NORTH-WESTERN JOURNAL OF ZOOLOGY 7 (1): pp.26-34 ©NwjZ, Oradea, Romania, 2011 Article No.: 101105 www.herp-or.uv.ro/nwjz

Zoogeography, diversity and altitudinal distribution of (: Formicidae) in the Mediterranean and the oro-Mediterranean parts of

Marko G. KARAMAN

Natural History Museum of Montenegro, P.O.Box 374, Podgorica 20000, Montenegro (Crna Gora). E-mail: [email protected]

Received: 22. June 2010 / Accepted: 14. December 2010 / Available online: 17. December 2010

Abstract. The species number, zoogeographical composition and vertical distribution of the fauna of the Mediterranean and the oro-Mediterranean part of Montenegro were investigated. A total number of 87 spe- cies and subspecies belonging to 28 genera and 4 subfamilies (Ponerinae, , Dolichoderinae, For- micinae) are recorded. The species belong to 15 zoogeographical elements within the three main zo- ogeographical zones: Mixed and deciduous forest zone; Mediterranean zone; and Coniferous high mountain- ous forest zone. Overall species richness is inversely proportional with the altitude. The Mediterranean ele- ments are dominant at the coastal region, and almost absent above 500 m altitude. The species richness of the mixed and deciduous forest zone was the highest at about 700 - 1100 m altitude. The coniferous element is scarce due to the sporadic true coniferous forests in the oro-Mediterranean zone of Montenegro.

Key words: ants, Formicidae, zoogeography, vertical distribution, Montenegro.

Introduction al. 1998, Karaman 1999, Karaman & Karaman 2006). Papers dealing with the zoogeography of the Zoogeography and vertical distribution of ants European myrmecofauna are very few, and refer in Balkan Peninsula, and Dinarides as part of Bal- to the biogeography of the North-European kans, up to now remained poorly known. The zo- (Baroni-Urbani & Collingwood 1977) and Polish ogeography of the myrmecofauna was studied (Chechowski et al. 2002) myrmecofauna. Similarly, only in Slovenia (Bračko 2007). The study of ants the species richness along the altitudinal gradient diversity in this region was recently a subject of of the European mountains has been poorly inves- research of several scientists: Bračko in Slovenia tigated, with only one study of the ant fauna of the (2000) and Croatia (2006); Atannasov & Dlusskij in Alps (Glasser 2006). This study showed that the Bulgaria (1992); Karaman in Macedonia (2000, number of species decreases with the altitude, as 2002); Markó et al. in Romania (2006); Petrov expected. Similar correlation was observed in the (2004), Karaman & Karaman (2003, 2007) in ; tropical regions (Brown 1973, Olson 1994, Sanders Karaman (2004; 2008), Karaman & Karaman (2005, 2002). In contrast, Sanders et al. (2003) observed a 2006) in Montenegro. The objectives of our re- significant increase of species richness of ants in search are to study the diversity, zoogeography arid canyons of Nevada at higher elevations. This and vertical distribution of Formicidae in the relationship was explained by lower temperatures coastal mountain region of Orjen, Lovćen and and higher precipitation at higher elevations in Rumija Mts. in Crna Gora (Montenegro). this ecosystem which favoured species richness of ants. Schlick-Steiner et al. (2008) found positive Materials and Methods correlation between ant species richness and hu- man population size and density, plant species Field collections of myrmecofauna in the Mediterranean richness and mean annual temperature, but nega- and the oro-Mediterranean part of Montenegro were tive correlation with latitudinal gradient in the conducted during the period between 1985 and 2005. In- myrmecofauna of different European countries. vestigated area included 115 localities from the narrow In comparison to Western Europe, Balkan Adriatic coast of Montenegro, 283 km long, and the chain of the coastal mountains: Orjen, Lovćen, and Rumija (Fig. fauna, including myrmecofauna, is insufficiently 1). Specimens were collected mostly from the nests. In the known. Previous myrmecological investigations of case the nests were not found, solitary specimens were this region included primarily faunistic studies collected from the ground, trees or shrubs. Material was with scarce ecological records (Müller 1923, preserved in 70% ethyl alcohol and deposited in collec- Zimmermann 1934, Petrov 1993, 1995, Karaman et tion of Natural History Museum of Montenegro, Pod- Zoogeography and distribution of ants (Hymenoptera: Formicidae) in Southern Montenegro 27

.

