN 0013-8738, Entomological Review, 2014, Vol. 94, No. 1, pp. 79–85. © Pleiades Publishing, Inc., 2014. Original Russian Text © G.S. Potapov, Yu.S. Kolosova, M.Yu. Gofarov, 2013, published in Zoologicheskii Zhurnal, 2013, Vol. 92, No. 10, pp. 1246–1252.
Zonal Distribution of Bumblebee Species (Hymenoptera, Apidae) in the North of European Russia G. S. Potapov, Yu. S. Kolosova, and M. Yu. Gofarov Institute of Ecological Problems of the North, Ural Branch, Russian Academy of Sciences, Arkhangelsk, 163000 Russia e-mail:
[email protected] Received June 13, 2013
Abstract—The bumblebee fauna of the North of European Russia includes 34 species. The trends in bumblebee diversity within the region are characterized. The species richness is the greatest in the middle taiga subzone and decreases from south to north, reaching its minimum in the Arctic tundra. Based on analysis of the zonal differentiation of bumblebee species, only four of them were found to be true arctic forms: Bombus (Alpinobombus) polaris, B. (Al.) balteatus, B. (Al.) hyperboreus, and B. (Pr.) lapponicus glacialis, whereas the rest belong to the temperate, boreal, and subboreal groups. DOI: 10.1134/S0013873814010096
searchers of the Institute of Ecological Problems of the North (Ural Branch of the Russian Academy of Sciences, Arkhangelsk) and the Northern (Arctic) Federal University (Arkhangelsk) in 1993–2012. The data on Novaya Zemlya (1909–1926) were based on collections of the Zoological Institute of the Russian Academy of Sciences (St. -Petersburg). The localities embraced by the study belong to the administrative districts of Arkhangelsk Province and Nenets Autonomous Area (Fig. 1).
Bumblebees (Hymenoptera, Apidae, Bombus Latr.) are one of the most numerous and widespread insect taxa in the north (Bolotov and Podbolotskaya, 2003). They constitute about 85–95% of the total number of individuals of Apoidea in the Eurasian tundras, and 55–70% in the taiga zone of Europe and West Siberia, (Panfilov, 1968), and play a significant role in the ecosystems as the main pollinators of entomophilous plants (Chernov, 1966, 1977). The geographic distribution of bumblebees reflects not only the modern ecological conditions of landscapes but also their changes in the past (Panfilov, 1957). Therefore, the study of the regional faunas is of special interest in the historical aspect, making it possible to specify the ways of formation of regional biocenoses (Bolotov and Podbolotskaya, 2003).
The collected bumblebees were identified using a number of published keys (Kruseman, 1945; Løken, 1973, 1984; Panfilov, 1978). Identifications were verified by studying reference collections at the Zoological Institute of the Russian Academy of Sciences, St.-Petersburg and the Natural History Museum, London. The subgenus- and species-level taxonomy follows the catalogues of the world fauna (Williams, 2012). Bombus (Bombus) lucorum is considered by us as a species complex since reliable identification of B. (Bo.) lucorum, B. (Bo.) cryptarum, and B. (Bo.) magnus is difficult without the use of molecular genetic data (Carolan et al., 2012; Williams, 2012). Bombus (Pyrobombus) lapponicus glacialis Sparre-Schneider, 1902, indicated by us for Novaya Zemlya, is considered by some experts as a distinct species or a subspecies of B. (Pr.) lapponicus isolated on the Arctic islands (Panfilov, 1978; Berezin, 1992, 1995; Kupyanskaya, 1995; Rasmont and Iserbyt, 2010). Data on the mean July air temperatures are borrowed from the Atlas of Arkhangelsk Province (1976). The data were statistically processed using MS Excel software.
Data on the fauna and ecology of bumblebees in various localities of the north of European Russia were reported in a number of publications (Bolotov and Podbolotskaya, 2003; Bolotov and Kolosova, 2006, 2007; The Environment…, 2007; Kolosova and Potapov, 2011a, 2011b). However, the problems of the zonal distribution of species in the region remained almost untouched. This communication presents the results of analysis of the zonal distribution of bumblebees in the north of European Russia, taking into consideration all the data accumulated by now. MATERIALS AND METHODS This work was based on the collections made by the authors in 2006–2011, and those made by the re79
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Fig. 1. Collection sites: 1, Ilinsko-Podomskoye; 2, Verkhnyaya Toima; 3, Shenkursk; 4, Konosha; 5, Kenozero National Park; 6, Mirnyi; 7, Bolshoi Bor; 8, Solovetsky Islands; 9, lower course of the Severnaya Dvina (Arkhangelsk and Kholmogory); 10, Pinega State Reserve; 11, Koida; 12, Nes; 13, Shoina; 14, Kolguev Island (Bugrino); 15, Naryan-Mar; 16, Pymvashor Tract; 17, Amderma; 18, Vaigach Island (Yangoto lake); 19, Novaya Zemlya Archipelago (Matochkin Strait coast, Malye Karmakuly).
