Divers ity an d spec ies compos ition of ants in arid and semi-arid regions of Iran

DISSERTATION for the degree of Dr. rer. nat. in the Faculty of Natural Science of the University of Ulm submitted by

Omid Paknia from Tehran

Ulm, 2010

Diversity and species

composition of ants in arid and semi-arid regions of Iran

DISSERTATION zur Erlangung des Doktorgrades Dr. rer. nat. der Fakultät für Naturwissenschaften der Universität Ulm vorgelegt von

Omid Paknia aus Teheran

Ulm 2010

AMTIERENDER DEKAN Prof. Dr. Axel Groβ ERSTGUTACHTER Prof. Dr. Elisabeth K. V. Kalko ZWEITGUTACHTER PD Dr. Martin Pfeiffer Tag der mündlichen Prüfung: 17.12.2010

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To mom, dad, Elham, Elham, Arman and Ashwin

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Contents

1 General introduction 1

Drylands 1 Biodiversity in drylands 1 Objectives of this study 4 Contributions of co-authors 10 References 11 2 Desert versus steppe: Multi-scale spatial patterns in diversity of ant communities in Iran 15

Abstract 15 Introduction 16 Methods 18 Results 22 Discussion 27 Acknowledgments 32 References 33 Appendix S1 36

3 Hierarchical partitioning of ant diversity: Implications for conservation of biogeographical diversity in arid and semi-arid areas 39 Abstract 39 Introduction 40 Methods 42 Results 47 Discussion 50 Acknowledgments 55 References 56 Appendix S1 59

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Appendix S2 61

4 Environmental and spatial drivers of ant species composition in a dryland ecosystem in Iran 63

Abstract 63 Introduction 64 Methods 66 Results 70 Discussion 75 Conclusion 78 Acknowledgments 79 References 79 Appendix S1 82

5 Productivity, solely, does not explain species richness of ants in a dryland habitat 85 Abstract 85 Introduction 86 Methods 88 Results 93 Discussion 98 Acknowledgments 101 References 101

6 A preliminary checklist of ants (Hymenoptera: Formicidae) of Iran 105

Abstract 105 Introduction 105 Material and Methods 105 Results 106 Discussion 110 Acknowledgments 111

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References 111 7 New records of ants (Hymenoptera: Formicidae) from Iran 114

Abstract 114 Introduction 114 Material and Methods 114 Results 116 Discussion 121 Acknowledgments 121 References 121

8 Two new species of genus Cataglyphis Foerster, 1850 (Hymenoptera: Formicidae) from Iran 124 Abstract 124 Introduction 124 Material and Methods 124 Taxonomy 125 Acknowledgments 130 References 130

9 Synthesis 132

References 136

Summary 138 English 138 German 141 Persian 144 General appendix 147 Acknowledgments 150 Curriculum Vitae 153 Declaration 157

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1 General introduction

DRYLANDS

Dryland ecosystems are defined by their arid conditions. As a consequence of this aridity, most drylands are unproductive and monotonous, leading many people to view them as wastelands. However, from the biologist’s view, drylands are laboratories of nature, where natural selection is exposed at its extremes (Ward, 2009). Over the last two million years climate changes of the earth have forced the drylands to minimize during the cold glacial periods and expand during the hot interglacials, leading finally to the current warming and aridization trend from mid-Holocene to date. Today, arid and semi-arid regions cover about 47% of the world’s land surface, when dry sub-humid regions are also included (United Nations Environment Program, 1992). By classical categorization arid lands are divided into three classes: Extreme arid, with less than 60-100 mm mean annual precipitation; Arid, with precipitation from 60-100 mm to 150-250 mm; Semi-arid, from 150-250 mm to 250-500 mm annual precipitation (Noy-Meir, 1973). Arid lands are distributed in all continents, excluding Antarctica; however, the Palearctic realm contains the largest sets of deserts in the world, a total 63 per cent of all deserts on the planet.

Biodiversity of drylands

It has been well demonstrated that species diversity is typically positively correlated with net primary productivity at large spatial scales (Waide et al., Chapter 1 - General Introduction

1999). On the basis of this fact, it is assumed that deserts are inhabited by relatively few species. In addition, not all plant and animal species have acquired the physiological capabilities to endure the stresses exerted by the high temperature fluctuations and scarcity of water found in dryland (Shmida et al., 1986). Despite of these logic assumptions for low species diversity in drylands, scientific studies show that species diversity of many desert animals such as ants, beetles and lizards and annual plants is quite high (Inouye, 1991; Polis, 1991). The main explanation for this apparent paradox is that, although primary productivity is low in drylands, these ecosystems tend to have higher ecological efficiencies of energy transfer along food changes (Kaspari et al., 2000). Additionally, it is suggested that detritivory in deserts is based on macro- detritivores such as termites and ants instead of bacteria (Shachak et al., 2005). As a result, macro-detritivores take the place of herbivores as the major group of primary consumers and this replacement has important implications for energy flow and species diversity in dryland ecosystems. The desert food webs are also ectothermic-based food webs, because the main primary and secondary consumers are ectotherm. Ectothermic-based food webs are energetically more efficient than endothermic-based food chains where the mammals and birds are primary and secondary consumers (Humphreys, 1979). Notably, certain taxa are more diverse in drylands compared to other ecosystems. These include predatory arthropods such as scorpions, tenebrionid beetles, ants and termites, snakes and lizards, and annual plants (Crawford, 1981).

As one of most successful taxa after their arising in the mid-Cretaceous about 120 million years ago, ants comprise approximately 20000 species (Ward, 2003). They have colonized all arid land biomes of the earth and are a dominant component of invertebrate assemblages in deserts and grasslands around the world (Hölldobler & Wilson, 1990). They are also functionally important elements in these ecosystems throughout the world (Bestelmeyer & Schooley, 1999). The seed-harvester ants of drylands alone can number from several hundred thousand to several million individuals per hectare and can have a total biomass comparable to that of small vertebrates (Went et al., 1972). This large number of seed-harvester ants can harvest up to several hundred thousand seeds per hectare per year and have a significant impact on energy flow. harvester ants

2 Chapter 1 - General Introduction are well known “ecosystem-engineers” that affect the relative abundance and species composition of annual plants and thus shape the landscape of their habitats (Johnson, 2001). Nevertheless, the ecological role of ants in arid regions is not restricted to seed-harvester ants. In arid regions, ants show strong interaction with many organisms and ecosystem processes (Bestelmeyer, 2005; Whitford et al., 2008). They also contribute to heterogeneity or patchiness of landscapes by producing nest mounds which increases the topographic complexity of the soil surface, plant diversity and density (Wilby et al., 2005). Ants, beside other burrowing desert animals, can influence soil structure and hydrology (Jones et al., 1993).

In spite of their large and fundamental contributions in arid and semi-arid regions, ants have remained poorly known in many arid and semi-arid regions of the Palearctic realm (Dunn et al., 2010: Fig. 3.2.1). The Palearctic arid and semi- arid regions cover a remarkable 16 million square kilometers, stretching from the Atlantic coasts of Morocco, to the Gobi in the Central Asia, and including 21 lowland desert ecoregions (Olson et al., 2001). In addition, these regions include both types of hot and cold deserts. Scarce faunistic and taxonomic studies on ants, which were carried out during early decades of the 20 century in these regions, were not followed by constructive and comprehensive modern faunistic studies, except for the Arabian Peninsula (Collingwood, 1985; Collingwood & Agosti, 1996), Turkmenistan (Dlussky et al., 1990), Kyrgyzstan (Schultz et al., 2006) and Mongolia (Pfeiffer et al., 2006). Fauna and taxonomy of a few taxa such as Cataglyphis and Camponotus have been also subject of modern revisions and studies (Agosti, 1990; Radchenko, 1996, 1997). Nevertheless, there is not even a rough number of ant species from these vast regions, contrary to other arid regions such as North America, Australia and Southern Africa which have been studied extensively (Andersen, 2007).

As a logical consequence, it is not surprising that the ecology of this taxon, in all its aspects, has been poorly investigated in the Palearctic dry regions. Specifically, so far, there has been only one comprehensive study on the ant community ecology conducted by Pfeiffer et al. (2003) in Mongolia. In that study, Pfeiffer and colleagues found a poor diversity of ants, a total of 26 species, in the Gobi desert and adjacent steppe regions. Although species

3 Chapter 1 - General Introduction composition of ants differed between the Gobi desert and adjacent steppe, ant species richness did not show any clear cut and significant pattern along the latitudinal gradient or along any climate parameter gradient such as precipitation or temperature, suggesting that ant species richness and composition in these harsh and cold arid ecosystems probably do not follow the observed patterns in other arid regions of the world (e.g. Davidson, 1977). Their findings finally urge to conduct more researches in cold arid ecosystems for a better understanding of the fundamental differences between cold and hot desert ecosystems and the differing impact of environmental parameters on niche pattern of ants in both systems (Pfeiffer et al., 2003). This thesis aims at adding comprehensive knowledge on ant diversity and assemblage organization in arid and semi-arid regions of Iran as a case study in dryland regions of Palearctic realm.

OBJECTIVE OF THIS STUDY

The general objective of this thesis was to broaden the understanding of the impact of environmental and spatial factors on species richness, diversity and assemblage organization of ants in arid and semi-arid regions in Iran. An extensive literature suggests that species richness and species composition of biota is strongly influenced by contemporary climate factors or productivity especially at large geographical scales (Pianka, 1966; Currie, 1991; Hawkins et al., 2003; Qian, 2010). Nevertheless, the magnitude of this influence varies among different taxa, ecosystems and even different spatial extent or grains (Whittaker et al., 2001; Field et al., 2009). Furthermore, the importance of spatial ecological structure in the modern thinking of ecology has been widely recognized by ecologists (Legendre & Legendre, 1998; Fortin & Dale, 2005). In the ecological context, spatial structure of any ecological pattern should be considered, not only as a potential nuisance for sampling or statistical analyses, but also as functional requirement (Borcard & Legendre, 2002).

Iran is located in south western Asia, at 25°-39° latitude. The Iranian dry lands form a part of the Plateau of Iran, an extensive highland region of the western Asia. The country comprises three distinct phytogeographical regions including

4 Chapter 1 - General Introduction the Hyrcanian, the Irano-Turanian and the Nubo-Sindian (Zohary, 1973; Zehzad et al., 2002). The Irano-Turanian is the largest phytogeographical region, covering 70 per cent of the country and the Hyrcanian is the smallest region covering only 4 per cent of the country at the southern coasts of the Caspian Sea. The remaining 26 per cent of the land’s country in northern coasts of Persian Gulf comprises the Nubo-Sindian phytogeographical region. As a part of a global conservation approach the country recently has been divided to fourteen smaller biogeographical units as “ecoregions” by Olson et al. (2001). The researches that are documented in this thesis were conducted in two of the Irano-Turanian and the Nubo-Sindian phytogeographical regions, comprising four main ecoregions: the Elburz Range forest steppe, the Central Persian desert basins, the Zagros Mountains forest steppe and the southern Iranian Nubo-Sindian deserts. All field work and ant data collection for this thesis was carried out during two intensive sampling seasons in 2007 and 2008 in fourteen national parks or protected areas (Figure 1). In the first sampling year, I collected ants in eight sampling sites along a long latitudinal gradient from Elburz forest steppe to Nubo-Sindian deserts (Figure 1). In the second sampling season, I collected ants in eight sampling sites within the Central Persian desert ecoregion. In both seasons, ants were collected by the pitfall traps.

As the main body of this dissertation, I present four ecological studies on ant communities in arid and semi-arid regions of Iran in chapters two through five:

In chapter 2, I investigate the multi-scale spatial patterns in ant diversity across a long latitudinal gradient. Multi-scale approaches are crucial in diversity and distributional analyses, because spatial patterns and processes which operate in communities may differ depending on the scales used (Noda, 2004). The main question of this study is whether spatial pattern of ant diversity differ among steppe and desert biomes of Iran. Applying a hierarchical approach, I demonstrate variations in species richness, diversity and equitability of ants, for the whole region and for two main biomes, steppe and desert, across fine-to- broad spatial grains. In addition, I demonstrate and compare the patterns of alpha and beta diversity among these major biomes. The main outline of this chapter is that 1) both, biome and ecoregional grains, contribute largely to overall species richness and diversity (high beta diversity in these grains) and 2) the ant

5 Chapter 1 - General Introduction diversity is higher in the steppe biome than the desert biome only at the largest grain.

Chapter 3 deals with the conservation biogeography of ants in arid regions of Iran. Because ants have been used widely as a “surrogate” of biodiversity (Andersen, 1997; Andersen & Majer, 2004; Bennett et al., 2009), our view can be extended to other taxa when applying ants as a “surrogate” taxon for other taxa.

40° N

Caspian Sea 0150 300 Km 38° N 1 2 36° N 3 4

5 Iran 34° N 6 11 7 10 9 8 32° N 12 30° N

13 Central Persian desert basins 14 28° N Elburz Range forest steppe South Iran Nubo-Sindian desert 26° N Zagros Mountains forest steppe Persian Gulf

44° E 46° E 48° E 50° E 52° E 54° E 56° E 58° E 60° E 62° E

Figure 1 Map of Iran, documenting four main ecoregions that were sampled in two sampling years: red circles indicate sample sites of 2007 and blue triangles indicate sample sites of 2008.

My focal question for this research is whether the World Wildlife Foundation’s ecoregional map which has been broadly applied for conservation purposes is the appropriate scale for conserving ants in arid regions of Iran (Olson et al., 2001). Although this map has been successfully used for other taxa: birds, mammals and trees (e.g. Mcdonald et al., 2005), this study is the first attempt to evaluate it for ant conservation. In addition, I investigate whether each ecoregion represent a

6 Chapter 1 - General Introduction distinct ant community composition. Hierarchical partitioning of diversity is also applied for this research, but across different levels. Hierarchical cluster analysis is used to assess the similarity of ant species composition among ecoregions. Results of this study suggest that ecoregions represent the proper scale for conserving ant diversity. However, ecoregions differ in their hierarchical partitioning patterns; as a result, different strategies should be considered for different ecoregions.

Chapter 4 explores the impact of climate factors and spatial structure on ant species composition along the latitudinal gradient. The central question of this study is how much of variation in ant species composition is explained by contemporary climate factors, such as precipitation and temperature range and which spatial structure of ant communities exists in our study region. Recent studies have demonstrated that environmental and spatial factors do not have large impacts on ant communities in tropical forests (Mezger & Pfeiffer, 2010; Vasconcelos et al., 2010). But very little is known about the impact of environment factors and spatial structures on ants outside of tropical regions. Applying variation partitioning by Correspondence Canonical Analysis (CCA), and spatial analysis, I disentagle the variation in ant species composition of different plots into climate and spatial components. The outcome of this research is that ant community composition is influenced largely by environmental and spatial factors.

In chapter 5, I report the first comprehensive study on ant species richness across the latitudinal and environmental gradients in the Central Persian desert basins. For animals, deserts and semi-deserts are productivity-controlled ecosystems. But studies which have been conducted in dry lands mostly have not revealed a positive correlation between ant species richness and productivity or its surrogates (e.g. rainfall) (e.g. Morton & Davidson, 1988; Delsinne et al., 2010). In this study, I evaluate the hypothesis that the productivity – species richness model is the best model to explain the ant species richness gradient in the Central Persian desert. I apply multimodel inference to assess support for several competing models which are thought to be responsible for the observed geographical variation of species richness of ants. My results suggest that the productivity model is the best model. However, a strong support for the

7 Chapter 1 - General Introduction tolerance – productivity model, which consists of temperature range and productivity factors, also was observed; indicating that species richness is well explained only by taking this model into account. This model predicts that ant species richness is determined by a tradeoff between impact of productivity and temperature range in the semi-cold Central Persian desert.