Figure 1. A– Location of Montenegro in Southeastern Europe; B– The map of investigated area in Montenegro. Maps created using DIVA-GIS Version 5.4 (Hijmans et al. 2004).

gorica, Montenegro and in Karaman's collection. Taxo- nean zone; and three species (3.45%) belong to the nomic nomenclature followed that of Bolton et al. (2006), coniferous zone. Quantitatively, the most impor- Schlick-Steiner et al. (2006a: Tetramorium cf. caespitum), tant zoogeographical elements are: Central and Schlick-Steiner et al. (2006b: Messor cf. structor) and Seifert Southern-European (14 spp., 16.09%), Euro-Cau- et al. (2009: hellenica Finzi, 1926). The total of 1130 samples of ants’ colonies were analysed. casian (14 spp., 16.09%) and North-Mediterranean The species are classified into zoogeographical and elements (11 spp., 12.65%). West Palaearctic (9 vegetation zones, and appropriate zoogeographical ele- spp., 10.34%), Transpalaearctic (7 spp., 8.04%), ments. Zoogeographical elements were determined from Euro-Siberian (5 spp, 5.74%), while Holomediter- the literature (Czechowski et al. 2002). ranean elements (4 spp, 4.59%) have a minor role. Correlation between the altitude and (1) the total The least important are Balkan (2 spp, 2.29%) and number of species, and (2) the number of species belong- the Southeast-European elements (1 spp., 1.14%) ing to zoogeographical zones and elements were tested using non-parametric Spearman Correlations, The analy- (Figs 3 and 4). ses were run in STATISTICA 9.1 (StatSoft, Inc., 2010). Vertical distribution of species in investigated area is presented in Table 2 and Figures 5 and 6. Results of the Spearman Rank Order Correlations Results analyses are shown in Table 3. There is a high level of a negative correlation between total spe- Within the studied region, 85 species and two sub- cies number of ants and altitude. Such negative species, belonging to 28 genera and four subfami- correlation is also observed in the number of lies (Ponerinae, Myrmicinae, Dolichoderinae and speices in the investigated zoogeographical zones ) were collected. These species belong and corresponding elements. The only exceptions to three main zoogeographical zones correspond- are the Transpalaearctic (south) and the Southern ing to the three main vegetation zones of the European elements which exhibited no correla- Palaearctic: 1. Mixed and deciduous forest zone; 2. tion, and the Boreo Montane which had a positive Mediterranean zone; and 3. Coniferous high correlation with the altitude. mountainous forest zone. The correlation between total number of spe- The Mixed and deciduous forest zone includes cies and altitude is not linear, exhibiting an inter- nine zoogeographical elements, the Mediterranean mediate peak between 700 and 1100 m a.s.l. (Fig. zone six elements, and Coniferous high moun- 5). While the number of species belonging to the tainous forest zone includes one element (Table 1). Mediterranean zone decreases almost linearly, that The cosmopolitan zoogeographic element does not of the mixed and deciduous forest zone follows belong to any of the zones. the total species richness pattern (Fig. 6). The More than half of the investigated ant species Euro-Caucasian, European and the Central and (56 spp., 64.37%) is associated with the class of the Southern European elements show two elevation mixed and deciduous forest zone; almost one third peaks, between 100 and 300 m a.s.l., and between (26 spp., 29.88%) are assigned to the Mediterra- 700 and 1100 m a.s.l. The number of the Holo-

28 Karaman, M.G.

Table 1. Zoogeographical composition of the ant fauna of the Mediterranean and the oro- Mediterranean part of Montenegro.

