Åâðàçèàòñêèé ýíòîìîë. æóðíàë 2(4): 235242 © EUROASIAN ENTOMOLOGICAL JOURNAL, 2003 Distribution patterns of ants in different natural zones and landscapes in Kazakhstan and West Siberia along a meridian trend Õàðàêòåð çîíàëüíîãî è ëàíäøàôòíîãî ðàñïðåäåëåíèÿ ìóðàâüåâ íà ìåðèäèîíàëüíîì ðàçðåçå ÷åðåç Çàïàäíóþ Ñèáèðü è Êàçàõñòàí Zh.I. Reznikova Æ.È. Ðåçíèêîâà Laboratory of insect ecology, Institute of Systematics and Ecology of Animals, Russian Academy of Sciences, Siberian Branch, (runze str. 11, Novosibirsk 630091 Russia; Novosibirsk State University, Pirogova str. 2, Novosibirsk 630090 Russia. URL: http://fen.nsu.ru/~rezzhan Ëàáîðàòîðèÿ ýêîëîãèè íàñåêîìûõ, ÈÑÝÆ ÑÎ ÐÀÍ, óë. Ôðóíçå 11, Íîâîñèáèðñê 630091 Ðîññèÿ; Ôàêóëüòåò åñòåñòâåííûõ íàóê, Íîâîñèáèðñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, óë. Ïèðîãîâà 2, Íîâîñèáèðñê 630090 Ðîññèÿ. Abstract. Distribution patterns of 70 ant species in Introduction the KazakhstanWest Siberian along a meridian trend from deserts to the southern taiga is described. The (inding assembly rules, i.e. orderliness in the pat- arid zone, as well as the forest-steppe zone, support tern of species coexistence as well as the mechanisms most diverse ant fauna. Natural habitats of more than that affect coexistence, is one of the major challenges half of the observed species are entirely restricted to in community ecology. This knowledge is extremely one zone. Those species which possess wide natural important for species conservation since key species habitats covering several native zones display similar are often closely associated with other members of the patterns of zonal change of habitats [in a sense of Bey- species community and, in some situations, may be less Bienko, 1930]. Assemblages in which ormica pratensis viable in the absence of these other members. was dominant were considered as the main example. Ants, with their great abundance and easily ob- Such communities maintain a constancy of species compo- served social behaviour, are superb organisms for stud- sition from the forest zone to the southern boundaries ying the organization of species communities. Domi- of the steppe zone. This should be taken into account nance hierarchy among different ant species is one of when considering the conservation of ants, not only of the most complex mechanisms that increases species key species but also of ant communities as a whole. richness. Kaczmarek [1953] was the first to use the terms «dominant», «subdominant» and «influent» in Ðåçþìå. Âûÿâëåíî çîíàëüíîå è ëàíäøàôòíîå their ethological sense to explain ant community struc- ðàñïðåäåëåíèå 70 âèäîâ ìóðàâüåâ íà ìåðèäèîíàëü- ture. Although there have been numerous studies on íîì ðàçðåçå ÷åðåç Çàïàäíóþ Ñèáèðü è Êàçàõñòàí è community structure [e.g. Brain, 1965; Wilson, 1971; ïðîàíàëèçèðîâàí õàðàêòåð çîíàëüíîé ñìåíû ìåñ- Greenslade, 1971; Reznikova, 1971, 1983, 1998a,b, òîîáèòàíèé ðàçíûõ âèäîâ ïî Áåé-Áèåíêî [Bey- 1999; Galle, 1975; (ox, (ox, 1982; Elmes, Lepage, Bienko, 1930] è ñòðóêòóðà ìíîãîâèäîâûõ ñîîáùåñòâ. 1994; Deslippe, Savolainen, 1995], surprisingly little Ìàêñèìàëüíîå âèäîâîå áîãàòñòâî îòìå÷åíî äëÿ ïó- work has been devoted to investigating species compo- ñòûííîé è ëåñîñòåïíîé çîí. Âûÿñíåíî, ÷òî âèäû sition. Meanwhile stable long standing ant communi- ìóðàâüåâ ñ øèðîêèìè àðåàëàìè îáëàäàþò ñõîäíû- ties have been revealed: Dlussky [1981] has investigat- ìè òåíäåíöèÿì çîíàëüíîé ñìåíû ìåñòîîáèòàíèé.  ed such communities in deserts and named them êà÷åñòâå íàèáîëåå ïîêàçàòåëüíîãî ïðîàíàëèçèðî- «co-adaptive complexes», Andersen [1993, 1997] and âàíà ñòðóêòóðà ñîîáùåñòâ ñ äîìèíèðîâàíèåì ëóãî- Herbers [1994] described patterns of organization of âîãî ìóðàâüÿ ormica pratensis. Ïðîäåìîíñòðèðî- ant communities in different arid zones in Australia âàíî, ÷òî ýòè ñîîáùåñòâà ñîõðàíÿþò îñíîâó and compared them with those in North America, and âèäîâîãî ñîñòàâà è ñòðóêòóðíóþ öåëîñòíîñòü íà Savolainen et al. [1989] have investigated many boreal áîëüøåé ÷àñòè àðåàëà äîìèíàíòà, îò ëåñíîé çîíû ant communities. äî ïîëóïóñòûíü. Âûÿâëåííûå çàêîíîìåðíîñòè äîë- There is a very intriguing situation in West Siberia æíû áûòü ïðèíÿòû âî âíèìàíèå ïðè ðàçðàáîòêå and Kazakhstan where the ethological dominant of ïîäõîäîâ ê ñîõðàíåíèþ ìóðàâü¸â, òàê êàê ìåæâè- steppe ant communities ormica pratensis is äîâîå îêðóæåíèå, ñ åãî ñïåöèôè÷åñêèìè ñâÿçÿìè, spread from the northern taiga to deserts. In our previ- ñîñòàâëÿåò åñòåñòâåííóþ áèîòè÷åñêóþ ñðåäó äëÿ ous studies [Stebaev, Reznikova, 1972; Reznikova, äîìèíèðóþùèõ âèäîâ. 