Because the ant fauna of the Iran is poorly known, an intensive study of ant taxonomy was crucial for success of the community ecology investigations presented here, which lead to three additional chapters on taxonomy and faunistic of Iranian ants for my thesis, which are already published.

In chapter 6, I report the first species list for ants of Iran, issued in Myrmecological News. Although the first records on the Iranian ants were published more than 100 years ago (e.g. Forel, 1904a, b), the ant fauna of this country remains poorly known. Preparing this species list was an urgent task because during the last decade many ant records were published by Iranian myrmecologists in local journals or national conferences, which needed to be sorted and evaluated. In addition, there have been old records from the country that needed to be revised. As a result, this species list can be seen as the first attempt to compiling and revising the ant records from Iran, which will be a helpful reference for myrmecologists that are interested in Asian ants.

Chapter 7 records 32 species and six genera new to Iran after the publication of the basis data (chapter 6). These records are on basis of 35000 specimens which I had collected in two sampling years in 2007 and 2008. Ants were sorted and identified in the first round in Ulm University. Further identifications were carried out the Museum and Institute of Zoology, Polish Academy of Sciences, in Warsaw, together with Prof. Alexander Radchenko (National Academy of Sciences of Ukraine).

In chapter 8, I represent, as co-author, the description of two new ant species to science. Both species belong to the genus Cataglyphis, Cataglyphis stigmatus sp. nov. and C. pubescens sp. nov. These two species have been collected in my 2007 and 2008 field works, from Nubo-Sindian and Central Persian desert ecoregions, respectively. C. stigmatus differs from known species of the bicolor

8 Chapter 1 - General Introduction group by its yellow colour. C. pubescens differs by the dense and long depressed pubescence on the head and alitrunk.

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Publications of the thesis and contributions of co-authors At the date of this thesis delivery, chapters 2 and 3 of this dissertation have been accepted or have been under revision for publishing in scientific journals. Chapters 5 and 6 have been prepared for submission. Chapters 6 to 8 have already been published in scientific journals. All chapters are closed entities that can be read independently of each other. However, this structure results in some redundancy in content among the chapters.

• Chapter 2: Paknia O. and Pfeiffer M. (revised) Desert versus steppe: Multi-scale spatial patterns in diversity of ant communities in Iran. Insect Conservation and Diversity. I am the first author of this paper. I am responsible for the study design, field work, data collection, data analyses and writing. PD Dr. Pfeiffer is the co- author as he provided important advice and comments on all aspects of the study and the manuscript. This combination of authors and contributions is the same for three next chapters (3-5) of this dissertation.

• Chapter 3: Paknia O. and Pfeiffer M. (accepted) Hierarchical partitioning of ant diversity: implications for conservation of biogeographical diversity in arid and semi-arid areas. Diversity and Distributions.

• Chapter 4: Paknia O. and Pfeiffer M. (in preparation) Environmental and spatial drivers of ant species composition in a dryland ecosystem in Iran.

• Chapter 5: Paknia O. and Pfeiffer M. (in preparation) Productivity, solely, does not explain diversity in a dryland habitat.

• Chapter 6: Paknia O., Radchenko A., Alipanah H. and Pfeiffer M. (2008). A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News: 11, 151-159.

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I am the first author of this paper. I am responsible for the study design and providing data from Iranian museums and for literature searching and collecting information. I am responsible for writing the manuscript. Prof. Radchenko provided data from European natural history museums, re- identified doubtful species, read the manuscript and gave important input on the manuscript. Dr. Alipanah, provided data from her collection, read the manuscript and gave valuable comments on it. PD Dr. Pfeiffer read the manuscript and gave valuable advice on all aspects of the study.

• Chapter 7: Paknia O., Radchenko A. and Pfeiffer M. (2010). New records of ants (Hymenoptera: Formicidae) from Iran. Asian Myrmecology: 3, 29 - 38. I am first author of this paper. I have designed the study and wrote the manuscript. I am also responsible for the first round of species identification. The final identification process was carried out in the Museum and Institute of Zoology of Polish Academy of Sciences in Warsaw, under supervision of Prof. Radchenko. He also provided valuable comments on the manuscript. PD Dr. Pfeiffer provided advice on all aspects of the study.

• Chapter 8: Radchenko A. Paknia O. (2010). Two new species of the genus Cataglyphis Foerster (Hymenoptera, Formicidae) from Iran. Annales Zoologici: 60, 69 - 76. In this paper, I am the second author. I provided basal data set for these two new species. I read the manuscript and provided comments on it. Prof. Radchenko is the first author as he is responsible for the study design, describing the new species and writing the paper.

REFERENCES:

Agosti, D. (1990) Review and reclassification of Cataglyphis (Hymenoptera, Formicidae). Journal of Natural History, 24, 1457-1505

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Andersen, A.N. (1997) Using ants as bioindicators: multiscale issues in ant community ecology. Conservation Ecology, 1, 8. Available from the Internet. URL: http://www.consecol.org/vol1/iss1/art8/ Andersen, A.N. (2007) Ant diversity in arid Australia: a systematic overview. Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80. (ed. by R.R. Snelling, B.L. Fisher and P.S. Ward), pp. 91-51. Andersen, A.N. & Majer, J.D. (2004) Ants show the way Down Under: invertebrates as bioindicators in land management. Front Ecological Environment 2, 291- 298 Bennett, J., Kutt, A., Johnson, C. & Robson, S. (2009) Ants as indicators for vertebrate fauna at a local scale: an assessment of cross-taxa surrogacy in a disturbed matrix. Biodiversity and Conservation Bestelmeyer, B.T. (2005) Does desertification diminish biodiversity? Enhancement of ant diversity by shrub invasion in southwestern USA. Diversity and Distributions, 11, 45-55 Bestelmeyer, B.T. & Schooley, R.L. (1999) The ants of the southern Sonoran desert: community structure and the role of trees. Biodiversity and Conservation, 8, 643-657 Borcard, D. & Legendre, P. (2002) All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices. Ecological Modelling, 153, 51-68 Collingwood, C.A. (1985) Hymenoptera: Fam. Formicidae of Saudi Arabia. Fauna Saudi Arabia, 7, 230-302 Collingwood, C.A. & Agosti, D. (1996) Formicidae (Insecta: Hymenoptera) of Saudi Arabia (Part 2). Fauna Saudi Arabia, 15, 300-385 Crawford, C.S. (1981) Biology of desert invertebrates Springer Verlag, Berlin. Currie, D.J. (1991) Energy and large-scale patterns of animal- and plant-species richness. The American Naturalist, 137, 27 Davidson, D.W. (1977) Species diversity and community organization in desert seed- eating ants. Ecology, 58, 711-724 Delsinne, T., Roisin, Y., Herbauts, J. & Leponce, M. (2010) Ant diversity along a wide rainfall gradient in the Paraguayan dry Chaco. Journal of Arid Environments, 74, 1149-1155 Dlussky, G.M., Soyunov, O.S. & Zabelin, S.I. (1990) Ants of Turkmenistan. Ylym Press, Ashkhabad. Dunn, R.R., Guenard, B., Weiser, M.D. & Sanders, N.J. (2010) Geographical gradients. Ant Ecology (ed. by L. Lach, C.L. Parr and K.L. Abbot). Oxford University Press, Oxford. Field, R., Hawkins, B.A., Cornell, H.V., Currie, D.J., Diniz-Filho, J.A., Guégan, J., Kaufman, D.M., Kerr, J.T., Mittelbach, G.G., Oberdorff, T., O'brien, E.M. & Turner, J.R.G. (2009) Spatial species-richness gradients across scales: a meta- analysis. Journal of Biogeography, 36, 132-147 Forel, A. (1904a) Dimorphisme du mâle chez les fourmis et quelques autres notices myrmécologiques. Annales de la Société Entomologique de Belgique, 48, 421- 425 Forel, A. (1904b) Note sur les fourmis du Musée Zoologique de l'Académie Impériale des Sciences à St. Pétersbourg. Ezheg. Zool. Muz., 8, 368-388 Fortin, M.J. & Dale, M.R.T. (2005) Spatial Analysis: A Guide for Ecologists. Cambridge University Press, Cambridge, UK.

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Hawkins, B.A., Field, R., Cornell, H.V., Currie, D.J., Guegan, J.-F., Kaufman, D.M., Kerr, J.T., Mittelbach, G.G., Oberdorff, T., O'brien, E.M., Porter, E.E. & Turner, J.R.G. (2003) Energy, water, and broad-scale geographic patterns of species richness. Ecology, 84, 3105-3117 Hölldobler, B. & Wilson, E.O. (1990) The ants. Belknap Press of Harvard University Press, Cambridge, Massachusetts, USA. Humphreys, W. (1979) Production and respiration in animal populations. Journal of Animal Ecology, 48, 427-453 Inouye, R. (1991) Population biology of desert annual plants. The Ecology of Desert Communities (ed. by G.A. Polis), pp. 27-54. The University of Arizona Press, Tucson. Johnson, R.A. (2001) Biogeography and community structure of North American seed-harvester ants. Annual Review of Entomology, 46, 1-29 Jones, C.G., Lawton, J.H. & Shachak, M. (1993) Organisms as ecosystem engineers. Oikos, 69 Kaspari, M., O'donnell, S. & Kercher, J.R. (2000) Energy, density, and constraints to species richness: Ant assemblages along a productivity gradient. American Naturalist, 155, 280-293 Legendre, P. & Legendre, L. (1998) Numerical Ecology. Elsevier, Amsterdam. Mcdonald, R., Mcknight, M., Weiss, D., Selig, E., O'connor, M., Violin, C. & Moody, A. (2005) Species compositional similarity and ecoregions: Do ecoregion boundaries represent zones of high species turnover? Biological Conservation, 126, 24-40 Mezger, D. & Pfeiffer, M. (2010) Partitioning the impact of abiotic factors and spatial patterns on species richness and community structure of ground ant assemblages in four Bornean rainforests. Ecography, doi: 10.1111/j.1600- 0587.2010.06518.x Morton, S.R. & Davidson, D.W. (1988) Comparative structure of harvester ant communities in arid Australia and North America. Ecological Monographs, 58, 19-38 Noda, T. (2004) Spatial hierarchical approach in community ecology: a way beyond high context-dependency and low predictability in local phenomena. Population Ecology, 46, 105-117 Noy-Meir, I. (1973) Desert ecosystems: Environment and producers. Annual Review of Ecology and Systematics, 4, 25-51 Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D'amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. Bioscience, 51, 933-938 Pfeiffer, M., Chimedregzen, L. & Ulykpan, K. (2003) Community organization and species richness of ants (Hymenoptera: Formicidae) in Mongolia along an ecological gradient from steppe to Gobi desert. Journal of Biogeography, 30, 1921-1935 Pfeiffer, M., Schultz, R., Radchenko, A., Yamane, S., Woyciechowski, M., Ulykpan, A. & Seifert, B. (2006) A critical checklist of the ants of Mongolia (Hymenoptera: Formicidae). Bonner Zoologische Beitraege, 55, 1-8 Pianka, E.R. (1966) Latitudinal gradients in species diversity: A review of concepts. American Naturalist, 100, 33-46

13 Chapter 1 - General Introduction

Polis, G.A. (1991) Desert communities: an overview of patterns and processses. The Ecology of Desert Communities (ed. by G.A. Polis), pp. 1-26. The University of Arizona Press, Tucson. Qian, H. (2010) Environment–richness relationships for mammals, birds, reptiles, and amphibians at global and regional scales. Ecological Research, 25, 629-637 Radchenko, A.G. (1996) A key to the ant genus Camponotus (Hymenoptera, Formicidae) in Palearctic Asia. [In Russian.]. Zoologicheskii Zhurnal, 75, 1195-1203 Radchenko, A.G. (1997) A review of ants of the genus Cataglyphis Foerster (Hymenoptera, Formicidae) from Asia. Entomological Review (Washington), 77, 684-698 Schultz, R., Radchenko, A.G. & Seifert, B. (2006) A critical checklist of ants of Kyrgyzstan (Hymenoptera: Formicidae). Myrmecologische Nachrichten, 8, 201-207 Shachak, M., Gosz, J.R., Steward, T.A.P. & Perevolotsky, A. (2005) Biodiversity in drylands toward a unified framework. Oxford University Press, New York. Shmida, A., Evenari, M. & Noy-Meir, I. (1986) Hot desert ecosystems an integrated view. Hot deserts and Arid shrublands (ed. by M. Evenari, A. Shmida and I. Noy-Meir), pp. 379-387. Elsevier Science Publications, Amsterdam. United Nations Environment Program (1992) Convention on Biological Diversity. United Nations, New York, NY. Vasconcelos, H.L., Vilhena, J.M.S., Facure, K.G. & Albernaz, A.L.K.M. (2010) Patterns of ant species diversity and turnover across 2000 km of Amazonian floodplain forest. Journal of Biogeography, 37, 432-440 Waide, R.B., Willing, M.R., Steiner, C.F., Mittelbach, G., Gough, L., Dodson, S.I., Juday, G.P. & Parmenter, R. (1999) The relationship between productivity and species richness. Annual Review of Ecology and Systematics, 30, 257-300 Ward, D. (2009) The biology of deserts. Oxford University Press, Oxford; New York. Ward, P.S. (2003) Ants. Current Biology, 16, R152-R155 Went, F.W., Wheeler, J. & Wheeler, G.C. (1972) Feeding and digestion in some ants (Veromessor and Manica). Bioscience, 22, 82-88 Whitford, W.G., Barness, G. & Steinberger, Y. (2008) Effects of three species of Chihuahuan Desert ants on annual plants and soil properties. Journal of Arid Environments, 72, 392-400 Whittaker, R.J., Willis, K.J. & Field, R. (2001) Scale and species richness: Towards a general, hierarchical theory of species diversity. Journal of Biogeography, 28, 453-470 Wilby, A., Boeken, B. & Shachak, M. (2005) The impact of animals on species diversity in arid-land plant communities. Biodiversity in Drylands: Toward a unified Framework (ed. by M. Shachak, J.R. Gosz, T.A.P. Steward and A. Perevolotsky), pp. 189-205. Oxford University Press, New York. Zehzad, B., Kiabi, B.H. & Madjnoonian, H. (2002) The natural areas and landscape of Iran: An overview. Zoology in the Middle East, 26, 7-10 Zohary, M. (1973) Geobotanical foundations of the Middle East. Gustav Fischer Verlag, Stuttgart.

14

2 Desert versus steppe: Multi-scale spatial patterns in diversity of ant communities in Iran

Omid Paknia and Martin Pfeiffer In revision, Insect Conservation and Diversity

ABSTRACT 1 Knowing the patterns of biodiversity is the first and inevitable step for any conservation program. Nevertheless, the diversity pattern of many insect taxa is unknown in vast regions of desert and steppe regions of Asia. In this study, we investigated the variation of ant species diversity, equitability, and turnover across fine-to-broad scales among two major steppe and desert biomes in Iran. 2 We sampled ants with a hierarchical sampling design across 10 degrees of latitudinal gradient. We calculated species richness, Shannon diversity, and evenness and partitioned species richness and Shannon diversity into its components, viz. alpha and beta across levels. 3 Alpha species richness increased across latitudinal gradient but beta species richness did not associate with latitude. Species richness and diversity measures increased across the hierarchical levels because of an increase of their beta components in both biomes. Both biomes represented distinctive faunal groups. Beta values were significant at all levels in both biomes. A Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

large amount of beta diversity within biomes was explained by distance. Only at the highest level species richness and Shannon diversity of steppe biome were significantly higher than those of in desert biome. 4 Significant values of beta species richness and beta diversity across all hierarchical levels suggests that ants are highly localised in small patches of habitats in both biomes, which make them vulnerable to extinctions due to habitat disturbance.

