Åâðàçèàòñêèé ýíòîìîë. æóðíàë 2(4): 235–242 © EUROASIAN ENTOMOLOGICAL JOURNAL, 2003 Distribution patterns of in different natural zones and landscapes in Kazakhstan and West 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 ecology, Institute of Systematics and Ecology of , 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 species in Introduction the Kazakhstan–West 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 it’s 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 1–3 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, 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 4–6 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 &&&&& & M. sabuleti K. Arn. P M. lobicornis Nyl. PPPPP P P S M. deplanata Ruzs. PPWWPPPP&VV V,S M. schenñki Em. PPPPPT & M. sulcinodis nigripes S Messor clivorum Ruzs. PPPWPPP V Messor denticulatus K.-Ug. PP Messor marikovskii K. Arn. PP M. aralocaspius infumatus K.-Ug. PP V,P Tetramorium caespitum L. T P PWWWWWWW V & &WWW T. schneideri Em. VW T. indocile Sanschi WT T. inerme Mayr VT P Leptothorax tuberum &abr. & L. nassonovi Ruzs. PP P L. acervorum &abr. PPP L. muscorum Nyl. VVVV L. balchaschensis K. Arn. VVV L. serviculus Ruzs. PP Solenopsis fugax Latr. PPWVV&&&&&& Crematogaster subdentata Mayr &V Cardioconlyla elegans Em. & C. stambuloffi Ruzs. & Pheidole pallidula Nyl. P Strongylognathus christophi Em. PP ormica rufa L. WVP . polyctena &oerst. WWW P P P S P,S . aquilonia Yarr. TP . lugubris Zett. T . pratensis Retz. VVPPPWWWWVVVVV T,SSS,V . sanguinea Latr. WW T P P P 238 Zh.I. Reznikova

Table 1. (continuation). Òàáëèöà 1. (ïðîäîëæåíèå).

Species / Numbers of profiles1 2 3 4 5 6 7 8 9 101112131415161718

ormica cinerea Mayr P . fusca L. WWW P P S . picea Nyl. PV S,V . rufibarbis &abr. PPW W T TW . cunicularia glauca Ruzs. W P P PWWWW WS,VS,V . pressilabris Nyl. WPPP . exsecta Nyl. VV&PPPPV T . brunneonitida Dluss. VT,S Camponotus herculeanus L. TTPPP C. saxatilis Ruzs. WWW P C. vagus Scop. &T,V C. japonicus Mayr VTT C. interjectus Mayr PSP C. turñestanus Ern. Andre. PPP TTWV C. turñestanicus Mayr V C. lameerei Mayr & C. fedchenkoi Mayr T Lasius fuliginosus Latr. PPP & & L. niger L. WWWWWW &&,P& & V VS, VS,V L. flavus &abr. WWWWWW & & V V V L. alienus &oerst. P P PWPWPW W WSS,P Cataglyphis aenescens Nyl. PWWWWW W W VS,T V C. setipes turcomanica Em. P C. pallida Mayr P Proformica epinotalis K.-Ug. &PVS,PS P. mongolica Dluss. T Polyergus rufescens Latr. PPVV Plagiolepis sp. TPP TVW S,P Dolychoderus quadripunctatus L. & Tapinoma sp. VV SS general number of species 15 15 21 23 24 24 20 22 17 17 12 15 18 23 19 16 13 13