1975, 1982, 1994, 1998a,b; Reznikova, Kulikov 1978; 236 Zh.I. Reznikova Reznikova, Bogatyreva 1984; Reznikova, Samoshilo- Ayaguz, Kurchum, Kolguti, Black Irtysch and Ile, as va, 1981] we demonstrated that . pratensis, dominat- well as the Alakol and Balkhash basins were investi- ing in species communities, has a fundamental influ- gated (see (ig. 1 and Table 1). There were 18 valley ence on the spatial distribution, daily activity, as well profiles and 5 main geomorphological levels within as composition and quantity of prey in those ant spe- each profile: lower and upper flood-plains, lower and cies which dwell in its feeding territories. The situa- upper terraces and the divide. tion mentioned above provides an opportunity to con- Small-size ant species nests were counted in 5x5 m sider changes in the structure of ant communities grouping plots. There were 15 plots per each geomorphologic around the same dominant species along huge area. level. Anthills of the subgenus ormica s.str. as well as The results of long term investigation of ant fauna underground nests of the genus Camponotus were count- and community structure in West Siberia and Kaza- ed on 2 km record belts, three for each geomorphologic khstan are presented here. Earlier the ant fauna in Sibe- level. Inventories were completed by thorough direct ria was studied mainly in forest biomes. Investigations searching. of Ruzsky [1905, 1907], Karavajev [1912], Kuznet- To estimate the constancy of species communities, zov-Ugamsky [1926], Dmitrienko, Petrenko [1976] only assemblages dominated by . pratensis were in- were devoted to the forest environment of Tomsk, Omsk vestigated. (or 10 of the 18 valley profiles such assem- and Tobolsk, the Yenisei valley and the Baikal Basin. blages were detected on 6 plots (each of area Marikovsky [1979] studied ant fauna of desert regions 1600 m2) 60 plots in total. To compare the structure in Kazakhstan. The ant fauna in the greater part of the of the different assemblages, a relative value was used: area under investigation here was unknown. the number of nests of each species per one feeding territory of . pratensis. This value was recalculated Materials and methods for the same standard area corresponding to the aver- age feeding territory of . pratensis, namely, 800 m2. The ant fauna was investigated along a meridian The average values from 6 measurements are provided in trend across the southern part of the West Siberian Table 2. Plain and adjacent regions to the south. To estimate the constancy of ant species communities, the work con- Results centrated on the communities dominated by ormica pratensis. ANALYSIS O( THE ANT (AUNA The valleys of Irtysch and of some of its tributaries The forest zone (see profiles 13 in Table 1). The (Ishim, Ui, Tui, Tara and Char) and of some rivers inspected territory embraces the southern taiga sub- flowing into Balkhash Lake and Zaysan Lake, namely zone and the birch-aspen forests in the subtaiga sub- zone. Mixed pine forests and high bogs occupy the divides. Herbaceous meadows are usual in the flood- plains. The greatest species richness has been recorded in the birch-aspen forests. Among them there are typi- cal forest species such as ormica rufa, . polyctena, . aquilonia, Myrmica lobicornis and Camponotus vagus, as well as those species which are more usual in the southern regions: . pratensis, . cunicularia, M. scabrinodis and Tetramorium caespitum. Species richness diminishes in the southern taiga because of the disappearance of those species more typical of the forest-steppe, such as . cunicularia and T. caespitum, which were not encountered north of forest-steppe boundaries. The forest-steppe zone (profiles 46 in Table 1). Mosaics of vegetation and soils are characteristic of this zone in the West Siberian Plain due to a combina- tion of high summer temperature and much soil water lying on salt loam. There are mixed grass meadows in the flood-plains and carnation-grass meadows and swamps in the watersheds. The species richness in this zone is higher than in neighbouring ones. The most typical species are . pratensis, . sanguinea, . cu- nicularia, . rufibarbis, M. scabrinodis, M. salina, Tetramorium caespitum and Lasius alienus. In this ig. 1. Location of valley transects 1-18 [from: Reznikova, 1999] (explanations as in Tabl. 1). zone boreal species dwell in proximity to steppe and Ðèñ. 1. Ëîêàëèçàöèÿ äîëèííûõ ïðîôèëåé 1-18 [èç: Rezni- Turan-steppe species. Apart, species with rather differ- kova, 1999] (îáîçíà÷åíèÿ ñì. òàáë. 1). ent preferences for moisture and heat dwell in this Distribution patterns of ants in different natural zones 237 Table 1. Ant diversity in different zones and landscapes. Òàáëèöà 1. Ðàñïðåäåëåíèå ìóðàâü¸â â ðàçëè÷íûõ ïðèðîäíûõ çîíàõ è ïîäçîíàõ. Species / Numbers of profiles1 2 3 4 5 6 7 8 9 101112131415161718 P, Myrmica rubra L. VVPPPP P P V Ò,S M. ruginodis Nyl. TTW V M. bergi Ruzs. && M. rugulosa Nyl. VVPPV& T M. stangeana Ruzs. VV M. slovaca Sad. P M. scabrinodis Nyl. VPPPPPP S M. salina Ruzs. PW &&&&&