Keywords: Ecological gradient, evenness, Formicidae, hierarchical partitioning, Iran, Shannon diversity, species richness.

INTRODUCTION

A key research agenda in insect ecology is to explain geographic patterns of species richness. Insects are major components of the animal kingdom in many ecosystems. Nevertheless, a reverse bias in documentation of animal species richness and their patterns at the global scale has been noticed and warned (Clark & May, 2002). In spite of a remarkable growing attempt to overcome this dilemma during last decade, a considerable geographical bias for this endeavour has remained among well and less developed countries in the temperate regions and the tropics. As a consequence, this inadequacy has left vast regions and habitats with no research on pattern of diversity in many insect taxa in addition to other organisms (Myers et al., 2000). Arid and semi-arid regions of the Palearctic realm in the Southwest Asia and the North Africa, stretching from Atlantic coasts of Morocco to the Gobi desert in the Central Asia, are instances of poor studied regions in the diversity patterns of insects. Among many insect taxa, ants have been chosen very often as a model organism to demonstrate geographical patterns of insects in many ecosystems around the world. This is because of their multifunctional roles in ecosystem services, as herbivore, predator or scavenger (Hölldobler & Wilson, 1990). High number of species in many ecosystems and considerable change in species richness and composition across

16 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants environmental gradients are other favourable aspects of this taxon for ecological studies (Gotelli & Ellison, 2002). The geographical pattern of ant species richness has been demonstrated well in some regions such as the North and the South America and Australia (e.g. Davidson, 1977; Andersen, 1997; Vasconcelos & Vilhena, 2006). Further, ants have been employed as indicator of disturbance or surrogate of biodiversity in these regions (e.g. King et al., 1998). However, arid regions of Asia and the North Africa have been critically underrepresented, by only one remarkable study on ant species richness and assemblage organizations in the Gobi steppe and deserts (Pfeiffer et al., 2003). The present study aims to provide further information on the fauna and geographical pattern of ant diversity in arid and semi-arid regions of Asia across a 1200 km north-to-south transect. In particular, the recognition of species richness and diversity patterns is important, because it is considered as a fundamental point for any scientific act in conservation biology (Bestelmeyer et al., 2003; Leather et al., 2008). “Species richness” as the main proxy of biodiversity has been at the heart of many community structure studies across scales or along gradients (e.g. Sanders et al., 2007; Dahms et al., 2010). This is mostly attributable to the simplicity of describing communities at different scales by species richness and its broadly understood meaning. However, species richness does not present any information about the abundance of species, the evenness of communities across the sampling region or the dominant species. Furthermore, for precisely modelling of spatial patterns of species richness and diversity a spatial hierarchical approach that takes several spatial scales into account is useful (Noda, 2004), because spatial patterns of diversity and interacting processes, which operate in communities, may differ depending on the scale(s) used (see Sanders et al., 2007). So far, there are only few studies on ants across long transects, dealing with main components of diversity viz. alpha and beta diversity (e.g. Pfeiffer et al., 2003; Vasconcelos et al., 2010). In addition, no hierarchical multi-scale approach has been used for the demonstration of ant diversity patterns. Applying the spatial hierarchical partitioning approach, the objective of the present work is to explore diversity patterns of ants among two major biomes in Iran, steppe and desert. Steppe and desert biomes of Iran cover nearly 95% of the total area of the country (Zehzad et al., 2002). These two regions differ particularly in their contemporary climate conditions (with steppe biome having higher precipitation rate),

17 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants vegetation complexity, topography (e.g. mean of elevation), history (e.g. paleoclimate), geology, and flora (Zohary, 1973). From the conservation perspective, both steppe and desert biomes have been under disturbance pressure in different ways during last decades. Desert areas have been used as military training exercises since many years ago, which have resulted in disturbance of habitats. Steppe biome, however, faced to deforestation of huge areas, generating high rates of habitat loss and fragmentation. In addition, it has been predicted that climate change will have a significant impact on both steppe and desert biomes of the country through higher temperature and changes in precipitation regimes (Evans, 2009). The first purpose of this study was to demonstrate patterns of ant species richness, Shannon diversity, and evenness across hierarchical levels for the whole region and its two main biomes, desert and steppe. Our second aim was to partition diversity (gamma) into its components (alpha and beta) across hierarchical levels to explore their patterns and relationships from the finest to broadest grains for the whole region and the two main biomes. The third objective of the study was to compare spatial patterns of ant diversity among steppe and desert biomes. Specifically we tested two hypotheses: 1) the highest proportion of ant diversity is generated at the highest spatial levels for the whole region, viz. among biomes; 2) the number of species and diversity is higher in steppe biome than in desert biome at all levels.

METHODS Study area Two main steppe ecoregions, Zagros Mountains forest steppe in the west of the country and the Elburz Range forest steppe in the north of Iran, are characterized by semi-arid climate, cold winters, mild summers, and a wide range annual precipitation from 123 mm to 860 mm (Iranian weather organization). The Zagros Mountains forest steppe includes oak-dominant deciduous forests and pistachio-almond forests and shows a diversified steppe flora. The Elburz Range forest steppe is covered partly by remnants of Juniper forests and several shrub species such as pistachio and cotoneaster. The groundcover includes communities of Artemisia and Astragalus (Zohary, 1973). The largest desert region in Iran, the Central Persian desert basins ecoregion is surrounded from the west and the north by Zagros and Elburz steppe

18 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants regions, respectively. The region is characterized by low annual precipitation, 122 mm to 127 mm, hot summers and cold winters and contains the typical flora of Iran’s deserts. The South Iran Nubo-Sindian deserts and semi-deserts ecoregion is located in the northern coastal plain of the Persian Gulf and contains a small part of the Sudanian phytogeographical region (Zohary, 1973). This region is characterised by hot summers and mild winters. We have taken our samples inside these four mentioned ecoregions (Figure 1).

40°0'0"N

0200100 Caspian Sea Kilometers 38°0'0"N 1 2 36°0'0"N 3 4 34°0'0"N Iran

5 32°0'0"N

6 30°0'0"N

7 Central Persian desert basins 8 28°0'0"N Elburz Range forest steppe South Iran Nubo-Sindian desert Persian Gulf Zagros Mountains forest steppe 26°0'0"N 44°0'0"E 46°0'0"E 48°0'0"E 50°0'0"E 52°0'0"E 54°0'0"E 56°0'0"E 58°0'0"E 60°0'0"E 62°0'0"E Figure 1 Map of Iran depicting four ecoregions and eight study sites. The numbers indicate the following regions: (1) Mirzabailu, (2) Khoshyelagh, (3) Turan, (4) Kavir, (5) Kolahghazi, (6) Dena, (7) Mond, (8) Naiband.

Ant sampling Epigaeic ant assemblages were sampled during May to July of 2007 in 16 sample sites within eight nature reserves along a latitudinal transect through Iran (Figure 1). Pitfall trapping method was used for sampling ants, an appropriate technique for short-term investigations of open habitats and epigaeic ant species (Agosti et al., 2000). At each study site, two pitfall trap transects were randomly laid out that were at least 1000 m and at most 2000 m apart. Each transect comprised 30 pitfall traps spaced at 10 m

19 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants intervals. Pitfall traps were left open for 72 hours. Samples were fixed in 96% ethanol, brought to the lab, cleaned from debris and soil, and sorted for species. Ants were classified to species when it was achievable (Paknia et al., 2010), or were assigned to morphospecies (later referred to as “species”). Voucher specimens are held in the AntBase.Net Collection of the Institute of Experimental Ecology of University of Ulm (http://www.antbase.net) and in the private ant collection of the corresponding author. Sampling design Ants were collected by a nested sampling design. Eight sites were nested within four ecoregions located within two biomes, viz. desert and steppe (Table 1 and Figure 1). At each site, two transects were set up. Each transect was considered as a “fine grain” at the lowest level of our hierarchical design, the two transects of one site as the “locale grain”, the two local grains of one ecoregion as the “ecoregion grain” and, at the higher level, the two steppe ecoregions were pooled as the “steppe biome” and two desert and semi-desert ecoregions were pooled as the “desert biome”. The whole transect was considered as the “regional” grain. Therefore, our sampling design contained five spatial levels: fine, locale, ecoregional, biome and regional. According to the “scale concept” of Whittaker et al. (2001) in this nested design the extent of the study remained constant across scales, but the size of grain or focus doubles in size from one scale to the next higher scale by pooling samples. Data analysis

We used the Mao-Tao function for computing the observed species richness (Sobs) curves. We computed Chao2’s for the expected number of species curves and also singletons and doubletons curves for the observed rarity. Singletons are species known from a single occurrence and doubletons are species known from two occurrences. All curves were computed by ESTIMATES 7.5.2 software (Colwell, 2005) for the both biomes and for the whole region. ESTIMATES package was also used to compute confidence intervals of observed species richness across fine-to-broad grains. Shannon-Wiener has been used widely as a diversity measure in ecological studies. However, recent conceptual studies have revealed that these classical indices, indeed are not diversities but “entropies” (Jost, 2006). As a diversity measure for our study, we employed the exponential of Shannon entropy. This diversity measure applies the concept of “effective number of species” (ENS), instead of traditional diversity indices (Jost, 2006). Each diversity index can be transformed into its ENS (or number

20 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants equivalent). Briefly, for all diversity indices, ENS is identical to species richness if all species of a sample have the same occurrence (Jost, 2006). Contrary to species richness, which is a diversity measure of the order 0 and is sensitive to rare species, Shannon diversity is a diversity measure of the order 1, weights all species by their frequency and reflects both commonness and rarity of a given community (Jost, 2007). In this paper, we refer to the exponential of Shannon entropy as “Shannon diversity” – not to be confounded with the “Shannon-Weiner index”. Shannon diversity was calculated as eH, with

ln

Where pi = the proportion of occurrences in the ith species. The SPADE software (Chao & Shen, 2003) was used for calculating Shannon diversity (eH) and its lower and upper bounds of confidence intervals. For both measure of species richness and Shannon diversity, 95% confidence intervals have been used for determination of significant differences among sites across fine-to-broad levels. Evenness (E) has been calculated as E=eH/S, where S is the number of observed species in a sample. We demonstrated the relationship between latitudinal gradient and alpha and beta species richness of ants with the Pearson correlation test. The beta-Simpson index

(βsim) was used as measure of turnover among pitfall transects across altitude gradient, with respect to its focusing on compositional differences and its independence from the species richness gradient (Koleff et al., 2003). Significance of faunal group among two biomes was assessed by ANOSIM and Mantel correlation computed in R (R Development Core Team, 2007) using the “vegan” package (Oksanen et al., 2009). We applied hierarchical partitioning for exploring the pattern of diversity components and the magnitude of turnover across scales. Partitioning of species richness was assessed by its additive definition (Sα + Sβ = Sγ) and the partitioning of Shannon H H diversity by its multiplicative definition (e α × β = e γ). The main advantage of the additive function for species richness is that both alpha and beta have the same units (Gering et al., 2003). The multiplicative definition was used for Shannon diversity because the Shannon diversity is concave only in the multiplicative definition when samples are weighted by relative number of individuals in each sample (see Jost 2007

21 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants for details). In a nested hierarchical approach, the mean of the alpha diversity of communities or samples at the lowest level is considered merely as an alpha, whereas at higher levels, gamma diversity at a given level is equal to the mean of alpha diversity at the next higher level. The difference between the alpha diversity at a given level and the alpha diversity at the next higher level is considered as the beta diversity of that given level. The total species richness or gamma species richness in this study can be stated as γ species richness total = α fine + β fine + β locale + β ecoregion + β biome.

The total Shannon diversity can be stated as γ Shannon diversity total = α fine × β fine × β locale

× β ecoregion × β biome. The contrasting partitions of species richness and Shannon diversity can be explained by patterns of species rarity and dominance, respectively (Crist et al., 2003). This is because Shannon diversity takes the abundance of species into account, but species richness is free of abundance measures and reacts to rare species. We employed the program PARTITION v. 3 (Veech & Crist, 2009) to determine the statistical significance of differences in observed and expected diversity components, alpha and beta, by individual-based randomization tests with 10000 repetitions. In our study, the beta of species richness at each level was the number of species, but the beta of Shannon diversity ranged from 1, when two communities have identical species composition, to 2, when two communities have two distinctive species compositions.

RESULTS

Whole region We found a total of 69 species along the latitudinal transect, including several new records for the country and one new species for the world (Paknia et al., 2010; Radchenko & Paknia, 2010). From a total of 17 recorded ant genera, four genera, Cataglyphis, Messor, Camponotus and Monomorium, comprised > 50% of total species richness. The three most widespread species were Cataglyphis nodus BRULLÉ,

1833, Lepisiota semenovi (RUZSKY, 1905), and Messor intermedius SANTSCHI, 1927.

The top three most abundant species were Monomorium indicum kusnezovi SANTSCHI,

1928, Cataglyphis niger (ANDRÉ, 1881), and Tetramorium striaventre MAYR, 1877. The observed species richness accumulation curve for the whole region reached an obvious asymptote close to the Chao2 estimated species richness curve, showing that sampling by pitfall trap method was relatively complete (Figure 2a). The number of

22 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants singletons did not approach to zero, suggesting that there have been several species which occurred only by one incidence in the pooled total inventory.

Whole region a) 70

60

50

40 Sobs Singletons Doubletons 30 Chao2 Number of species

20

10

0 0 200 400 600 800 1000 1200 1400 1600 1800 Number of species occurences

b) 50 Biomes

40

30 Sobs desert Singletons desert Doubletons desert Chao2 desert 20 Sobs steppe

Number of species Singletons steppe Doubletons steppe Chao2 steppe 10

0 0 200 400 600 800 1000 Number of species occurences Figure 2 Sample-based species accumulation curves of the observed species richness, estimated species richness (Chao2) and curves of singletons and doubletons at two levels; a) whole region and b) biome levels.

23 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

Furthermore, the higher ratio of “singletons” to “doubletons” indicates that there have been probably missing species in our whole region inventory. Ant species richness within pitfall transects varied from 8 to 14 (Table 1), and increased across the latitude (r = 0.475, df = 13, t = 1.95, P = 0.07; Figure 3a). However, beta species richness across the latitude did not follow specific pattern (r = - 0.391, df = 13, t = - 1.53, P = 0.14; Figure 3b).

Table1 Species diversities of ants from fine scale to regional scale. The following parameters were obtained from our samples: number of species occurrences (SOCs), observed species (Sobs), lower bound of 95 % confidence interval of Sob (95 % LB), upper bound of 95 % confidence interval of Sob (95 % UB), observed Shannon diversity (eH), lower bound of 95 % confidence interval of eH (95 % LB), upper bound of 95 % confidence interval of eH (95 %

UB), observed evenness (Eobs), number of unique and duplicate species.