Indications. 1–18 — valley profiles in the vicinity of the following villages and towns (see ig. 1): South taiga: 1 — the village Ust-Ischim, 2 — the village Tevris, 3 — the village Artin; Northern forest-steppe: 4 — the village Krutaya Gorka; Southern forest- steppe: 5 — the village Cherlak, 6 — the town Karasuk; Steppe: 7 — the town Pavlodar, 8 — the town Ayagus; Semidesert: 9 — the town Charsk, 10 — the village Kurchum, 11 — nord-west Zaysan land, 12 — lower sream of Kolguti valley, 13 — Black Irtysh valley, the village Buran; Desert: 14 — Ili valley, village Bakanas, 15 — Chu valley, the village urmanovka; Mountain slopes and mountain valleys: 16 — upper stream of Kolguti river, 17 — slopes of Kungey and Tarbagatay ranges, 18 — Zhalmara valley. P — plain, divide; T — middle terraces, V — valleys, M — flood plains, W — widely spread, S — mountain slopes. Îáîçíà÷åíèÿ. 1–18 — äîëèííûå ïðîôèëè â îêðåñòíîñòÿõ ñëåäóþùèõ íàñåë¸ííûõ ïóíêòîâ (ñì. ðèñ. 1): þæíàÿ òàéãà: 1 — ñ. Óñòü- Èøèì, 2 — ñ. Òåâðèç, 3 — ñ. Àðòûí; ñåâåðíàÿ ëåñîñòåïü: 4 — ñ. Êðóòàÿ Ãîðêà; þæíàÿ ëåñîñòåïü: 5 — ñ. ×åðëàê, 6 — ã. Êàðàñóê; ñòåïü: 7 — ã. Ïàâëîäàð, 8 — ã. Àÿãóç; ïîëóïóñòûíÿ: 9 — ã. ×àðñê, 10 — ñ. Êóð÷óì, 11 — ñåâåðî-çàïàäíûé áåðåã îç. Çàéñàí, 12 — íèæíåå òå÷åíèå ð. Êîëãóòû, 13 — äîëèíà ׸ðíîãî Èðòûøà, ñ. Áóðàí; ïóñòûíÿ: 14 — äîëèíà ð. Èëè, ñ. Áàêàíàñ, 15 — äîëèíà ð. ×ó, ñ. Ôóðìàíîâêà; ãîðíûå ñêëîíû è äîëèíû ãîðíûõ ðåê: 16 — âåðõíåå òå÷åíèå ð. Êîëãóòû, 17 — ñêëîíû õð. Êóíãåé è Òàðáàãàòàé, 18 — äîëèíà ð. Æàëìàðà. P — ïëàêîðû, Ò — ñðåäíèå òåððàññû, V — äîëèíû, M — ïîéìû, W — øèðîêîå ðàñïðîñòðàíåíèå, S — íèçêîãîðíûå ìåæäóðå÷üÿ. Distribution patterns of ants in different natural zones 239 zone. They are able to contend with the climate of the The deserts (profiles 14–15 in Table 1). There are forest tundra (Leptothorax acervorum, Myrmica lobi- northern sand desert landscapes in the middle and up- cornis) and combine with the extremely xerophilous stream of Ile, in the Sari Ishik Sands: saxauls grow on Cataglyphis aenescens and Myrmica deplanata. the plains, willow underbushes and salt-tree with canes The steppe zone (profiles 7, 8 in Table 1). In the grow in the flood-plains, and salt meadows occur on northern outskirts of this zone there is a transitional the terraces. strip with the birch-aspen forest islets within xerophil- The ant species richness is higher here than in the ous vegetation composed mainly of feather-grass and neighbouring northern zones; in all, 24 species were sheep’s fescue. Dry steppe extends in the south from found (Table 1). On the plains, besides the common Pavlodar. In the Ayagus valley there are low steppe species of semi-deserts and steppes (such as Campono- hills covered with feather grass, sheep’s fescue and tus turcestanus, Cataglyphis aenescens, Proformica wormwood. Typical inhabitants of the divides are Cata- epinotalis, Plagiolepis sp.), there are several species glyphis aenescens, Myrmica deplanata, M. schenki, which are only found in this zone, such as Tetramori- Messor clivorum, Leptothorax serviculus, L. nassonovi, um schneideri, Camponotus fedchenkoi, Messor ar- Tetramorium caespitum, Lasius alienus, ormica cu- alocaspius infumatus and M. denticulatus. nicularia and . rufibarbis. Some species which dwell The mountain slopes and valleys (profiles 16–18 here in the valleys are also encountered in the forest- in Table 1). In the lower and middle reaches of the steppe and even in forest divides from adjacent zones: Kolguti river on the inland slopes Messor clivorum, Lasius niger, L. flavus, ormica fusca, . exsecta, Myrmica deplanata, Cataglyphis aenescens, Campono- . polycten and Myrmica rubra. On the sand dunes of tus turcestanus, C. japonicus, Tetramorium caespitum, the high Ayagus flood-plain, Proformica epinotalis and ormica pratensis, . cunicularia and L. niger occur. ormica brunneonitidae occur, the latter being found Ant diversity increases in the upper Kolguti. The near- by Dlussky [1967] in Mongolia and Tuva. ness of the Mongol part of Altai and Northern China The semi-deserts (profiles 9–13 in Table 1). In the influences the ant fauna here. There are numerous nests Zaysan hollow there is xerophytic vegetation composed of . brunneonitida, . picea and Camponotus vagus. of wormwood and sheep’s fescue and common salt- It is interesting that . uralensis replaces . pratensis wort plant communities on plains and in the flood- in the vicinity of the mountain lake Markacol. In the plains. Ant species’ communities in the flood-plains Karabuga valley, in the internal Tarbagatay foothills, differ somewhat from those of the divides. In the flood- Myrmica deplanata, M. lobicornis, M. schenki, . pi- plains the species composition is similar to that of the cea, . mesasiatica, . rufibarbis and Tetramorium divides in the more northern forest-steppe zone. The caespitum were found. The last two species, together following species are common: Myrmica rubra, with numerous . pratensis and Camponotus vagus, M. salina, Solenopsis fugax, ormica pratensis, were encountered on the meadows in the vicinity of the . cunicularia and Lasius fuliginosus. In the poplar foothills. On the arid low front Tarbagatay foothills, forest on the flood-plain of the Black Irtysch Cam- only Camponotus turcestanus and Plagiolepis sp. were ponotus vagus and Dolichoderus quadripunctatus were found. The ant fauna of the river valleys from the found, species which are known from deciduous for- Alakol and Balkhash basins includes Turan species ests of the Caucasus and the Crimea. The typical com- (Myrmica bergi, M. stangeana, Tetramorium inerme, munity of the plains and low mountains is composed of Crematogaster subdentata, Camponotus turcestanicus, Camponotus turcestanus, Messor clivorum, Proformi- C. lameerei), Kazakhstan species (Tetramorium indo- ca mongolica, Myrmica deplanata, Plagiolepis sp. and cile, Leptothorax balchaschensis) and Mediterranean Tapinoma sp. species (Cardiocondyla elegans). The specific feature of the ant fauna in undercur- rent of Black Irtysh and Kurchum is the high diversity THE CONSTANCY O( SPECIES COMMUNITIES DOMI- and density of Camponotus species: C. vagus is numer- NATED BY ORMICA PRATENSIS ous in the flood-plains and low terraces, C. japonicus Comparing Table 1 and Table 2, one can see that is numerous in the salt meadows of middle terraces, those species which have wide natural habitats, central C. turcestanus on the plains, and C. interjectus on the parts of which fall in the forest-steppe and the steppe, slopes of low mountains. are included in the same species communities not only The North-West Zaysan land is the most arid area within the central parts of their natural habitats but also here. The small river Kolguti flowing down from Kur- in the forest zone and in the northern deserts (e.g. chum, dries up and bores into the crumbly alluvium. In ormica pratensis, . cunicularia, Lasius alienus, Myr- its flood-plain there are meadows composed of Span- mica salina, M. deplanata, M. scabrinodis, Tetramori- ish liquorice and grisebach. Myrmica bergi, M. stange- um caespitum). When analysing the inter-relationships ana, M. salina, M. schencki, ormica cunicularia, La- of the ethological dominant . pratensis with other sius niger and L. flavus are found here, as well as species lacking protected food territories, one can see Tetramorium caespitum and Cataglyphis aenescens in that near the edge of its natural habitat, . pratensis the higher parts. On the slopes and plains, only 3 spe- becomes a member of species’ communities differing cies were encountered: C. aenescens, Messor clivorum from those in the central part of its area. Thus, in the and Plagiolepis sp. northern part of the area this species enters an impover- 240 Zh.I. Reznikova