RESULTS AND DISCUSSION The bumblebee fauna of the north of European Russia comprises 34 species (table). The species composition of bumblebees in the study region changes
considerably from the middle taiga to the Arctic tundra. We observed a distinct latitudinal trend of species richness of bumblebees corresponding to the gradient ENTOMOLOGICAL REVIEW Vol. 94 No. 1 2014
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of mean July air temperatures, which indicates that only a small number of species could successfully adapt to the Arctic and Subarctic environments (Fig. 2), in spite of numerous adaptations to extreme climatic conditions present in bumblebees. Such adaptations include facultative endothermy and the specific traits of the life cycle, namely annual colonies and overwintering of single females (Pekkarinen, 1988). The development of these adaptations may be related to the initially montane habitats of bumblebees (Hines, 2008). Similar results were earlier obtained for many other taxa, for instance, butterflies and ground beetles (Chernov and Penev, 1993; Chernov and Tatarinov, 2006). The distribution of organisms at high latitudes is most strongly correlated with the mean air temperatures of July (Chernov and Matveyeva, 2002). Under the conditions of total deficit of heat, the ecophysiological value of any rise of temperature is increased (Chernov and Matveyeva, 2002). Therefore, the landscape-zonal structure of the Arctic and Subarctic faunas is characterized by abrupt changes of many parameters along short segments of the latitudinal gradient. As the heat deficit increases, the correlation between the species richness of individual taxa and the summer air temperatures approaches the maximally possible values (Chernov, 2002). Besides differences in the species composition of bumblebees, the dominance status of some species also changes along the latitudinal transect. This can be accounted for by the influence of climatic factors, since each species due to its biological peculiarities is confined to one or several natural zones. On the other hand, a great influence on the bumblebee taxocenes is exerted by the local biotopic conditions, which impedes analysis of connections between their distribution and climate. The bumblebee species typical of the zonal tundra landscapes are B. (Al.) polaris, B. (Al.) hyperboreus, B. (Al.) balteatus, and B. (Pr.) lapponicus. They are common in the Arctic and Subarctic and are considered as true arctic species by some researchers (Skorikov, 1937; Chernov, 1966; Løken, 1973). However, in the forest-tundra and southern tundra these species do not always occupy the dominant position in the topical complexes of bumblebees (Kolosova and Potapov, 2011). These biotopes often have a large fraction of species that are largely associated with the ENTOMOLOGICAL REVIEW Vol. 94 No. 1 2014
Fig. 2. Relation between the number of species in local bumblebee faunas and the mean July air temperature.
taiga zone but extend into the tundra along the valleys of big rivers, which serve as migration pathways for bumblebees (Kolosova and Potapov, 2011). In ascribing different bumblebee species of the tundra zone to certain zonal groups, we used the classification based on the pattern of their distribution (including the optimum) across the main groups of the northern territories; thus, the species were characterized as hyperarctic, euarctic, hemiarctic or hypoarctic (Chernov, 1978; Chernov and Matveyeva, 2002). In this classification, special significance is attached to distribution of species in zonal communities, quantitative participation in them, and the degree of colonization of different landscapes. Analysis of descriptions of zonal groups (Chernov, 1978) shows that none of the bumblebee species belong to the hyperarctic category. It is under the conditions of polar deserts that hyperarctic forms reveal high cenotic activity, whereas in the tundra zone they live only under intrazonal conditions, in the coldest biotopes. It is evident that polar deserts are not optimal habitats for such poikilothermic organisms as bumblebees. Nevertheless, this does not mean that bumblebees are absent in the polar desert zone. For instance, B. (Al.) polaris was recorded in the north of Greenland (Skorikov, 1937) and on Devon Island in the Canadian Arctic Archipelago (Chernov, 2004). The definition of a euarctic species implies that the optimum conditions for it can be found in the arctic tundras and the northern part of typical tundras. Such species are distributed as far northwards as the polar deserts, whereas in the plain regions they do not reach the southern boundaries of the Subarctic. In their turn, hemiarctic species are absent or scarce in the arctic