H e 95 % LB 95 % UB Eobs Unique Duplicate ﭜڮ ᾤ Plot or level name SOCs Sobs 95 % LB Fine level Mirzabailu A 83 11 7 15 6.68 5.51 7.84 0.61 4 1 Mirzabailu B 128 14 10 16 9.78 8.53 11.03 0.7 2 1 Khoshyelagh A 87 10 8 11 5.83 4.78 6.89 0.58 2 2 Khoshyelagh B 160 13 11 13 9.64 8.68 10.59 0.74 1 0 Kolahghazi A 78 8 8 8 5.34 4.4 6.28 0.67 0 3

Steppe biome Kolahghazi B 105 9 9 9 6.38 5.43 7.33 0.71 0 0 Dena A 143 13 11 13 9.69 8.7 10.68 0.75 1 2 Dena B 105 9 8 9 6.42 5.55 7.3 0.71 1 0 Turan A 166 12 7 13 8.08 7.32 8.85 0.67 3 1 Turan B 165 12 7 13 7.78 6.98 8.58 0.65 3 0 Kavir A 119 8 8 8 6.14 5.49 6.8 0.77 0 1 Kavir B 143 10 9 10 7.68 6.94 8.42 0.77 0 2 Mond A 75 10 7 13 7.23 6.17 8.28 0.72 3 0 Mond B 116 9 5 11 5.56 4.84 6.29 0.62 3 0 Desert biome biome Desert Naiband A 89 10 7 12 6.18 5.12 7.25 0.62 2 1 Naiband B 81 9 9 9 7.69 6.9 8.48 0.85 1 0 Locale level Mirzabailu 211 16 14 17 9.2 8.12 10.28 0.58 2 3 Khoshyelagh 248 15 13 16 9.94 8.95 10.94 0.66 1 1

Kolahghazi 183 10 10 10 7.00 6.18 7.81 0.7 0 0

Steppe biome Steppe biome Dena 248 14 12 15 10.1 9.23 10.96 0.72 1 2

Turan 331 12 11 12 8.82 8.24 9.4 0.74 0 1 Kavir 262 11 11 11 7.29 6.69 7.91 0.66 0 1 biome Desert Desert Mond 191 15 11 19 9.37 8.3 10.43 0.62 4 0 Naiband 170 13 10 16 7.79 6.83 8.74 0.6 3 0 Ecoregional level Elburz steppe 459 24 21 26 13.18 12.03 14.32 0.55 3 3 Zagros steppe 431 24 23 25 17.08 15.89 18.28 0.71 1 2 Central desert 593 17 17 17 11.39 10.72 12.07 0.67 0 0 Nubo desert 361 24 18 30 14.03 12.82 15.26 0.58 5 1 Biome level Steppe biome 890 45 43 47 29.24 27.63 30.86 0.65 3 3 Desert biome 954 37 31 43 19.77 18.63 20.92 0.53 5 1 Regional level Total transect 1844 69 65 73 38.9 37.25 40.57 0.56 6 4

24 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

a) 14

13

12

11

10 Alpha species richness species Alpha

9

8

27 28 29 30 31 32 33 34 35 36 37 Latitude

b) 1.0

0.8

0.6

0.4 Beta species richness species Beta

0.2

0.0

27 28 29 30 31 32 33 34 35 36 37 Latitude Figure 3 Spatial variation of a) alpha ant species richness and b) beta ant species richness across the ~ ten degree latitudinal gradient. A LOESS smoothing was fitted on ant species richness in each graph to show the detailed trend of species richness across desert-steppe- desert-steppe gradient.

25 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

For the whole region, records of species richness and Shannon diversity revealed a constant trend of increasing species richness across levels (Table 1). Highest evenness was observed at the finest grain and decreased as the level increased. Hierarchical partitioning showed that the highest portion of β diversity in species richness occurred in β biome, indicating that the highest turnover of species regardless of their abundances happened among biomes. However, in Shannon diversity partitioning, β ecoregional represented the greatest part (Table 2), suggesting that the highest turnover of diversity occurred among ecoregions.

Table 2 Spatial partitioning of species richness and Shannon diversity of ant assemblages across a hierarchically study in Iran as calculated by randomization with PARTITION v.3. All observed values differed significantly from the expected values at the 0.0001 level, marked by asterisk (***). Alpha components were lower than expected, while beta components were higher than expected at all scales, except β locale of the whole region species richness.

Partition/Scale Species richness Shannon diversity Observed Expected Observed Expected Whole region β biome 28.6 *** 4.7 1.6 3*** 1.02 β ecoregion 18.9 *** 6.1 1.77*** 1.04 β locale 8.5 8.2 1.56*** 1.08 β fine 2.4 *** 10.4 1.18 *** 1.16 α fine 10.7 *** 39.6 7.33 *** 28.55

Steppe β ecoregion 21 *** 2.5 1.96 *** 1.03 β locale 10.07 *** 3.8 1.64 *** 1.06 β fine 2.65 *** 6.1 1.19 *** 1.13 α fine 11.28 *** 32.6 7.65 *** 23.54

Desert β ecoregion 17.33 *** 3.6 1.61 *** 1.02 β locale 7.16 *** 4.1 1.49 *** 1.05 β fine 2.27 *** 4.6 1.17 *** 1.09 α fine 10.24 *** 24.6 7.07 *** 16.84

Desert versus steppe Forty five species were sampled in steppe biome and 37 species in desert biome. Thirteen species belonging to eleven genera were in common between two biomes. Temnothorax, Lasius, Proformica and Aphaenogaster occurred only in steppe biome, and Paratrechina and Pachycondyla occurred only in desert biome. Camponotus with six species was the most speciose genus in the steppe, while in desert Cataglyphis with nine species was the most species rich genus. Three top abundant species which occurred only in the steppe biome were Cataglyphis cugiai, Crematogaster sp. 1, and

26 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

Cataglyphis kurdestanicus PISARSKI, 1965, and three top abundant species which occurred only in the desert biome were Cataglyphis cinnamomeus (KARAVAIEV,

1910), C. lividus (ANDRÉ, 1881), and C. ruber (FOREL, 1903). In the steppe biome, the observed species richness and Chao2 estimator were more asymptotic and closer together than in desert biome (Figure 2b). The singletons and doubletons curves of the steppe biome were hump-shaped and declined at higher sampling intensity. For desert biome, both curves were almost flat. None of singletons and doubletons curves approached to zero. Ant faunal dissimilarity among biomes was significantly greater than those within the biomes (ANOSIM statistic, R = 0.3348, P < 0.005). Species richness and Shannon diversity increased across the levels in both steppe and desert biomes. Evenness, in the steppe biome, decreased from fine level to ecoregional level, but it increased at the biomes level (Figure 4a). In the desert biome, however, it decreased across all levels (Figure 4b). Contrary to our assumption that species richness and diversity would be higher in steppe biome than in desert biome at all scales, the observed species richness and diversity of desert and steppe differed significantly only at the biome level (Table 1). The highest proportion of β-species richness and β-Shannon diversity for both biomes occurred at the highest scale, among ecoregions within biomes. All values were higher than expected (Table 2). The lowest component of β-species richness and β- Shannon diversity for both biomes were observed among transects within locales (Table 2). At this scale observed values of species richness were lower than expected, but those of Shannon diversity were higher than expected. The amount of α-species richness and α-Shannon diversity were much lower than expected by chance in both biomes (Table 2). Dissimilarity of ant compositions within both biomes associated strongly and significantly with distance (Mantel r steppe = 0.78 P = 0.001 vs. Mantel r desert = 0.63, P = 0.008).

DISCUSSION

The key result of this study is that patterns of species richness, diversity and evenness are generally the same in the both studied biomes; however, steppe has a greater species richness and diversity at the highest level. Additionally, species richness and

27 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants diversity of ants in the region is generated mainly by turnover of species among large grains, ecoregions and biomes.

50 1.0 Species richness (a) steppe 45 Shannon diversity Evenness 40 0.8

35

30 0.6

25 Evenness 20 0.4

15 Effective number of species

10 0.2

5

0 0.0 Fine Local Ecoregional Biome Spatial scales

40 1.0 Species richness (b) desert Shannon diversity 35 Evenness 0.8 30

25 0.6

20 Evenness 0.4 15

Effective number of species of number Effective 10 0.2

5

0 0.0 Fine Local Ecoregional Biome Spatial scales Figure 4 Mean of species richness, Shannon diversity and evenness of ants on four spatial levels for a) steppe biome and b) desert biome. The scaling of species richness and Shannon diversity (left Y axis) differ from that of evenness (right Y axis).

28 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

Both β species richness and β Shannon diversity proportions were the largest at the two highest levels. These results are consistent with the findings of Summerville et al. (2003) and Gering et al. (2003) for temperate forest Lepidoptera and arboreal Coleoptera, respectively; and propose a general pattern of determination of insect diversity and composition by broad scale factors. The large turnover of species at the biome level and significant higher dissimilarity among biomes than within biomes indicate that both biomes represent distinctive species compositions. In a comparable study, Pfeiffer et al. (2003) also found different species composition in steppe and desert of Mongolia, as the steppe assemblage consisted of cold resistant species of Formica and Myrmica and desert assemblages were dominated by hot climate specialists of Cataglyphis, Messor. In a big contrast to their findings, steppe regions of our study contained no member of Formica and Myrmica and, instead, were dominated by arboreal Camponotus and Crematogaster ants. This clear cut difference is mainly because of the very cold temperatures which occur in the steppe regions of Mongolia compared to Iranian steppe regions. In addition, oak and junipers forests of Iranian steppe supply the preference niche of arboreal ants with nesting sites, food resource or microhabitats, while steppe regions of Mongolia are dominated by shrubs (Pfeiffer et al., 2003). Vasconcelos and Vilhena (2006) also found a low similarity between ant species composition of forest and savanna in the Brazilian Amazon. These results confirm that distinctive ant assemblages are particularly related to their habitat types, especially vegetation types (Andersen, 1983). The species richness and diversity of steppe and desert regions differed significantly at the highest spatial grain. The composition of ants in steppe biome included several ant genera which did not occur in desert biome or occurred with a low number of species. In addition to arboreal ant genera such as Camponotus and Crematogaster which distributed mainly in the steppe biome, cryptic species such as Plagiolepis were also more diverse in steppe than desert biome; and cold climate specialists such as Lasius only occurred in steppe biome. These findings can be attributed to greater habitat complexity in steppe biome, where the heterogeneity in topography (e.g. elevation) and vegetation (from short scrub plants to sparse forests) is higher than in desert biome. It has been demonstrated that heterogeneity in topography or vegetation structure has a great impact on ant species distribution and composition in arid and semi-arid regions (Andersen, 1983; Sanders et al., 2003). These explanations mirror the old idea that habitat heterogeneity could be the most important factor that shapes

29 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants species-area relationships (Macarthur & Wilson, 1967), suggesting that larger areas usually have more habitat heterogeneity and which is generally positively correlated with species richness and diversity (Tews et al., 2004). In contrast to Shannon diversity and species richness, the mean of evenness decreased across levels for the whole region and both biomes as well. Within a single grain, species richness and evenness have been shown to have a strong positive correlation for invertebrate (Stirling & Wilsey, 2001). However, the relationship of species richness or diversity and evenness across different scales has been poorly documented (but see Leponce et al., 2004). Leponce et al. (2004) showed that evenness of ants decrease with increasing sampling grain. Evenness decreases from smaller grain to a larger grain if some species have restricted distributions while others are generalists with wide distribution (Wilson et al., 1999). In our study, the restricted distribution of 45 species to fine or locale scales and the wide distribution of a few other species, e.g.

Monomorium indicum kusnezovi, C. nodus and Lepisiota semenovi (RUZSKY, 1905), is probably the major reason for the decrease of evenness when the grain increases. This conclusion is supported by the decrease of commonness and increase of rarity of species across scales (Paknia & Pfeiffer, unpublished data). Furthermore, our results add weight to the suggestion that insects species commonness and evenness are determined at finer spatial grains (Summerville et al., 2003). Partitioning of species richness and Shannon diversity to α and β components in steppe and desert biome showed a similar trend, indicating those factors or parameters that are responsible for diversity of ants at the different levels are similar in both biomes. The greatest fraction of β species richness and β Shannon diversity occurred at the ecoregional level, as a consequence of large distance between ecoregions of each biome. A strong correlation between ant assemblages’ dissimilarity and geographical distance has been documented by other researchers in dry and tropical biomes (Pfeiffer et al., 2003; Vasconcelos et al., 2010). In particular, in the steppe biome, species composition and evenness changed very largely from the Elburz ecoregion to the Zagros ecoregion (Table 1), although, both ecoregions comprised the same number of species (24 species). These two ecoregions are separated by the vast Central desert that can act as a biogeographical barrier for dispersal of ants among these two ecoregions (Figure 1). Species composition of the two ecoregions within desert biome also differed notably, as they are also separated by Zagros mountain ranges and a great distance which exists among sample sites of these ecoregions.

30 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

These desert ecoregions differed also considerably in other aspects such as climate conditions and phytogeographical structure (Zohary, 1973; Breckle, 2002). In essence, Central Persian desert basin ecoregion of Iran shows similar features as the cold deserts of Central Asia, but the Nubo Sindian ecoregion is related to the hot Arabian deserts (Breckle 2002). Faunistically speaking, this study can be seen as a substantial contribution to ant fauna of Iran. Although, all collected species were not identified to species level, the description of one species to science and the addition of thirteen new records to the recently published ant species list of the country (Paknia et al., 2008), indicate the poor knowledge about the faunas occurring in different regions of the country and the need of further sampling efforts in these regions.

Implications for conservation The role of ants in engineering and maintaining ecosystem integrity in arid regions has been emphasized by researchers (e.g. Whitford, 1996). As a result, conserving ant assemblages in arid regions should be a priority not only because of themselves, but also because of sustainability of arid ecosystems. In particular, our observations from the region suggest that the steppe regions of Iran are under more pressure of anthropologic disturbances such as overgrazing and farming than deserts. Impact of future climate change will also be harder in montane steppe regions, as marginal regions of deserts, by increasing of dryness and threats of desertification (Ezcurra, 2006) and shifting of land use by humans from drier regions to these regions (Evans, 2009). Furthermore, significant beta species richness and beta diversity across fine-to-broad hierarchical levels in both biomes suggest that many species have restricted distributions in their specific habitats across the long latitudinal gradient. This localized distribution will make many ant species vulnerable to local or regional extinctions (Dunn, 2005). But because the fauna of the region has not been understood well, there could be even global extinctions for endemic ants which are distributed in a narrow habitat in arid and semi-arid regions of Iran.

31 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

ACKNOWLEDGMENTS The kind support of Elisabeth K.V. Kalko (head of Experimental Ecology Institute, Ulm University) made this work possible. We are grateful to the staff of the Iranian Department of the Environment for allowing this study to be conducted in the natural reserves of Iran. We thank Alexander Radchenko for extensive help and advice with ant species identification and Theresa Jones for language correction. Financial support from the German Academic Exchange Service (DAAD) for the PhD study of OP in Germany is gratefully acknowledged.