Zoogeographical zones Zoogeographical element number of species % Zone of the mixed and de- Holarctic 3 3.45 ciduous forest Transpalaearctic (south) 7 8.04 West Palaearctic 9 10.34 Euro-Caucasian 14 16.09 Euro-Siberian 5 5.75 European 3 3.45 Central and Southern-European 14 16.09 Southeast-European 1 1.15 56 Mediterranean zone Southern-European 3 3.45 Holomediterranean 4 4.59 North-Mediterranean 11 12.64 Balkan 2 2.29 East-Adriatic 3 3.45 Endemics of Montenegro 3 3.45 26 Zone of the coniferous Boreo Montane 3 3.45 high mountainous forest Cosmopolitan 2 2.29 Total number of species 87

Figure 2. Zoogeographical composition of the ant fauna

of the Mediterranean and the oro-Mediterranean part of Montenegro. 1- Zone of mixed and deciduous forest; Figue 4. Number of species in zoogeographical elements 2- Mediterranean zone; 3- Zone of coniferous high belonging ot the Mediterranean zone. 1- Southern- mountainous forest; 4- Cosmopolitan. European; 2 – Holomediterranean; 3- North-Mediterra- nean; 4- Balkan; 5- East-Adriatic; 6- Endemics of Mon- tenegro.

mediterranean and the North Mediterranean spe- cies decreases with the altitude, from the sea level to 400 m a.s.l., and the same species are only spo- radically present at higher altitudes. The species associated with the Boreo-montane element occur only above 1100 m a.s.l.

Figue 3. Number of species in zoogeographical elements Discussion belonging to the Zone of mixed and deciduous forest. 1- Holarctic; 2- Transpalaearctic (south); 3- West Palae- arctic; 4- Euro-Caucasian; 5- Euro-Siberian; 6- Euro- Zoogeographical composition pean; 7- Central and Southern-European; 8- Southeast- Present fauna of Formicidae in Montenegro is in European. strong correlation with the paleogeographic

Zoogeography and distribution of ants (Hymenoptera: Formicidae) in Southern Montenegro 29

changes and climatic-ecological fluctuations of this Sićevo Gorge, and Demir Kapija crag (Lopatin & region. Matvejev 1996). The present fauna of the oro-Mediterranean The West-Palaearctic element survived the and the Mediterranean parts of Montenegro is the glaciations in the refuges of Northern Africa, Asia combination of faunas with different origin. It in- Minor or Balkan Peninsula, and after the retreat of cludes the old Tertiary fauna, the fauna formed the last continental glaciers most of them spread during the glacial period, and the fauna that set- again northwards in the Central Europe. Now tled the mountains after the retreat of the last gla- they are distributed in the Middle East, North Af- ciers. During the glaciation, as well as the rica and in the , some of these interglacial periods, Balkan Peninsula played a species settled also towards the Central Europe. very important role as a bridge between Europe The Euro-Caucasian element is distributed in and Asia Minor, where many of these species sur- the open deciduous forests and dry grasslands of vived glaciations. Consequently, Balkan Peninsula the Southern and partly the Central Europe, as contains several preserved glacial refuges far from well as in the forest zone and the forest-steppe the sea coast, such as the canyon of Cijevna River, zone of Asia Minor. These species during the gla-

Table 2. Vertical distribution of the ants in the Mediterranean and the oro-Mediterranean part of Montenegro. Species arranged in alphabetical order. (ZGE - zoogeographical elements: H – Holarctic; TP(s) - Transpalaearctic (south); WP - West Palaearctic; EC – Euro Caucasian; ES - Euro-Siberian; E – European; CSE - Central and Southern- European; SeE - Southeast-European; SE - Southern-European; HM – Holomediterranean; NM - North-Mediterra- nean; B – Balkan; EA - East-Adriatic; EnM - Endemics of Montenegro; BM - Boreo Montane; C – Cosmopolitan).