Table 2. Number of ant’ nests per one feeding territory of the dominant /ormica pratensis in different zones (average values recalculated per the same square). Òàáëèöà 2. ×èñëî ãí¸çä ìóðàâü¸â ðàçíûõ âèäîâ, ïðèõîäÿùèõñÿ íà îäèí êîðìîâîé ó÷àñòîê /ormica pratensis â ðàçíûõ çîíàõ (ïðèâåä¸ííîå ê îäèíàêîâîé ïëîùàäè).

Number of ant's nests / numbers of profiles 1245679101314 ormica fusca 1.2±0.4 2.3±0.8 . cunicularia 4.2±1.3 5.3±1.8 5.7±2.1 7.3±2.4 8.3±3,2 5.7±1.9 . rufibarbis 6.2±2.9 5.1±2.2 Lasius niger 1.7±0.8 2.3±0.7 L. alienus 4.7±1.9 11.8±4.9 12.2±5.4 10.2±2.7 12.1±3.4 7.5±3.3 8.1±3.1 Myrmica ruginodis 5.1±1.8 6.2±2.8 M. scabrinodis 8.2±3.1 6.7±2.8 8.9±3.7 7.3±4.2 M. schenki 8.3±3.5 3.9±1.8 6.2±1.9 5.3±1.4 6.2±0.9 M. salina 10.4±3.8 12.7±5.2 8.2±1.9 5,2±1.3 6.2±2.4 4.1±2,2 M. deplanata 4.2±2.1 5.2±3.4 10.2±3.9 8.9±2.7 5.6±1.9 5.1±0.9 13.2±4.1 M. bergi 3.1±0.9 Tetramorium caespitum 4.3±0.8 6.3±3.4 10.2±4.3 15.2±4.8 11.1±3.8 6.1±1.8 7.1±3.5 3.1±1.3 T. inerme 4.2±0.9 T. indocile 6.1±2.3 T. schneideri 1.7±0.5 Camponotus turcestanicus 2.1±0.7 Plagiolepis sp. 2.2±0.9