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tundra subzone, reach the maximum abundance in the middle and southern part of the typical tundra subzone, and extend as far southwards as the forest-tundra (Chernov, 1978). In the literature (Chernov, 1978; Chernov and Matveyeva, 2002; Berezin and Tkacheva, 2007), B. (Al.) polaris and B. (Al.) hyperboreus were considered euarctic, and B. (Al.) balteatus, a hemiarctic species. The status of B. (Pr.) lapponicus lapponicus is questionable since it is sometimes described as a hemiarctic (Chernov, 1978; Berezin and Tkacheva, 2007) and sometimes as a hypoarctic species (Chernov and Matveyeva, 2002). The category of hypoarctic forms includes species with an optimum in southern tundra, forest-tundra, and northern taiga (especially mountain) regions; besides, such species also penetrate into the southern parts of the typical tundra subzone (Chernov, 1978). The subspecies B. (Al.) lapponicus lapponicus may occupy an intermediate position between these categories. According to the classification of Pekkarinen and Teräs (1993), it belongs to the arcto-boreal complex, i.e., is distributed across the entire tundra zone and partly extends into the taiga. The bumblebee B. (Pr.) lapponicus glacialis occurs in the north of Europe only on the Novaya Zemlya Archipelago. Besides this territory, it was also recorded on Wrangel Island (Berezin, 1992, 1995), i.e., it inhabits only the Arctic tundra zone and is therefore a typical euarctic species, unlike the nominotypical subspecies B. (Pr.) lapponicus lapponicus. The temperate latitudes of the study region are mainly inhabited by polyzonal species, for instance, B. (Th.) pascuorum (table). According to the published data (Pekkarinen and Teräs, 1993; Byvaltsev, 2009), most species in the north of European Russia belong to the temperate group (table), i.e., they are distributed from the forest-tundra to the steppe zone (Gorodkov, 1984). The arcto-temperate and boreal species are less represented. Of the subboreal species, only B. (Th.) humilis is present. Thus, species richness of bumblebees in the north of European Russia is the greatest in the middle taiga subzone, diminishes significantly from south to north, and reaches its minimum in the Arctic tundra. The true arctic species are B. (Al.) polaris, B. (Al.) balteatus, B. (Al.) hyperboreus, and B. (Pr.) lapponicus glacialis, whereas the nominotypical subspecies B. (Pr.) lapponicus lapponicus belongs to the arcto-boreal complex. The remaining species belong to the arctotemperate, temperate, boreal, and subboreal groups.
ACKNOWLEDGMENTS The work was financially supported by the Presidential grant MD-4164.2011.5, the Russian Foundation for Basic Research (grants 13-04-00652, 12-04-00594-a, 11-04-98815-r_sever_a, 11-04-98817r_sever_a, 13-04-10107-k), the Ural Branch of RAS (grants 12-P-5-1014, 12-U-5-1022, 12-M-45-2062, 12-5-7-009), the Federal Target Program “Cadres” (8660, 8774, 14132211023), the “Young Scientists of Pomorye 2013” Project (05-2013-03a), and the Departmental research program 546152011. REFERENCES 1. Atlas of Arkhangelsk Province (Moscow, 1976) [in Russian]. 2. Berezin, M.V., “Relations between Bumblebees and Polar Foxes on Wrangel Island,” in Cenotic Relations in Tundra Ecosystems (Nauka, Moscow, 1992), pp. 90–99 [in Russian]. 3. Berezin, M.V., “Bumblebees in Arctic Ecosystems,” in Ecosystems of the North: Structure, Adaptation, Stability. Proc. of All-Russia Workshop, Petrozavodsk, 26–28 October 1993 (Moscow State Univ., Moscow, 1995), pp. 43–57 [in Russian]. 4. Berezin, M.V. and Tkacheva, E.Yu., “On the Study of the Fauna and Ecology of Bumblebees (Hymenoptera: Apidae, Bombini) in the West of Putorana Plateau,” in Biodiversity of the Ecosystems of the Putorana Plateau and Adjoining Territories (Putorana Nature Reserve, Moscow, 2007), pp. 234–245 [in Russian]. 5. Bolotov, I.N. and Kolosova, Yu.S., “The Formation of Topical Complexes of Bumblebees in the North Taiga Karst Landscapes of the Western Russian Plain,” Ekologiya, No. 3, 173–183 (2006). 6. Bolotov, I.N. and Kolosova, Yu.S., “Local Faunas of Bumblebees (Hymenoptera: Apidae, Bombini) in the North of European Russia: the South Timan,” Vestnik Pomor. Univ. Ser. Estestv. Tochnye Nauki, No. 1, 28–39 (2007). 7. Bolotov, I.N. and Podbolotskaya, M.V., “Local Faunas of Bumblebees (Hymenoptera: Apidae, Bombini) in the North of European Russia. The Solovetsky Islands,” Vestnik Pomor. Univ. Ser. Estestv. Tochnye Nauki, No. 1 (3), 74–87 (2003). 8. Byvaltsev, A.M., Bumblebees (Hymenoptera: Apidae, Bombini) in the Forest-Steppes and Steppes in the South of the West Siberian Plain: the Fauna and Population. Candidate’s Dissertation in Biology (Novosibirsk, 2009). 9. Carolan, J.C., Murray, T.E., Fitzpatrick, U., et al., “Color Patterns do not Diagnose Species: Quantitative Evaluation of a DNA Barcoded Cryptic Bumblebee Complex,” PLoS ONE, No. 7, 662–667 (2012). ENTOMOLOGICAL REVIEW Vol. 94 No. 1 2014
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