32 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

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Agosti, D., Majer, J.D., Alonso, L.E. & Schultz, T.R. (2000) Ants: Standard methods for measuring and monitoring biodiversity. Smithsonian Institution Press, London. Andersen, A.N. (1983) Species diversity and temporal distribution of ants in the semi- arid mallee region of northwestern Victoria. Australian Journal of Ecology, 8, 127-137 Andersen, A.N. (1997) Functional groups and patterns of organization in North American ant communities: a comparison with Australia. Journal of Biogeography, 24, 433-460 Bestelmeyer, B.T., Miller, J.R. & Wiens, J.A. (2003) Applying species diversity theory to land management. Ecological Applications, 13, 1750-1761 Breckle, S.-W. (2002) Salt deserts in Iran and Afghanistan. Sabkha Ecosystems (ed. by H.-J. Barth and B. Boer), pp. 109-122. Kluwer Academic Publishers, Dordrecht. Chao, A. & Shen, T.J. (2003) Program SPADE (species prediction and diversity estimation). Program and user's guide published at http://chao.stat.nthu.edu.tw In: Clark, J.A. & May, R.M. (2002) Taxonomic Bias in Conservation Research. Science, 297, 191b-192 Colwell, R.K. (2005) EstimateS: Statistical estimation of species richness and shared species from samples. Version 7.5 User's Guide and application published at: http:purl.oclc.org/estimates. Crist, T.O., Veech, J.A., Gering, J.C. & Summerville, K.S. (2003) Partitioning species diversity across landscapes and regions: a hierarchical analysis of α, β, and γ diversity. The American Naturalist, 162, 734-743 Dahms, H., Mayr, S., Birkhofer, K., Chauvat, M., Melnichnova, E., Wolters, V. & Dauber, J. (2010) Contrasting diversity patterns of epigeic arthropods between grasslands of high and low agronomic potential. Basic and Applied Ecology, 11, 6-14 Davidson, D.W. (1977) Species diversity and community organization in desert seed- eating ants. Ecology, 58, 711-724 Dunn, R.R. (2005) Modern insect extinctions, the neglected majority. Conservation Biology, 19, 1030-1036 Evans, J. (2009) 21st century climate change in the Middle East. Climatic Change, 92, 417-432 Ezcurra, E. (2006) Global deserts outlook. United Nations Environment Programme, Nairobi, Kenya. Gering, J.C., Crist, T.O. & Veech, J.A. (2003) Additive partitioning of species diversity across multiple spatial scales: Implications for regional conservation of biodiversity. Conservation Biology, 17, 488-499 Gotelli, N.J. & Ellison, A.M. (2002) Biogeography at a regional scale: Determinants of ant species density in New England bogs and forests. Ecology, 83, 1604- 1609 Hölldobler, B. & Wilson, E.O. (1990) The ants. Belknap Press of Harvard University Press, Cambridge, Massachusetts, USA. Jost, L. (2006) Entropy and diversity. Oikos, 113, 363-375 Jost, L. (2007) Partitioning diversity into independent alpha and beta component. Ecology 88, 2427-2439

33 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

King, J.R., Andersen, A.N. & Cutter, A.D. (1998) Ants as bioindicators of habitat disturbance: validation of the functional group model for Australia's humid tropics. Biodiversity and Conservation, 7, 1627-1638 Koleff, P., Gaston, K.J. & Lennon, J.J. (2003) Measuring beta diversity for presence- absence data. Journal of Animal Ecology, 72, 367-382 Leather, S.R., Basset, Y. & Hawkins, B.A. (2008) Insect conservation: finding the way forward. Insect Conservation and Diversity, 1, 67-69 Leponce, M., Theunis, L., Delabie, J.H.C. & Roisin, Y. (2004) Scale dependence of diversity measures in a leaf-litter ant assemblage. Ecography, 27, 253-267 Macarthur, R.H. & Wilson, E.O. (1967) The Theory of Island Biogeography. Princeton University Press, Princeton. Myers, N., Mittermeier, R.A., Mittermeier, C.G., Da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853-858 Noda, T. (2004) Spatial hierarchical approach in community ecology: a way beyond high context-dependency and low predictability in local phenomena. Population Ecology, 46, 105-117 Oksanen, J., Kindt, R., Legendre, P., O'hara, B., Simpson, G.L., Peter, S., Stevens, M.H., H. & Wagner, H. (2009) vegan: community ecology package. R package version 1.15-4. (http://cran.r-project.org/). Paknia, O., Radchenko, A., Alipanah, H. & Pfeiffer, M. (2008) A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News, 11, 151-159 Paknia, O., Radchenko, A. & Pfeiffer, M. (2010) New records of ants (Hymenoptera: Formicidae) from Iran. Asian Myrmecology, 3, 29-38 Pfeiffer, M., Chimedregzen, L. & Ulykpan, K. (2003) Community organization and species richness of ants (Hymenoptera: Formicidae) in Mongolia along an ecological gradient from steppe to Gobi desert. Journal of Biogeography, 30, 1921-1935 R Development Core Team (2007) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. (http://www.R-project.org). Radchenko, A. & Paknia, O. (2010) Two new species of the genus Cataglyphis Foerster, 1850 (Hymenoptera: Formicidae) from Iran. Annales Zoologici (Warsaw), 60, 69-76 Sanders, N.J., Lessard, J.-P., Fitzpatrick, M.C. & Dunn, R.R. (2007) Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. Global Ecology and Biogeography, 16, 640-649 Sanders, N.J., Moss, J. & Wagner, D. (2003) Patterns of ant species richness along elevational gradients in an arid ecosystem. Global Ecology & Biogeography, 12, 93-102 Stirling, G. & Wilsey, B. (2001) Empirical relationships between species richness, evenness, and proportional diversity. American Naturalist, 158, 286-299 Summerville, K.S., Boulware, M.J., Veech, J.A. & Crist, T.O. (2003) Spatial variation in species diversity and composition of forest Lepidoptera in eastern deciduous forests of North America. Conservation Biology, 17, 1045-1057 & .Rger, K., Wichmann, M.C., Schwager, M¶أTews, J., Brose, U., Grimm, V., Tielb Jeltsch, F. (2004) Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures. Journal of Biogeography, 31, 79-92

34 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

Vasconcelos, H.L. & Vilhena, J.M.S. (2006) Species turnover and vertical partitioning of ant assemblages in the Brazilian Amazon: A comparison of forests and savannas. Biotropica, 38, 100-106 Vasconcelos, H.L., Vilhena, J.M.S., Facure, K.G. & Albernaz, A.L.K.M. (2010) Patterns of ant species diversity and turnover across 2000 km of Amazonian floodplain forest. Journal of Biogeography, 37, 432-440 Veech, J.A. & Crist, T.O. (2009) PARTITION: software for hierarchical partitioning of species diversity, version 3.0. published at http://www.users.muohio.edu/cristto/partition.htm. Whitford, W.G. (1996) The importance of the biodiversity of soil biota in arid ecosystems. Biodiversity and Conservation, 5, 185-195 Whittaker, R.J., Willis, K.J. & Field, R. (2001) Scale and species richness: Towards a general, hierarchical theory of species diversity. Journal of Biogeography, 28, 453-470 Wilson, J.B., Steel, J.B., King, W.M. & Gitay, H. (1999) The effect of spatial scale on evenness. Journal of Vegetation Science 10, 463-468 Zehzad, B., Kiabi, B.H. & Madjnoonian, H. (2002) The natural areas and landscape of Iran: An overview. Zoology in the Middle East, 26, 7-10 Zohary, M. (1973) Geobotanical foundations of the Middle East. Gustav Fischer Verlag, Stuttgart.

35 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

Appendix S1. List of ant species and their occurrence (number of occurrences) or their absent (-) in sixteen pitfall trap transects from north to south of Iran. Numbers indicate site numbers on the map (Figure 1). The two transects within each site were named by ‘a’ or ‘b’ according to their arrangement along North-South latitudinal transect. Ant species 1a 1b 2a 2b 3a 3b 4a 4b 5a 5b 6a 6b 7a 7b 8a 8b Aphaenogaster gibbosa (LATREILLE, 1798) - 1 ------Camponotus aethiops (LATREILLE, 1798) ------17 8 - - - - Camponotus fedchenkoi MAYR, 1877 - 2 - 19 ------Camponotus gestroi EMERY, 1878 ------13 1 - - - - Camponotus interjectus MAYR, 1877 * 1 3 11 1 ------Camponotus oasium FOREL, 1890 ------13 23 15 1 - - Camponotus staryi PISARSKI, 1971 * ------2 - - - - - Camponotus turkestanicus EMERY, 1877 - - - - 2 1 ------Cardiocondyla sp. 1 - 6 ------Cardiocondyla sp. 2 - - - - 17 11 ------Cardiocondyla sp. 3 ------6 9 ------Cataglyphis cf. foreli (RUZSKY, 1903) - - 1 - 1 28 ------Cataglyphis cinnamomeus (KARAVAIEV, 1910) * - - - - 22 28 - 10 ------Cataglyphis cugiai MENOZZI, 1939 * 24 17 28 30 ------Cataglyphis emeryi KARAVAIEV, 1911 - - - - 25 25 ------Cataglyphis frigidus persicus (EMERY, 1906) ------18 - - - - - Cataglyphis isis (FOREL, 1913) ------5 7 - - Cataglyphis kurdestanicus PISARSKI, 1965 * ------29 30 - - - - Cataglyphis lividus (ANDRÉ, 1881) ------23 22 - - - - 13 - - - Cataglyphis niger (ANDRE, 1881) ------27 27 29 29 ------Cataglyphis nodus BRULLE, 1833 - 5 - - 28 1 ------20 9 Cataglyphis ruber (FOREL, 1903) ------1 28 16 12 Cataglyphis stigmatus RADCHENKO & PAKNIA # ------5 - 6 11 Crematogaster sp. 1 10 19 26 24 ------Crematogaster sp. 2 - - 2 16 ------1 - - - - - Crematogaster sp. 3 ------10 5 ------Crematogaster sp. 4 ------18 - - - - - Crematogaster sp. 5 ------1 - Lasius cf. alienus (FOERSTER, 1850) 1 ------Lepisiota frauenfeldi (MAYR, 1855) - - 1 4 ------4 Lepisiota semenovi (RUZSKY, 1905) - - - 14 9 5 12 17 - 6 - - 9 - - - Lepisiota sp. 3 ------3 - Lepisiota sp. 4 ------

36 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

Messor cf. caducus (VICTOR, 1839) ------13 - - Messor cf. striaticeps (ANDRÉ, 1883) ------1 - - Messor denticulatus SANTSCHI, 1927 20 25 ------Messor intermedius SANTSCHI, 1927 - - - - 1 1 - 4 20 17 ------Messor pratennatus ARNOLDI, 1970 * - - 7 12 ------Messor semirufus EMERY, 1925 ------2 4 ------Messor turcmenochorassanicus ARNOLDI, 1977 * ------9 3 - - - - Messor subgracilinodis ARNOLDI, 1970 * - - - - 1 5 - 2 2 6 ------Messor variabilis KUZNETSOV-UGAMSKY, 1927 * - 7 - - - - 2 2 ------Messor sp. 10 ------8 7 Monomorium cf. destructor (JERDON, 1851) ------2 10 Monomorium dentigerum (ROGER, 1862) * ------18 30 - - Monomorium indicum kusnezovi SANTSCHI, 1928 - - - - 30 30 30 30 - 30 - - - 30 - 8 Monomorium sp. 4 ------7 ------Monomorium sp. 5 ------5 - - Monomorium sp. 6 ------7 - - - Monomorium sp. 7 ------29 19 Pachycondyla sennaarensis (MAYR, 1862) ------3 - Paratrechina sp. 1 ------1 Pheidole sp. 1 - - 2 ------Pheidole sp. 2 ------6 4 ------Pheidole sp. 3 ------21 - - - - Plagiolepis sp. 1 2 ------Plagiolepis sp. 2 - - 2 3 ------Plagiolepis sp. 3 ------3 - 2 4 - - 1 - - - Plagiolepis sp. 4 ------4 8 - - - - Plagiolepis sp. 5 ------3 5 - - - - Proformica epinotalis KUSNETSOV-UGAMSKY, 1927 * 4 7 - 8 ------Tapinoma erraticum (LATREILLE, 1798) 1 5 - 3 ------Tapinoma sp. 2 ------1 1 - Temnothorax semenovi (RUZSKY, 1903) 1 1 ------2 - - - - - Tetramorium armatum SANTSCHI, 1927 8 7 8 3 5 3 ------Tetramorium schneideri EMERY, 1898 * 11 23 ------Tetramorium striaventre MAYR, 1877 * - - - 23 25 27 16 20 ------Tetramorium sp. 4 ------14 6 - - - - Tetramorium sp. 5 ------1 - - -

37 Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants

38

3 Hierarchical partitioning of ant diversity: Implications for conservation of biogeographical diversity in arid and semi- arid areas

Omid Paknia and Martin Pfeiffer Accepted for publication in Diversity & Distributions

ABSTRACT

Aim The scale of observation is important in detecting the spatial variation of biological assemblages, which should be taken into consideration for an appropriate plan of biogeographical conservation. We investigated whether 1) World Wildlife Fund’s ecoregion units are the appropriate scale for conserving ant diversity in Iran, 2) each ecoregion represents a distinct ant community composition, and 3) patterns of diversity partitioning differ among four ecoregions.

Location Iran, a sampling transect along four arid and semi-arid ecoregions.

Methods We applied hierarchical partitioning to data collected from a nested sampling design including four hierarchical levels: “local”, “landscape”, “ecoregional”, and “whole-region”. Observed alpha and beta diversity components were compared with values of null distributions. Hierarchical cluster analysis was applied to evaluate similarity of ant species composition among ecoregions. Chapter 3 – Hierarchical Partitioning of Ant Diversity

Results Partitioning of whole-region species richness showed that 85% of the species richness was generated by beta diversity among ecoregions and landscapes. The highest value of diversity was generated by beta diversity among ecoregions. Unlike whole-region partitioning, separate partitioning within each ecoregion revealed that beta component among localities contributed to species richness of each ecoregion. Ecoregions showed different patterns of diversity partitioning. The alpha component contributed largely to the total diversity of two ecoregions, but for two other ecoregions, beta component contributed more than alpha component. Cluster analysis identified four discrete ant species compositions; however, it split landscapes of one ecoregion into two distinct groups.

Main conclusions Whole-region diversity partitioning indicates that ecoregions represent the appropriate scale for conserving ant diversity, and that each ecoregion has a distinct ant fauna. However, different conservation strategies should be considered for different ecoregions due to the differing scales of variation within them. Boundaries of ecoregions remain a subject for further studies. The influence of climate change on ecoregional boundaries should be considered and should be predicted with respect to future conservation maps.

Keywords: beta diversity, conservation biogeography, Iran, Shannon diversity, species richness, WWF ecoregions.

INTRODUCTION

The emphasis of conservation biologists has recently shifted from the conservation of single species within a habitat to the conservation of entire communities within landscapes, ecoregions, or biomes (e.g. Devictor & Robert, 2009). As a result, conservationists try to determine the spatial and temporal distribution of species within large areas, and how the main conservation threats (e.g. invasive species, disturbance or climate change) can change these patterns. However, ecological patterns and processes of communities in nature are recognized as being scale- dependent (Andersen, 1997b; Whittaker et al., 2005).

40 Chapter 3 – Hierarchical Partitioning of Ant Diversity

Conservation biologists are increasingly interested in patterns and consequences related to scale variation (e.g. Ribeiro et al., 2008; Foxcroft et al., 2009). One of the fundamental aspects of a community that varies with spatial scale is its species composition. Variation in species composition can be studied along a range of scales by using a spatial hierarchical sampling design and diversity partitioning (e.g. Crist et al., 2003; Noda, 2004; Lindo & Winchester, 2009). The idea of diversity partitioning originated with Whittaker (1960), who described alpha and beta as components of the total diversity, gamma, and linked them to a spatial scale. Whittaker (1972) demonstrated two potential relationships of these components: a multiplicative relationship (γ = α × β) and an additive relationship (γ = α + β). The additive partitioning of species diversity received attention following Lande's (1996) use of the breakdown of total species richness or diversity additively into its alpha and beta components. Later studies have revealed more suitable methods for hierarchical partitioning of diversity (see Jost, 2007). The main advantage of hierarchical diversity partitioning is that it provides insight into the effects of scale on patterns of diversity. The hierarchical partitioning approach has been used for conservation purposes (Gering et al., 2003; Tylianakis et al., 2006; Ribeiro et al., 2008; Fahr & Kalko, 2010). For example, Gering et al. (2003) have used it to evaluate beetle diversity in temperate forests. Tylianakis et al. (2006) have employed this technique to examine bee and wasp diversity across a tropical land use gradient, and Ribeiro et al. (2008) have applied this method to determine the effect of habitat fragmentation on diversity patterns of butterflies. In the present study, we used this approach to evaluate the variation of ant diversity between and within four ecoregions of Iran.