Altitude (meters above sea level) No. Species ZGE 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900

1. Aphaenogaster epirotes Emery, 1895 NM

2. Aphaenogaster finzii Müller, 1921 EA

3. Aphaenogaster muelleriana Wolf, 1915 EA

4. Aphaenogaster obsidiana Mayr, 1861 EC

5. Aphaenogaster subterranea (Latreille, 1789) WP

6. Bothriomyrmex adriacus Santschi, 1922 EA

7. Camponotus aethiops (Latreille, 1798) WP

8. Camponotus dalmaticus (Nylander, 1848) B

9. Camponotus fallax (Nylander, 1856) WP

10. Camponotus lateralis (Olivier, 1792) EC

11. Camponotus ligniperdus (Latreille, 1802) BM

12. Camponotus piceus (Leach, 1825) NM

13. Camponotus truncatus (Spinola, 1808) EC

14. Camponotus vagus (Scopoli, 1763) ES

15. Cardiocondyla elegans Emery, 1869 NM

16. Cataglyphis nodus Brullé, 1833 EC

17. Crematogaster auberti savinae Zimmermann, 1934 EnM

18. Crematogaster gordani Karaman, 2008 EnM

19. Crematogaster ionia Forel, 1911 NM

20. Crematogaster jehovae Forel, 1907 NM

21. Crematogaster montenigrinus Karaman, 2008 EnM

22. Crematogaster schmidti (Mayr, 1853) NM

23. Crematogaster sordidula mayri (Mayr, 1853) B

24. Dolichoderus quadripunctatus (Linné, 1771) CSE

25. Formica cunicularia Latreille, 1798 CSE

30 Karaman, M.G.

Table 2. (continued).

Altitude (meters above sea level) No. Species ZGE 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900

26. Formica fusca Linnaeus, 1758 TP(s)

27. Formica gagates Latreille, 1798 EC

28. Formica polyctena Foerster, 1850 ES

29. Formica pratensis Retzius, 1783 TP(s)

30. Formica rufibarbis Fabricius, 1793 ES

31. Hypoponera eduardi (Forel, 1894) TP(s)

32. alienus (Foerster, 1850) H

33. Lasius distinguendus (Emery, 1916) CSE

34. Lasius emarginatus (Olivier, 1791) EC

35. Lasius flavus (Fabricius, 1782) TP(s)

36. Lasius fuliginosus (Latreille, 1798) TP(s)

37. Lasius jensi Seifert, 1982 CSE

38. Lasius lasioides (Emery, 1869) HM

39. Lasius myops (Forel, 1894) CSE

40. Lasius niger (Linné, 1758) H

41. Lasius paralienus Seifert, 1992 E

42. Lasius platythorax Seifert, 1991 ES

43. Lepisiota nigra (Dalla Tore, 1893) NM

44. Leptothorax acervorum (Fabricius, 1793) BM

45. Liometopum microcephalum (Panzer, 1798) EC

46. Manica rubida (Latreille, 1802) BM

47. Messor aralocaspius (Ruzsky, 1902) NM

48. Messor denticulatus Ugamski, 1927 NM

49. Messor cf. structor (Latreille, 1798) EC

50. Messor wasmanni Krause, 1910 NM

51. Monomorium monomorium Bolton, 1987 HM

52. Monomorium pharaonis (Linné, 1758) C

53. Myrmecina graminicola (Latreille, 1802) WP

54. Myrmica hellenica Finzi, 1926 CSE

55. Myrmica sabuleti Meinert, 1861 ES

56. Myrmica scabrinodis Nylander, 1846 TP(s)

57. Pheidole pallidula (Nylander, 1849) C

58. pygmaea (Latreille, 1798) CSE

59. Plagiolepis taurica Santschi, 1920 TP(s)

60. Plagiolepis xene Stärcke, 1936 CSE

61. Ponera testacea Emery, 1895 E

62. Prenolepis nitens (Mayr, 1852) SeE

63. Proceratium melinum (Roger, 1860) EC

64. Solenopsis banyulensis Bernard, 1950 NM

65. Solenopsis fugax (Latreille, 1798) WP

66. Stenamma debile (Foerster, 1850) WP

67. Tapinoma erraticum (Latreille, 1798) WP

68. Tapinoma madeirense Emery, 1925 CSE

69. Tapinoma nigerrimum (Nylander, 1856) EC

Zoogeography and distribution of ants (Hymenoptera: Formicidae) in Southern Montenegro 31

Table 2. (continued).