Indications. Numbers of profiles correspond to Table 1 and ig. 1. Îáîçíà÷åíèÿ. Íîìåðà äîëèííûõ ïðîôèëåé ñîîòâåòñòâóþò òàáëèöå 1 è ðèñ. 1. ished variant of a forest species community. Actually, (M. scabrinodis, M. salina, M. schencki), almost the such steppe species as . cunicularia, . rufibarbis, same for Lasius alienus and 10–15 nests of T. caespi- L.alienus, M. salina, M. deplanata and T. caespitum tum. In semi-desert and desert landscapes there is a do not extend into the forest zone. At the same time the somewhat different proportion as a result of a lack of forest species Myrmica ruginodis, M. rubra, M. rugu- L. alienus and decreasing numbers of T. caespitum but, losa, ormica fusca and Lasius niger are numerous in in general, a constant image of species community is the typical zonal landscapes, where . pratensis is maintained. rather rare. Thus, . pratensis is only observed with small numbers of . fusca, M. ruginodis (and M. schenki Discussion in some cases) and L. niger (profiles 1, 2 in Table 2) . On the contrary, in the flood-plains of the desert rivers, Analysing ant distributions across the vast region . pratensis enters an enriched variant of species com- covering several zones, one can see that the arid zone munity. Here are not only steppe species but also those as well as the forest-steppe zone support more diverse which dwell in intrazonal landscapes together with . ant fauna than other zones: 29 species were found in pratensis, namely Myrmica bergi, Tetramorium inerme, the deserts (valley profiles 14, 15) and 31 species in T. indocile, T. schneideri, Plagiolepis sp., and Cam- the forest steppe (profiles 4–6). (or comparison, there ponotus turcestanicus . Therefore, the species richness are 24 species in the forest zone (profiles 1–3) and 25 in the communities dominated by . pratensis increas- in the semi-desert (profiles 9–13). es from north to south, from 3 to 9 species. The ranges of more than half of the observed species It is interesting that in the central part of . praten- are restricted to a single zone; for example, Myrmica sis’ natural habitat, not only is species composition ruginodis, ormica rufa, . aquilonia and . lugubris maintained, but also the numerical proportion of nests coincide with the forest zone, while Leptothorax nas- of different species which dwell in the feeding territo- sonovi and L. serviculus only coincide with the steppe. ries of the dominant. There are 4–8 mounds of ormi- The greater part of the group of species with locally ca cunicularia or . rufibarbis, 5–12 Myrmica nests restricted distributions consists of stenothermic Turan Distribution patterns of ants in different natural zones 241 species coinciding with the deserts, such as Tetramori- have overlapping ecological niches. This should be um schneideri, T. inerme, T. indocile, Myrmica stan- taken into account when considering ant conservation, geana and all species of the genus Messor. not only of key species but also of ant communities as a It is interesting to consider patterns of distribution whole. in those species which possess exceptionally vast natu- ral habitats covering several zones; for example, or- Acknowledgments mica pratensis, . cunicularia, Tetramorium caespi- tum, Lasius niger and Myrmica deplanata are I am grateful to Prof. Donald Michie and Dr Annie widespread from the forest to the arid zones. It is Lamb for their helpful comments on this paper. This work probable that the main mechanism for the expansion of was supported by a grant from the Russian (und for Basic range in these species is the zonal change of habitats, Research (02–04–48386) and from the fund of Dr (rank as discovered by Bey-Bienko [1930] for Acrididae Salter (Humanethologie und Humanwissenschaftliches Zen- when he considered their distributions in West Siberi- trum der Ludwig — Maximilians Universitaet, Muenchen). an and Zaisan Plains. In those ant species which have wide distributions, central parts of which lie in the References forest-steppe and steppe zones, the zonal change of habitats is displayed both in southern and northern Andersen A.N. 1993. Ant Communities in the Gulf Region of Australia’s Semi-arid Tropics: Special Composition, Patterns of directions. Organisation, and Biogeography // Australian Journal of Zool- Thus, . pratensis in the steppe and in the southern ogy. Vol.41. 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