Ecoregions have been defined as areas containing a distinct assemblage of natural communities whose boundaries approximate the original extent of natural communities before major land-use change (World Wildlife Fund's ecoregions map: Olson et al., 2001) or have been delineated based on biophysical features such as climate zones (e.g. Bailey's ecoregions map: Bailey, 1998). The main use of these maps has been the provision of a global framework for biodiversity conservation. In particular, the World Wildlife Fund’s (WWF) ecoregions are claimed to be appropriate units of areas for biodiversity conservation because they mirror the distribution and biogeographic boundaries of species and communities of animals and plants across the entire planet (Olson et al., 2001; Wikramanayake et al., 2002).

41 Chapter 3 – Hierarchical Partitioning of Ant Diversity

However, to date we are unaware of any attempt to apply these maps to conserving the diversity of ants.

Ants have been subject of many conservation studies, either as organisms that have a negative impact on ecosystems, particularly as introduced species (e.g. Tsutsui & Suarez, 2003; Paknia, 2006) or as organisms affected by habitat disturbance (Dahms et al., 2005; Seppä, 2008; Brühl & Eltz, 2010). In addition, they have been adopted as bio-indicators to determine the environmental response to ecological disturbance associated with anthropogenic land-use (e.g. King et al., 1998; Andersen et al., 2002). Andersen proposed that ants can also be considered a “surrogate” for biodiversity (Andersen, 1997b). Surrogates are used for conservation planning when spatial information about the distribution of biodiversity is not completely known (Rodrigues & Brooks, 2007). Ants have been employed as surrogate taxa for monitoring biodiversity of invertebrates and vertebrates (Andersen, 1997a; Bennett et al., 2009), and they have been shown to provide valuable information for management-based monitoring (Underwood & Fisher, 2006). Although the debate with regard to the effectiveness of surrogate species for the planning of biodiversity conservation has a long history (see Rodrigues & Brooks, 2007), the application of this approach in areas where conservation studies are least developed, such as Iran, would be useful.

In this study, our aim has been to test three predictions. Assuming that the WWF’s ecoregional map accurately reflects the biogeographical units of the region, we predicted that 1) when applying a multi-scale diversity partitioning approach we would find the highest beta diversity of ants at the ecoregional scale, and 2) each ecoregion would have a distinct ecoregional ant species pool. In addition, we predicted that 3) each ecoregion would have different patterns of alpha and beta diversity, because each ecoregion has different habitat, climate, and natural history features.

METHODS

Study area and study sites

Arid and semi-arid regions of Iran comprise more than 95% of the total area of the country. We sampled assemblages of ants during May to July 2007 at eight study sites along a north-south transect (ca. 1200 km, approximately ten degrees of latitude),

42 Chapter 3 – Hierarchical Partitioning of Ant Diversity which included four arid and semi-arid ecoregions (Table 1 and Figure 1): the Elburz range forest steppe ecoregion (hereafter Elburz) comprising the southern and eastern slopes of the Elburz Mountains, which are steppe-like in vegetation and climate, 2) the “Central Persian desert basin” ecoregion (hereafter Central desert) containing the typical flora of Iran’s deserts, 3) the Zagros Mountains forest steppe ecoregion (hereafter Zagros) in which the vegetation comprises steppe forest and a dense ground cover of shrubs, and 4) the South Iran Nubo-Sindian deserts and semi-deserts ecoregion (hereafter Nubo), which comprises the northernmost belt of the Palaeotropical kingdom in Africa and south-western Asia (Zohary, 1973).

Turkmenistan Turkey 40°0'0"N

Afghanistan Iraq Iran 0300150 Caspian Sea Kilometers Pakistan 38°0'0"N 1 Saudi Arabia 2

36°0'0"N 3 4

34°0'0"N

5 32°0'0"N

6

30°0'0"N

7 28°0'0"N Central Persian desert basins 8 Elburz Range forest steppe South Iran Nubo-Sindian desert Persian Gulf Zagros Mountains forest steppe 26°0'0"N 44°0'0"E 46°0'0"E 48°0'0"E 50°0'0"E 52°0'0"E 54°0'0"E 56°0'0"E 58°0'0"E 60°0'0"E 62°0'0"E

Figure 1 Map of Iran depicting the four ecoregions and eight study sites. The numbers indicate the following study areas: (1) Golestan, (2) Khoshyelagh, (3) Turan, (4) Kavir, (5) Kolahghazi, (6) Dena, (7) Mond, (8) Naiband.

The studied areas represent broad variations of temperature, rainfall, and elevation. The annual mean daily air temperature of the study sites ranged from 13°C to 28°C, and the mean annual precipitation varied from 122 mm to 863 mm. The elevation along the sampling areas varied from 15 m to 2728 m (Table 1).

43 Chapter 3 – Hierarchical Partitioning of Ant Diversity

Ant sampling and sampling design

We used the pitfall trapping method for sampling ants; this method is appropriate for short-term investigations of open habitats and accurately represents epigaeic ant species diversity (Bestelmeyer et al., 2000). The traps were plastic cups, 65 mm in diameter and 100 mm in depth, filled two-thirds with a 1:1 mixture of propylene glycol and water to preserve the captured ants. Pitfall traps were left open for a 72- hour period. Samples were preserved in 96% ethanol, brought to the lab, and sorted to species. With the assistance of Alexander Radchenko and available keys, ants were identified to species when this was achievable, or assigned to morphospecies when they could not be matched to known species (see Appendix S1 in the supporting information). In this paper, we refer to morphospecies as “species”. Voucher specimens are held in the collection of the Institute of Experimental Ecology of University of Ulm (http://www.antbase.net) and in the private ant collection of the corresponding author.

Table 1 Descriptions of the eight study sites with location names, elevation, names of the respective ecoregions according to Olson et al. (2001), mean annual precipitation, and mean annual air temperatures. The climate data were obtained from the Iran Meteorological Organization. NP = national park, WLR = wildlife refuge, PA = protected area. Site numbers indicate numbers on the map (Figure 1).

Site Elevation Precipitation Mean daily air Plot name Ecoregion no. (m a.s.l.) mm year^-1 temp (C°) year^-1 1 Golestan NP 1217 Elburz Range forest steppe 185.3 13.1 2 Khoshyelagh WLR 1580 Elburz Range forest steppe 165.4 12.5 3 Turan NP 1190 Central Persian desert basins 126.9 16 4 Kavir NP 1073 Central Persian desert basins 122.3 18.4 5 Kolahghazi NP 1724 Zagros Mountains forest steppe 122.8 16.2 6 Dena PA 2728 Zagros Mountains forest steppe 862.9 15.2 7 Mond PA 17 South Iran Nubo-Sindian desert 241.6 27.1 8 Naiband PA 15 South Iran Nubo-Sindian desert 236.5 27.9

We partitioned ant sampling into four nested spatial levels. We held the extent of our study constant, but the size of grain was increased across levels by aggregating samples of the lower level at a higher level. At each study site we randomly set up two pitfall trap transects, at least 1000 m and at most 2000 m apart, in random directions. We determined the geographical positions of sampling transects by GPS (Garmin eTrex®). Each transect comprised 30 pitfall traps spaced at 10 m intervals. We considered each transect to be the “local” grain at the lowest level (local level) of our hierarchical design, the two transects of one site to be the “landscape” grain, and

44 Chapter 3 – Hierarchical Partitioning of Ant Diversity the two landscapes of one ecoregion to be the “ecoregional” grain. The whole extent of the study was considered the “whole-region” grain. With this sampling arrangement, we sampled each ecoregion with two sample sites (replications). Inter- site distances between replications within each ecoregion ranged from 101 km to 385 km, proportionate to the size of our vast ecoregions. In addition, we chose study sites that represented the typical ecological features and vegetation zones of their ecoregions.

Data analysis

We used the Mao Tao function of ESTIMATES 7.5.2 software (Colwell, 2005), a moment-based species richness estimator, for constructing species accumulation curves. We applied these curves for measuring the sampling effort.

In order to test whether our hierarchical partitioning model showed spatial autocorrelation for observed species richness or Shannon diversity, we produced connectivity matrices based on the transects’ position coordinates by using SAM v.3 software (Rangel et al., 2006) as follows. At the local level, we connected two transects within a landscape. At the landscape level, we connected each transect to the other three transects within an ecoregion. At the ecoregional level, we connected each transect within an ecoregion to transects of the other three ecoregions. These three connectivity matrices mirrored the way in which transects were used at these levels of diversity partitioning to estimate α and β components at that level. SAM v.3 software was also used to calculate Moran’s I values and their significant tests for three connectivity matrices of our hierarchy model.

Crist et al. (2003) showed the way that species richness and diversity can be partitioned across hierarchical scales. In addition, they introduced a computing program, PARTITION, and provided null hypotheses that can be tested to explore significant deviations of observed alpha and beta components of diversity from those expected by chance. We applied program PARTITION v.3 (Veech & Crist, 2009b) to partition species richness and diversity across hierarchical scales in our study and compared the results with those of a random distribution.

45 Chapter 3 – Hierarchical Partitioning of Ant Diversity

We partitioned species richness and Shannon diversity in multiplicative mode, which is appropriate for conservation studies (Jost et al., 2010). In this mode, the beta diversity as measured by species richness and Shannon diversity can be calculated by β = γ/α, if we know the total diversity of a region and average diversity within local species pools (Jost, 2007; Veech & Crist, 2009a). In our study, beta diversity could range from 1, when two species pools had identical species composition, to 2, when two species pools had no overlap in species composition. However, for the highest scale of the whole region, among ecoregions, the beta of species richness and Shannon diversity could range from 1 to 4, because we had four species pools at this level. We employed Shannon diversity, which is the exponential of Shannon entropy (the traditional Shannon-Wiener index) and is the true diversity, measured as the effective number of species, the number of species in an assemblage with an evenness of 1.0 (Jost, 2006). In order to test our predictions, we performed 10,000 randomizations to produce null model distributions for each component of species richness and Shannon diversity across all scales. We first partitioned the total observed species richness and diversity to its components across all scales and ran a randomization for the whole region. Second, we partitioned the observed species richness and diversity of each ecoregion separately. We applied separate randomizations for each ecoregion and used individual randomization tests to explore whether a significant departure of α and β components of our study occurred from a random distribution of individuals among samples. The observed value of each component was compared with the expected value obtained from the null distribution generated by the randomization procedure. We assessed statistical significance by using the proportion of null values that were greater or smaller than the observed number. For instance, if 5 out of 10,000 null values were different from our observed value, then the probability of obtaining (by chance) an estimate greater (or less) than the observed value was 0.0005. We examined the similarity of ant species compositions among the four ecoregions by hierarchical cluster analysis of the species occurrences matrix with the Bray-Curtis similarity index by program PC-ORD (Mccune & Mefford, 2006). We used complete linkage (furthest neighbour), instead of Ward’s method, because of an incompatibility of the latter with non-Euclidean distances (e.g. Bray-Curtis distance) (Mccune & Grace, 2002).

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RESULTS We sampled a total of 12,863 specimens of ants comprising 69 species, corresponding to 1844 species occurrences from the eight sampling sites. We recorded 24 species (confidence intervals (CI): 95% LB: 21 & 95%UB: 26) from the Elburz steppe, 17 species (95% LB: 17 & 95% UB: 17) from the Central desert, 24 species (95% LB: 23 & 95% UB: 25) from the Zagros steppe, and 24 species (95% LB: 18 & 95% UB: 30) from the Nubo steppe. The Central desert had significantly fewer species than the other three ecoregions. Our richness observation curves showed that we had sampled a large fraction of the true number of species present in each ecoregion and in the total region (Figure 2). Nevertheless, our sampling coverage might be biased by not capturing possible cryptic or litter species, especially in steppe ecoregions, as pitfall trapping is not a suitable method for sampling cryptic and litter ants (Bestelmeyer et al., 2000). As a result, the list of 69 collected species should not be considered as the total species richness of the studied region.

25

20

15

10 Species number

5 Elburz Central desert Zagros Nubo 0 0 100 200 300 400 500 Number of species occurrences

Figure 2 Sample-based accumulation curves of the observed ant species richness for the four studied ecoregions.

47 Chapter 3 – Hierarchical Partitioning of Ant Diversity

Spatial autocorrelation The proportion of Moran’s I for observed species richness was positive and insignificant at the local level (Moran’s I = 0.211, p = 0.637) but negative and insignificant at the two other levels: landscape level (Moran’s I = - 0.02, p = 0.959) and ecoregional level (Moran’s I = - 0.097, p = 0.892). Moran’s I values for Shannon diversity were negative and insignificant for all three levels: local level (Moran’s I = - 0.236, p = 0.757), landscape level (Moran’s I = - 0.077, p = 0.989), and ecoregional level (Moran’s I = - 0.051, p = 0.978). This implies that species richness and Shannon diversity of ants were not spatially autocorrelated along the spatial levels, indicating high heterogeneity of ant species composition along the transects. Whole region diversity partitioning For the whole region, hierarchical partitioning suggested that, at the two highest levels, landscape and ecoregional, an under-prediction of beta diversity was a dominant trend. At the local level, however, only Shannon diversity showed such a trend, and the difference in species richness between transects (β local) was insignificant.

The highest proportion of β-species richness (β ecoregional = 3.14) and of β-Shannon (β ecoregional = 2.88) was observed among ecoregions. The lowest values of β-species richness and β-Shannon were found between transects within landscapes, viz., 1.26 and 1.18, respectively (Table 2), with only Shannon diversity being significantly higher than expected. An over-prediction of alpha species richness and diversity was a consistent trend. Species richness within transects (α local = 10.38) was also lower than expected. When diversity was expressed as Shannon diversity, the proportion of α- diversity (α local) was 7.33. Ecoregion diversity partitioning Partitioning of total diversity across four ecoregions into its components, alpha and beta, revealed that most of the diversity components deviated from a null distribution (Table 2). In the Elburz steppe ecoregion half of the species richness (12 out of 24 species) occurred at the alpha level. For the Central desert steppe ecoregion the largest fraction of species richness was observed at α local (11 out of 17 species), and the turnover of species among transects was low and insignificant (Table 2). In contrast, in the Zagros steppe and Nubo desert ecoregions, more than half of the species richness was generated by the turnover of species among transects and among landscapes. In the Zagros steppe, β landscape showed a complete change in species

48 Chapter 3 – Hierarchical Partitioning of Ant Diversity composition at this level. Alpha Shannon diversity within transects for the four ecoregions ranged from 8.20 (Elburz) to 6.48 (Nubo). The greatest β-Shannon occurred at the highest scale, β landscape, in all ecoregions. The lowest values of species richness and Shannon diversity were found at β local in all four ecoregions (Table 2).

Table 2 Hierarchical multiplicative partitioning of α and β components for species richness and Shannon diversity. The P values were obtained by comparing the observed values with the distribution of values from 10000 randomizations with PARTITION v.3.

Species richness Shannon diversity Diversity Observed Expected P Observed Expected P Whole region α local 10.38 39.62 < 0.0001 7.33 29.8 < 0.0001 β local 1.26 1.26 0.565 1.18 1.14 < 0.0001 β landscape 1.68 1.16 < 0.0001 1.56 1.08 < 0.0001 β ecoregion 3.14 1.19 < 0.0001 2.88 1.06 < 0.0001 γ 69 69 38.93 Ecoregions Elburz α local 12 18.75 < 0.0001 8.2 12.12 < 0.0001 β local 1.29 1.15 < 0.0001 1.17 1.05 < 0.0001 β landscape 1.55 1.11 < 0.0001 1.37 1.03 < 0.0001 γ 24 13.14 Zagros α local 9.75 20.95 < 0.0001 7.11 15.64 < 0.0001 β local 1.23 1.09 < 0.0001 1.22 1.06 < 0.0001 β landscape 2 1.05 < 0.0001 1.98 1.03 < 0.0001 γ 24 17.17 Central desert α local 10.5 15.43 < 0.0001 7.48 10.9 < 0.0001 β local 1.1 1.08 0.146 1.08 1.03 < 0.0001 β landscape 1.47 1.02 < 0.0001 1.4 1.01 < 0.0001 γ 17 11.31 Nubo α local 9.5 17.93 < 0.0001 6.48 12.7 < 0.0001 β local 1.47 1.16 < 0.0001 1.32 1.07 < 0.0001 β landscape 1.71 1.15 < 0.0001 1.64 1.04 < 0.0001 γ 24 14.03

Ecoregion classification Cluster analysis identified two main clusters of ecoregions at the first step. The first cluster included the Elburz, Central desert, and Zagros ecoregions (Figure 3). The second cluster contained only the sites of the Nubo ecoregion. The pattern within the first cluster suggested the existence of three major groups. The Elburz and Central desert ecoregions separated into two well-defined groups, but transects of the Zagros ecoregions were divided into two subsets, the northern site clustering with the Central desert sites (Figure 3).