Altitude (meters above sea level) No. Species ZGE 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900

70. Tapinoma simrothi Krausse, 1911 HM

71. Temnothorax affinis Mayr, 1855 EC

72. Temnothorax crassispinus (Karavaiev, 1926) CSE

73. Temnothorax interruptus (Shenck, 1852) WP

74. Temnothorax nigriceps Mayr, 1855 EC

75. Temnothorax parvulus (Schenck, 1852) WP

76. Temnothorax racovitzai Bondroit, 1918 CSE

77. Temnothorax rottenbergi (Emery, 1870) SE

78. Temnothorax saxonicus (Seifert, 1995) CSE

79. Temnothorax sordidulus (Müller, 1923) SE

80. Temnothorax tuberum (Fabricius, 1775) E

81. Temnothorax unifasciatus (Latreille, 1798) CSE

82. Tetramorium cf. caespitum (Linné, 1758) H

83. Tetramorium diomedeum Emery, 1908 SE

84. Tetramorium ferox Ruzsky, 1903 EC

85. Tetramorium forte Forel, 1903 EC

86. Tetramorium impurum (Foester, 1850) CSE

87. Tetramorium semilaeve André, 1883 HM

Number of species: 70 39 22 19 14 11 12 25 27 22 11 22 12 9 2 6 2 1 Mediterranean zone 23 16 8 7 4 3 2 3 4 2 3 4 1 0 0 0 0 0 Mixed and deciduous forest zone 43 22 13 11 9 7 9 21 22 19 6 18 11 8 2 5 1 0 Coniferous high mountainous forest zone 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 1 1 Cosmopolitan 2 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0

cial period found refuge in the Near East, and Both species are introduced. The origin and the later, following the retreat of the glaciers, spread colonization pattern of these species are hard to along the Mediterranean coast and into the South- define. Thus, for a long time it was assumed that ern and the Central Europe. It is among dominant the origin of Monomorium pharaonis was Africa, but zoogeographical elements in our investigated area later studies suggested that this species probably (14 spp., 16.09%). originated in tropical Asia (Wetterer 2010). Part of the present ant fauna of Central Europe Twenty six species (29.88%) is associated with originates from the Balkan region. These species the Mediterranean habitats, and almost two thirds survived glacial periods (Würm I-III) in the ref- of species (56 spp., 64.37%) are associated with uges of the Balkan Peninsula (as described by mixed and deciduous forests (Fig. 2). Unexpected Savić 2008), where, due to geographical isolation is higher percentage of the non-Mediterranean and low capacity for long distance dispersal, iso- element in the myrmecofauna of the explored re- lated populations underwent speciation, and after gion. The coastal mountains, Lovćen, Orjen and the retreat of the glaciers, migrated northward as Rumija, rise almost immediately above the coastal distinct but closely-related species. They form one line, leaving very narrow zone with the typical of the most important element (the Central and Mediterranean climate. With the altitude of over Southern European elements) in investigated area 1000 meters, the climate on these mountains is (14 spp., 16.09%). modified under the influence of the continental The cosmopolitan ants’ element is represented and the alpine climate (Ćurčić et al. 2008) becom- by Monomorium pharaonis and Pheidole pallidula. ing colder and more rainy.

32 Karaman, M.G.

Table 3. Statistical parameters and significance levels (Spearman Rank Order Correlations) for the relations between total species numbers and species numbers of zoogeographical zones and ele- ments in relation to altitude steps (n = 19).

Spearman R t(N-2) p-value p-level All species -0.812146 -5.73921 0.000024 <0.001 Zoogeographical zones Mixed and deciduous forest zone -0.698595 -4.02559 0.000877 <0.001 Mediterranean zone -0.917890 -9.53689 0.000000 <0.001 Coniferous high mountainous forest zone 0.676475 3.78725 0.001471 <0.01 Cosmopolitan -0.870572 -7.29503 0.000001 <0.001 Zoogeographical elements Holarctic -0.907570 -8.9117 0.000000 <0.001 Transpalaearctic (south) 0.034011 0.1403 0.890061 n.s. West Palaearctic -0.837754 -6.3257 0.000008 <0.001 Euro-Caucasian -0.676810 -3.7907 0.001460 <0.01 Euro-Siberian -0.165109 -0.6902 0.499362 <0.05 European -0.536543 -2.6215 0.017866 <0.05 Central and Southern-European -0.580019 -2.9358 0.009235 <0.01 Southeast-European -0.707107 -4.1231 0.000710 <0.001 Southern-European -0.380875 -1.6984 0.107658 n.s. Holomediterranean -0.617588 -3.2376 0.004839 <0.01 North-Mediterranean -0.935192 -10.8880 0.000000 <0.001 Balkan -0.821390 -5.9377 0.000016 <0.001 East-Adriatic -0.676113 -3.7835 0.001483 <0.01 Endemics of Montenegro -0.584765 -2.9722 0.008546 <0.01 Boreo Montane 0.676475 3.7873 0.001471 <0.01 Cosmopolitan -0.870557 -7.2950 0.000001 <0.001

Figure 5. Pattern of total number of species in relation to Figure 6. Patterns of species numbers of zoogeographical altitude steps. zones in relation to altitude steps.

Vertical distribution of species a.s.l., corresponds to the transition zone between The largest number of species is located at the lowland and mountain ant communities. Most of transition zone between two, species rich commu- species from this transition zone belong to the nities (Lomolino 2001). In the studied area this zone of mixed and deciduous forests. The Euro- transition zone is between the Mediterranean and Caucasian species mainly occur below 1000 m alti- the Mixed and deciduous forests zones, within the tude, but Aphaenogaster obsidiana was found over altitude of 100-300 m a.s.l.. Then the total number 900 m a.s.l. only. On the other hand, the Euro- of species decreases with the altitude. However, Siberian elements do not show a significant nega- this decrease has a major intermediate peak be- tive correlation with altitude: Myrmica sabuleti and tween 700 and 1100 m a.s.l. and two minor peaks Lasius platythorax can be found from the sea level (Fig. 5). The major peak is between 700 and 1100 m to the 1400 m a.s.l.; Formica rufibarbis and F.

Zoogeography and distribution of ants (Hymenoptera: Formicidae) in Southern Montenegro 33 polyctena occur over 1100 m altitude, while Campo- and 1100 m a.s.l. These species are well adapted to notus vagus was found from the 0 to 1100 m a.s.l. the harsh conditions of higher altitudes in the Di- The species number in the Mediterranean zone narides. The species richness of the high moun- shows significant negative correlation with alti- tainous coniferous forest is negligible (3 spp., tude. At 500 m above the sea level, the number of 3.45%), because due influence of the Mediterra- species is negligible, depending on the geographi- nean climate that reaches all to the highest peaks cal position and the north-to-south exposure of the of the coastal mountains, the high mountainous locality. The species still sporadically nest above coniferous forests are only sporadically present in 1000 m altitude on the sites with the south expo- the investigated region. sure. Thus Temnothorax rottenbergi was found at