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Information Remaining (%) 100 75 50 25 0 ElburzA ElburzB ElburzC ElburzD CentralA CentralB CentralC CentralD ZagrosA ZagrosB ZagrosC ZagrosD NuboA NuboB NuboC NuboD

Figure 3 Hierarchical cluster analysis for 16 transects among four ecoregions of Iran. The four transects within each ecoregion were named by A, B, C, and D according to their arrangement along the N-S transect. The Bray-Curtis similarity coefficient was used as a measure for species turnover among the plots.

DISCUSSION The significant deviation of most diversity components from their expected values shows that ants are distributed non-randomly among and within scales in the present study. We have found that ecoregions are an appropriate scale for ant conservation, since the greatest variation in species composition of ants occurs at this level. Our findings add to a growing body of literature from insect community ecology suggesting that insect species richness is explained largely by species turnover among broad spatial grains (Summerville et al., 2003). Whole region diversity partitioning At the “whole-region” level, our fundamental finding is that more than 50 percent of total species richness and Shannon diversity of ants was produced by the turnover of species among ecoregions, indicating that the ecoregional scale plays a crucial role in structuring species composition of ground-dwelling ants in arid and semi-arid ecoregions of Iran. This result is consistent with the few comparable hierarchical partitioning studies that have been conducted on other insect taxa, viz. Coleoptera and Lepidoptera in deciduous forests (Gering et al., 2003; Summerville et al., 2003).

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Although the WWF ecoregions have been delineated based on species distribution (Wikramanayake et al., 2002), in our study region they also differ considerably in environmental conditions, vegetation structure, and climate history (Zohary, 1973; Breckle, 1983). Therefore, we suggest that broad-scale abiotic parameters that shape biogeographical history and contemporary environmental conditions are possibly responsible for this large turnover of ants among ecoregions. This view has been supported by detailed regression analyses of environmental parameters and ant community patterns (Paknia & Pfeiffer, 2010). Unlike species richness, which is sensitive to rare species, Shannon diversity weights all species by their frequency (Jost, 2007) and therefore reflects the “evenness” of a community (see Veech & Crist, 2009a). Thus, the highest values of β-species richness between the four ecoregions can be interpreted as the highest turnover of species composition, regardless of their frequency at this scale. The high value of β-Shannon diversity can be explained as the large changes in composition and abundances of species at the ecoregional level recorded in our study. The turnover of species among landscapes was also large and significantly higher than expected, indicating that beta species richness among landscapes also contributes significantly to the overall ant species richness in our studied region. The significant contribution of the landscape level to overall species richness suggests that our sampling replications within each ecoregion detected significant variation in species composition of ants in the ecoregions (e.g. in the Zagros ecoregion with the complete turnover of species composition). Thus shows our sampling design of two replications was sufficient for our study. The observed hierarchical pattern at the landscape and ecoregional levels is unlikely to be changed by additional sampling within ecoregions, because additional sampling would probably increase the β-species richness of the landscape level and, consequently, the β-species richness of the ecoregional level, as newly discovered species will most likely be rare species with restricted distributions and habitats within their specific ecoregions. Ecoregion diversity partitioning As we had expected, despite the overall similarities in the observed patterns of partitioned diversity in the four studied ecoregions, and despite the equal number of observed species for three ecoregions, the contribution of α and β components to the total species richness and diversity was different among ecoregions. These results suggest that different underlying mechanisms probably structure each of these

51 Chapter 3 – Hierarchical Partitioning of Ant Diversity assemblages. Ant species composition within an ecoregion can be structured by both abiotic and biotic factors. If localities of an ecoregion differ greatly enough in their habitat and climate conditions, they are likely to have large differences in their local ant community (Veech & Crist, 2007). In a scenario in which abiotic factors play a major role, probably only a few generalists can adapt to all or most of the localities, but a large number of species are restricted to specific localities, resulting in high values of beta diversity. This is a potential explanation for the observed pattern in the Zagros ecoregion, because the two landscapes of this ecoregion differed largely in habitat and climate (e.g., elevation difference: approx. 1000 m; precipitation difference: approx. 700 mm year^-1). A biotic explanation might highlight species dispersal and interaction within local communities. In such a scenario, the structure of local communities within an ecoregion may depend mainly on the existence or lack of co-dominant species among localities. Co-dominant species within local communities can structure the regional communities in a monotonous mode that reduces β-diversity (Cadotte, 2006). The observed patterns within the four ecoregions in our study are unlikely to be a result of abiotic or biotic influences in isolation, but a combination of the two. The Central desert ecoregion, with a large number of species and a great amount of diversity at the alpha level, had a relatively homogeneous ant species pool with a substantial number of species in common, in spite of large separation between landscapes. This pattern could be explained by the similar climate conditions of sampling localities within this ecoregion but also, probably, by the similar dominant or generalist species that they have in common. In this ecoregion, Monomorium indicum kusnezovi SANTSCHI, 1928 was a super-dominant species and occurred in 100% of traps. The relative influence of these factors remains to be tested in future work. Ecoregion classification The ecoregions of Southwest Asia, including Iran, were adapted from Zohary’s (1973) phytogeographical map of the region (Olson et al., 2001). Interestingly, cluster analysis separates the ant composition of the Nubo ecoregion sample sites well from the other three ecoregions. This pattern mirrors the documented phytogeographical pattern of our studied area, as the Nubo ecoregion harbours remnants of the Sudanian phytogeographical region in Iran and differs from the other three ecoregions, which belong to the Irano-Turanian phytogeographical region (Zohary, 1973). As a consequence, we found several North African or pantropical ants only within this

52 Chapter 3 – Hierarchical Partitioning of Ant Diversity ecoregion, such as Cataglyphis ruber (FOREL, 1903), Pachycondyla sennaarensis

(MAYR, 1862), and M. cf. destructor (JERDON, 1851). Cluster analysis does not distinguish the Zagros ecoregion as a homogeneous fauna. This can be explained by the complete turnover of species from the Kolahghazi National Park (See Figure S1a in Supporting Information) to the Dena protected area (see Figure S1b), with a beta diversity of almost 2.0 (Table 2), indicating two distinct ant species pools. On the other hand, the Kolahghazi National Park shared six species with the Central desert ecoregion (Nr. 4 in Figure 1) and clustered with the two landscapes of the Central desert ecoregion (Figure 3). Several possible interpretations can be proposed for this pattern. One explanation is that ants in the Kolahghazi National Park form a transitional species pool, because we have sampled ants close to the border of the Central desert (Figure 1). This is supported by the findings of Bestelmeyer and Wiens (2001) who demonstrated that a zoogeographical transition for ants might not be coincident with the corresponding phytogeographical transition but might be close to it. However, this explanation does not elucidate the large observed dissimilarity among landscapes of the Zagros ecoregion, unless we assume that the complete dissimilarity among the Zagros ant species pools at the landscape level is attributable to the large differences in topography, habitats, and climate heterogeneity at this level. Another explanation is that the currently suggested ecoregional border among the Zagros and Central desert ecoregions is not accurate, because it represents an old floristic border that has possibly changed during the last four decades, not because of land use, but because of changes in climate features, such as temperature or precipitation regimes (see Evans, 2009). A growing number of studies suggests that climate change affects the distribution ranges of biota (for review see Parmesan, 2006). This means that the borders of the WWF ecoregion maps, i.e., the boundaries of the distribution of the biota, might also change. The outcome of this explanation is that the ant composition of the Kolahghazi National Park indeed represents a Central desert ant community. Conservation strategies Our results show that a substantial amount of species richness and diversity is generated by beta diversity among ecoregions. This production of species richness and diversity is the direct consequence of a restricted distribution of many species to their ecoregions. As a result, anyone wanting to conserve the range of ant biodiversity will need to consider all four ecoregions in conservation plans, such as the creation of

53 Chapter 3 – Hierarchical Partitioning of Ant Diversity new parks and protected areas, especially if we consider ants a surrogate taxon reflecting patterns in many other plant and animal groups (Bennett et al., 2009). The different partitioning of species richness within each ecoregion has revealed that there is also significant variation within ecoregions at the local level with the exception of the Central desert (Table 2). This is indicated by the significant β species richness at the local level of ecoregions, in apparent contrast to the insignificant values among locales for the partitioning of the whole region (Table 2). This explicitly means that the turnover of species among locales within a given ecoregion significantly contributes to the total diversity in that ecoregion. As a result, the turnover of species among locales should also be taken into account in conservation planning. In addition, this outcome suggests that hierarchical partitioning should not be restricted to a whole-region scale, and that regional or geographical subsets should also be considered for further investigation. The observed patterns among the four ecoregions in our study strongly indicate that different conservation strategies are needed for the different ecoregions. For example, the depicted pattern in the Central desert ecoregion suggests that single areas can represent much of ant diversity. On the other hand, the observed patterns in the other ecoregions imply that conservation efforts should consider representing a range of landscapes. Although conservationists usually place most emphasis on “species richness” in their conservation plans, species diversity, which involves species frequencies, includes additional information that might be useful for diversity conservation. However, use of the wrong species diversity measures can lead to unalterable conservation mistakes. Shannon diversity has been suggested as a good abundance-based measure of diversity for conservation purposes (Jost et al. 2010). Use of this measure, together with species richness, gives a more informative picture of the studied diversity pattern. For example, in the Central desert ecoregion in the present investigation, beta species richness was not significant at the local level, but beta Shannon diversity was significantly higher than expected, indicating that occurrences of species fluctuate significantly between locales; however, the species composition does not. In this study, we have not directly inspected the validity of the WWF ecoregion map in Iran and for ants, but the largest turnover in species and in abundances of species at this level indicates that ecoregions correctly represent the biogeographic components of ants in Iran. However, as has been mentioned by ecologists (Magnusson, 2004;

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Whittaker et al., 2005), the validity of boundaries should be considered as a subject for further tests of their robustness (Williams et al., 1999). Our results from the ecoregional classification suggest that the natural turnover of ant species distribution from the Zagros ecoregion to the Central desert ecoregion differs from the delineated ecoregional boundary among these two ecoregions. Although we do not believe our sampling is sufficient for a concrete conclusion, we wish to draw the attention of conservationists to two points for further studies: 1) the coincidence of phytogeographical and zoogeographical boundaries should be evaluated in Iran and elsewhere, and 2) the influence of climate changes on ecoregional boundaries should be studied and anticipated to improve future conservation strategies (Araujo & Rahbek, 2006; Thomas, 2010).

ACKNOWLEDGMENTS The support of Elisabeth K.V. Kalko (Head of the Experimental Ecology Institute, Ulm University) made this work possible. Lou Jost and Jakob Fahr helped us in various ways to shape our ideas. We thank Alexander Radchenko for extensive help and advice with ant species identification. We thank associate editor and three anonymous reviewers for reading the manuscript and providing useful and constructive suggestions. We are most grateful to John Fellowes for his valuable comments and for language correction of the text. The research permits to work in natural reserves were kindly granted by the Department of the Environment of Iran. O.P. is a fellow of the German Academic Exchange Service (DAAD).

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Andersen, A.N. (1997a) Measuring invertebrate biodiversity: Surrogates of ant species richness in the Australian seasonal tropics. Memoirs of the Museum of Victoria, 56, 355-359 Andersen, A.N. (1997b) Using ants as bioindicators: Multiscale issues in ant community ecology. Ecology and Society, 1, 8, published at http://www.consecol.org/vol1/iss1/art8/ Andersen, A.N., Hoffmann, B.D., Müller, W.J. & Griffiths, A.D. (2002) Using ants as bioindicators in land management: simplifying assessment of ant community responses. Journal of Applied Ecology, 39, 8-17 Araujo, M.B. & Rahbek, C. (2006) How does climate change affect biodiversity? Science, 313, 1396-1397 Bailey, R.G. (1998) Ecoregions: the ecosystem geography of the oceans and continents. Springer Verlag, New York. Bennett, J., Kutt, A., Johnson, C. & Robson, S. (2009) Ants as indicators for vertebrate fauna at a local scale: an assessment of cross-taxa surrogacy in a disturbed matrix. Biodiversity and Conservation, 18, 3407-3419 Bestelmeyer, B.T., Agosti, D., Alonso, L.E., Brandao, C.R.F., Brown, W.L., Delabie, J.H.C. & Silvestre, R. (2000) Field techniques for study of ground-dwelling ants. Ants: Standard Methods for Measuring and Monitoring Biodiversity (ed. by D. Agosti, J.D. Majer, L.E. Alonso and T.R. Schultz), pp. 122-144. Smithson Institution Press, Washington, London. Bestelmeyer, B.T. & Wiens, J.A. (2001) Local and regional-scale responses of ant diversity to a semiarid biome transition. Ecography, 24, 381-392 Breckle, S.W. (1983) Temperate deserts and semi-deserts of Afghanistan and Iran. Temperate deserts and semi-deserts. Ecosystems of the world (ed. by N.E. West), pp. 271-319. Elsevier, Amsterdam. Brühl, C. & Eltz, T. (2010) Fuelling the biodiversity crisis: species loss of ground- dwelling forest ants in oil palm plantations in Sabah, Malaysia (Borneo). Biodiversity and Conservation, 19, 519-529 Cadotte, M.W. (2006) Dispersal and species diversity: A meta-analysis. The American Naturalist, 167, 913-924 Colwell, R.K. (2005) EstimateS: Statistical estimation of species richness and shared species from samples. Version 7.5 User's Guide and application published at: http:purl.oclc.org/estimates. Crist, T.O., Veech, J.A., Gering, J.C. & Summerville, K.S. (2003) Partitioning species diversity across landscapes and regions: a hierarchical analysis of α, β, and γ diversity. The American Naturalist, 162, 734-743 Dahms, H., Wellstein, C., Wolters, V. & Dauber, J. (2005) Effects of managment practices on ant species richness and community composition in grasslands (Hymenoptera: Formicidae). Myrmecological News, 7, 9-16 Devictor, V. & Robert, A. (2009) Measuring community responses to large-scale disturbance in conservation biogeography. Diversity and Distributions, 15, 122-130 Evans, J. (2009) 21st century climate change in the Middle East. Climatic Change, 92, 417-432 Fahr, J. & Kalko, E.K.V. (2010) Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales. Ecography, doi: 10.1111/j.1600-0587.2010.05510.x.