1183 m a.s.l. at the rocky summit of Vrsuta (Ru- mija Mt.), while Lasius lasioides, Solenop- sis Acknowledgements. Many thanks to Barsi Laszlo (De- banyulensis and Temnothorax sordidulus were found partment of Biology, Novi Sad, Serbia) for help in retriev- at 1260-1380 m a.s.l. in the rocky fields on the ing statistical data; to Vesna Karaman-Castro for a lin- Lovćen Mt. guistic improvements; to anonymous reviewers for help- Unlike the other zones, the zone of coniferous ful comments on an earlier version of the manuscript. high mountainous forests exhibited positive corre- lation between species number and altitude. Cam- ponotus ligniperda was found nesting only on Orjen References at 1100 m altitude; Leptothorax acervorum on Lov- Atannasov, N., Dlusskij, G.M. (1992): Fauna Bulgarica ćen at 1400 m a.s.l.; and Manica rubida on Orjen at (Hymenoptera, Formicidae). Aebidus Academiae Scientiarum 1600-1900 m. a.s.l. We did not find representati- Bulgaricae 22: 1-311. ves from this zone at 1200-1300 and at 1500 m Baroni Urbani, C., Collingwood, C.A. (1977): The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe. Acta a.s.l., but further field studies are needed to con- Zoologica Fennica 152: 1-34. firm our finding. Bolton, B., Alpert, G., Ward, P.S., Naskrecki, P. (2006): Bolton's The number of ant species asociated with the Catalogue of Ants of the World: 1758-2005. Harvard University Press, Cambridge, Massachusetts, London. Database available zone of coniferous high mountainous forests is on CD-ROM. very small because of the lack of the true high Bračko, G. (2000): Review of the Ant Fauna (Hymenoptera: mountainous coniferous forests. In our investigat- Formicidae) of Slovenia. Acta Biologica Slovenica 43(4): 37-54. Bračko, G. (2006): Review of the ant fauna (Hymenoptera: ed area this zone can be found only on Orjen Mt. Formicidae) of Croatia. Acta Entomologica Slovenica 14(2): 131- whose highest peak is at 1894 m a.s.l., where the 156. high mountainous coniferous forests are located. Bračko, G. (2007): Present status of the Slovenian ant fauna (Hymenoptera: Formicidae). Myrmecological News 10: 102. Brown, W. L. (1973): A comparison of the Hylean and Congo-West african rain forest ant faunas. pp. 161-185. In Meggers, B.J., Conclusion Ayensu, E.S. and Duckworth, W.D. (eds.) Tropical forest ecosystems in Africa and South America: a comparative review. Smithsonian Institution press, Washington. The myrmecofauna of investigated region belong- Czechowski, W., Radchenko, A., Czechowska, W. (2002): The ants ing to three main zoogeographical zones corre- (Hymenoptera, Formicidae) of Poland. Muzeum i Instytut Zoologii PAN, Warszawa 200pp. sponding with three main vegetation zones. The Ćurčić, B., Decu, V., Juberthie, C. (2008): Cave dwelling most number of species belong to: Central and invertebrates in Montenegro. pp. 35-55. In: Makarov, Southern European (14 spp., 16.09%), Euro-Cauca- Dimitrijević (eds), Advances in Arachnology and Developmental Biology. Papers dedicated to Prof. Dr. Božidar sian (14 spp., 16.09%) and North-Mediterranean Ćurčić. Inst.Zool., Belgrade; BAS, Sofia, Fac. Life sci., Vienna; (11 spp., 12.64%) element. SASA, Belgrade & UNESCO MAB Committee, Serbia. Vienna- The total number of species is inversely pro- Belgrade-Sofia, Monographs, 12. Glaser, F. (2006): Biogegraphy, diversity and vertical distribution of portional to the altitude. Most of the species prefer ants (Hymenoptera: Formicidae) in Vorarlberg, Austria. warmer types of habitats. The most numerous Myrmecologische Nachrichten 8: 263-270. species from the Mediterranean zone are distri- Hijmans, R.J., Guarino, L. Bussink, C. Mathur, P. Cruz, M. buted up to 500 m a.s.l., the Mixed and deciduous Barrentes, I., Rojas, E. (2004): DIVA-GIS Vsn. 5.0. A geographic information system for the analysis of species distribution data. forest elements up to 1600 m altitude, and the high Manual available at mountainous coniferous forest elements above Karaman, M., Karaman, G., Petrov, I. [1998 (1993)]: Contribution to the knowledge of the ants (Hymenoptera, Formicidae) of the 1600 m a.s.l.; Vrmac peninsula - Boka Kotorska (Montenegro). Bulletin of the The species richness of the Mixed and deci- Republic Institution for the Protection of Nature and the duous forest zone has the mid peak between 700 Museum of Natural History in Podgorica 26: 41-53.

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