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Foxcroft, L.C., Richardson, D.M., Rouget, M. & Macfadyen, S. (2009) Patterns of alien plant distribution at multiple spatial scales in a large national park: implications for ecology, management and monitoring. Diversity and Distributions, 15, 367-378 Gering, J.C., Crist, T.O. & Veech, J.A. (2003) Additive partitioning of species diversity across multiple spatial scales: Implications for regional conservation of biodiversity. Conservation Biology, 17, 488-499 Jost, L. (2006) Entropy and diversity. Oikos, 113, 363-375 Jost, L. (2007) Partitioning diversity into independent alpha and beta component. Ecology 88, 2427-2439 Jost, L., Devries, P., Walla, T., Greeney, H., Chao, A. & Ricotta, C. (2010) Partitioning diversity for conservation analyses. Diversity and Distributions, 16, 65-76 King, J.R., Andersen, A.N. & Cutter, A.D. (1998) Ants as bioindicators of habitat disturbance: validation of the functional group model for Australia's humid tropics. Biodiversity and Conservation, 7, 1627-1638 Lande, R. (1996) Statistics and partitioning of species diversity and similarity among multiple communities. Oikos, 76, 5-13 Lindo, Z. & Winchester, N. (2009) Spatial and environmental factors contributing to patterns in arboreal and terrestrial oribatid mite diversity across spatial scales. Oecologia, 160, 817-825 Magnusson, W.E. (2004) Ecoregion as a Pragmatic Tool. Conservation Biology, 18, 4-10 Mccune, B. & Grace, J.B. (2002) Analysis of ecological communities. MjM Software Design, Gleneden Beach. Mccune, B. & Mefford, M.J. (2006) PC-ORD. Multivariate Analysis of Ecological Data. Version 5.17. In. MjM Software, Gleneden Beach, Oregan, U.S.A Noda, T. (2004) Spatial hierarchical approach in community ecology: a way beyond high context-dependency and low predictability in local phenomena. Population Ecology, 46, 105-117 Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D'amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. Bioscience, 51, 933-938 Paknia, O. (2006) Distribution of the introduced ponerine ant Pachycondyla sennaarensis (Hymenoptera: Formicidae) in Iran. Myrmecological News, 8, 235-238 Paknia, O. & Pfeiffer, M. (2010) Determinations of ant community composition along a 1200 km latitudinal gradient in arid and semi-arid regions of Iran. In: XVI Congress of the International Union for the Study of Social Insects (eds. D. Nash, R., S.P.A. Den Boer, H.H. De Fine Licht and J.J. Boomsma), Copenhagen, Denmark available at http://www.iussi.org/iussi2010/ Parmesan, C. (2006) Ecological and evolutionary responses to recent climate change. Annual Review of Ecology, Evolution, and Systematics, 37, 637 Rangel, T.F.L.V.B., Diniz-Filho, J.A.F. & Bini, L.M. (2006) Towards an integrated computational tool for spatial analysis in macroecology and biogeography. Global Ecology and Biogeography, 15, 321-327 Ribeiro, D.B., Prado, P.I., Brown Jr, K.S. & Freitas, A.V.L. (2008) Additive partitioning of butterfly diversity in a fragmented landscape: importance of

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scale and implications for conservation. Diversity and Distributions, 14, 961- 968 Rodrigues, A.S.L. & Brooks, T.M. (2007) Shortcuts for biodiversity conservation planning: The effectiveness of surrogates. Annual Review of Ecology, Evolution, and Systematics, 38, 713-737 Seppä, P. (2008) Do ants (Hymenoptera: Formicidae) need conservation and does ant conservation need genetics? Myrmecological News, 11, 161-172 Summerville, K.S., Boulware, M.J., Veech, J.A. & Crist, T.O. (2003) Spatial variation in species diversity and composition of forest Lepidoptera in eastern deciduous forests of North America. Conservation Biology, 17, 1045-1057 Thomas, C.D. (2010) Climate, climate change and range boundaries. Diversity and Distributions, 16, 488-495 Tsutsui, N.D. & Suarez, A.V. (2003) The colony structure and population biology of invasive ants. Conservation Biology, 17, 48-58 Tylianakis, J.M., Klein, A.-M., Lozada, T. & Tscharntke, T. (2006) Spatial scale of observation affects α, β and γ diversity of cavity-nesting bees and wasps across a tropical land-use gradient. Journal of Biogeography, 33, 1295-1304 Underwood, E.C. & Fisher, B.L. (2006) The role of ants in conservation monitoring: If, when, and how. Biological Conservation, 132, 166-182 Veech, J.A. & Crist, T.O. (2007) Habitat and climate heterogeneity maintain beta- diversity of birds among landscapes within ecoregions. Global Ecology and Biogeography, 16, 650-656 Veech, J.A. & Crist, T.O. (2009a) PARTITION 3.0 user’s manual. (unpublished document). Veech, J.A. & Crist, T.O. (2009b) PARTITION: software for hierarchical partitioning of species diversity, version 3.0. published at http://www.users.muohio.edu/cristto/partition.htm. Whittaker, R.H. (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs, 30, 279-338 Whittaker, R.H. (1972) Evolution and measurement of species diversity. Taxon, 21, 213-251 Whittaker, R.J., Araujo, M.B., Jepson, P., Ladle, R.J., Watson, J.E.M. & Willis, K.J. (2005) Conservation Biogeography: assessment and prospect. Diversity and Distributions, 11, 3-23 Wikramanayake, E., Dinerstein, E., Loucks, C., Olson, D., Morrison, J., Lamoreux, J., Mcknight, M. & Hedao, P. (2002) Ecoregions in ascendance: reply to Jepson and Whittaker. Conservation Biology, 16, 238-243 Williams, P.H., Klerk, H.M. & Crowe, T.M. (1999) Interpreting biogeographical boundaries among Afrotropical birds: spatial patterns in richness gradients and species replacement. Journal of Biogeography, 26, 459-474 Zohary, M. (1973) Geobotanical foundations of the Middle East. Gustav Fischer Verlag, Stuttgart.

58 Chapter 3 – Hierarchical Partitioning of Ant Diversity

Supporting Information Table S1 List of ant species and their occurrence (+ present, - absent) in the eight studied sites of Iran. Numbers indicate site numbers on the map (Figure 1).

Ant species 1 2 3 4 5 6 7 8 Aphaenogaster gibbosa (LATREILLE, 1798) + ------Camponotus aethiops (LATREILLE, 1798) - - - - - + - - Camponotus fedchenkoi MAYR, 1877 ++------Camponotus gestroi EMERY, 1878 - - - - - + - - Camponotus interjectus MAYR, 1877 + + ------Camponotus oasium FOREL, 1890 - - - - - + + - Camponotus staryi PISARSKI, 1971 - - - - - + - - Camponotus turkestanicus EMERY, 1877 - - + - - - - - Cardiocondyla sp. 1 + ------Cardiocondyla sp. 2 - - + - - - - - Cardiocondyla sp. 3 - - - + - - - - Cataglyphis cf. foreli (RUZSKY, 1903) - - + - - - - - Cataglyphis cinnamomeus (KARAVAIEV, 1910) - - + + - - - - Cataglyphis cugiai MENOZZI, 1939 + + ------Cataglyphis emeryi KARAVAIEV, 1911 - - + - - - - - Cataglyphis frigidus persicus (EMERY, 1906) - - - - - + - - Cataglyphis isis (FOREL, 1913) ------+ - Cataglyphis kurdestanicus PISARSKI, 1965 - - - - - + - - Cataglyphis lividus (ANDRÉ, 1881) - - - + - - + - Cataglyphis niger (ANDRÉ, 1881) - - - + + - - - Cataglyphis nodus BRULLÉ, 1833 + - + - - - - + Cataglyphis ruber (FOREL, 1903) ------+ + Cataglyphis stigmatus RADCHENKO & PAKNIA ------+ + Crematogaster sp. 1 + + ------Crematogaster sp. 2 - + - - - + - - Crematogaster sp. 3 - - - - + - - - Crematogaster sp. 4 - - - - - + - - Crematogaster sp. 5 ------+ Lasius cf. alienus (FOERSTER, 1850) +------Lepisiota frauenfeldi (MAYR, 1855) ------+ Lepisiota semenovi (RUZSKY, 1905) - + + + + - + - Lepisiota sp. 3 - + ------Lepisiota sp. 4 ------+ Messor cf. caducus (VICTOR, 1839) ------+ - Messor cf. striaticeps (ANDRÉ, 1883) ------+ - Messor denticulatus SANTSCHI, 1927 + ------Messor intermedius SANTSCHI, 1927 - - + + + - - - Messor pratennatus ARNOLDI, 1970 - + ------Messor semirufus EMERY, 1925 - - - - + - - - Messor turcmenochorassanicus ARNOLDI, 1977 - - - - - + - - Messor subgracilinodis ARNOLDI, 1970 - - + + + - - - Messor variabilis KUZNETSOV-UGAMSKY, 1927 + - - + - - - - Messor sp. 10 ------+ Monomorium cf. destructor (JERDON, 1851) ------+ Monomorium dentigerum (ROGER, 1862) ------+ - Monomorium indicum kusnezovi SANTSCHI, 1928 - - + + + - + + Monomorium sp. 4 - - - - + - - - Monomorium sp. 5 ------+ - Monomorium sp. 6 ------+ - Monomorium sp. 7 ------+ Pachycondyla sennaarensis (MAYR, 1862) ------+

59 Chapter 3 – Hierarchical Partitioning of Ant Diversity

Paratrechina sp. 1 ------+ Pheidole sp. 1 - + ------Pheidole sp. 2 - - - - + - - - Pheidole sp. 3 - - - - - + + - Plagiolepis sp. 1 + ------Plagiolepis sp. 2 - + ------Plagiolepis sp. 3 - - - + + - - - Plagiolepis sp. 4 - - - - - + - - Plagiolepis sp. 5 - - - - - + - - Proformica epinotalis KUSNETSOV-UGAMSKY, 1927 + + ------Tapinoma erraticum (LATREILLE, 1798) + + ------Tapinoma sp. 2 ------+ + Temnothorax semenovi (RUZSKY, 1903) + - - - - + - - Tetramorium armatum SANTSCHI, 1927 + + + - - - - - Tetramorium schneideri EMERY, 1898 + ------Tetramorium striaventre MAYR, 1877 - + + + - - - - Tetramorium sp. 5 - - - - - + - - Tetramorium sp. 6 ------+ -

60 Chapter 3 – Hierarchical Partitioning of Ant Diversity

(a)

(b)

Figure S1 Kolahghazi National Park (a) and Dena protected area (b) in the Zagros Mountains forest steppe ecoregion. The first area is a xeric shrubland, unlike the second area, which is a sparse oak forest.

61 Chapter 3 – Hierarchical Partitioning of Ant Diversity

62

4 Environmental and spatial drivers of ant species composition in a dryland ecosystem in Iran

Omid Paknia and Martin Pfeiffer In preparation for publication

ABSTRACT

Clarification of the mechanisms that cause diversity in communities is a key task in current community ecology. Niche differentiation, dispersal limitation, and stochastic processes have been suggested as the main sources of biodiversity in communities. In this study we have quantified the variation in ant assemblages relating to environmental and spatial factors. Our main hypothesis, as we explored a long transect across harsh arid and semi-arid heterogeneous environments, was that variation in species composition is largely explained by environmental and spatial parameters. Using pitfall trap methods for sampling ants, along a north-south transect of ca. 1200 km through arid and semi-arid areas of Iran, we applied canonical analysis to disentangle the relative importance of environmental and spatial processes. We used principal coordinates of neighbor matrices (PCNM) to model spatial processes. Our parsimonious environmental and spatial models jointly explain 62% of the variation in the community composition of ants. Community composition is dominantly controlled by the spatial-dependent environment component (39%), including the variables annual rainfall, summer rainfall, and temperature range. The Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages environment-independent spatial component explains 23% of total variation. A large number of species are highly localized. The hypothesis that ant species composition is influenced largely by environmental and spatial variables is supported by our variation partitioning analysis. However, the significant contribution of spatial processes suggests that ant community composition is also influenced by dispersal, especially over short distances when environmental conditions are suitable.

Keywords: beta diversity, contemporary climate, dispersal, PCNM analysis, variation partitioning, Iran.

INTRODUCTION

Over the years, most efforts in community ecology have focused on alpha diversity and its causes. As a result, our current ecological knowledge about the causes of alpha diversity is superior to that known about beta diversity (e.g. Hawkins et al., 2003). Beta diversity and the factors that contribute to it have crucial functions in community ecology theories such as the regional vs. local control of diversity (Ricklefs, 1987), and key implications for proper conservation management (Jost et al., 2010). The main currently postulated hypotheses about the origin of beta diversity suggest that a number of biological, ecological, and stochastic processes drive it. A large body of studies, mostly conducted in environmentally heterogeneous habitats, has demonstrated that the variation in species composition is a niche-based process and is strongly influenced by environmental conditions (e.g. Bray & Curtis, 1957). Alternatively, variation in species composition has been suggested to be a result of neutral and stochastic processes. This hypothesis gives a particular role to dispersal, whereas individuals of all species are considered equal in their other abilities (Hubbell, 2001). The main conclusion of this hypothesis is a distance decay of similarity patterns or a positive autocorrelation among local assemblages because of dispersal limitation.

Despite the recent growing body of research devoted to understanding its origins, beta diversity remains the subject of ongoing studies in community ecology, because of the

64 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages scarce documentation of beta diversity patterns based on systematic and standardized inventories especially for terrestrial invertebrates at large scales (Vasconcelos et al., 2010). Moreover, there are a small number of studies which have considered the impact of spatial structure on beta diversity (Baselga, 2008), while spatial structuring plays a fundamental role in each studied ecosystem, either as potential nuisance for sampling and statistical testing, or as a functional necessity. Determination and explanation of geographical structure affiliated with ecological communities is an essential issue in contemporary ecology (Borcard & Legendre, 2002; Griffith & Peres-Neto, 2006).

The aim of the present study is to determine the origins of variation among ant assemblages along a latitudinal transect in a dryland ecosystem in Iran. Ants are an ideal taxa for evaluating the ecological impact of environmental and spatial factors, because they are important terrestrial insects, especially in arid regions (Johnson, 2001), and because ant species richness and composition varies widely among regions. Patterns of alpha diversity of ants have been demonstrated at a wide range of scales. These studies show that alpha diversity in ants correlates with climate parameters such as temperature, rainfall, and productivity (e.g. Kaspari et al., 2003; Sanders et al., 2007; Dunn et al., 2009). Nevertheless, there is a surprising lack of knowledge about the impacts of both, the spatial structure and environment, on the patterns of beta diversity of ants, restricted to a few studies which have been carried out in tropical forests. From tests of the variation partitioning among spatial and environmental variables across a long transect in the Amazonian floodplain, Vasconcelos et al. (2010) showed that environmental and spatial heterogeneity is weakly responsible for fluctuations in ant assemblages, and a large fraction of observed variation remained unexplained. Similar results have been obtained by Mezger and Pfeiffer (2010) who have found that environmental parameters have only a limited influence on the assemblage of ants in various types of primary rainforest on Borneo.

In this study, for the first time, we have evaluated the influence of environmental and geographical factors on variations in the assemblages of ants across a long transect in arid and semi-arid regions. We aim to investigate the following hypotheses. First, as we explored a long transect across harsh arid and semi-arid heterogeneous environments, we hypothesize that variations in ant species composition are

65 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages significantly and largely explained by environmental parameters, rather than by spatial parameters or remaining unexplained. Second, in consequence of the environmental heterogeneity and dispersal limitation to short distances, we hypothesize that a large number of species are associated with particular habitats.

METHODS

Study area, study sites, and sampling design

Iran is located in the mid-latitude band of arid and semi-arid regions of the old world and is divided into three main phytogeographical regions (Zohary, 1973). In this study, we sampled ants from two vast phytogeographical regions (Table 1). We sampled ant assemblages during May to July of 2007 along a north-south transect of ca. 1200 km at 16 sites within eight protected area (Figure 1). We captured epigaeic ants by pitfall trapping. In each site, we laid out one transect of pitfall traps. In each transect, we placed 30 pitfall traps with a 10-m interval between traps. All pitfall traps were left open for 3 days. Two sites within each protected areas were at last 1000 m and at most 2000 m apart. For spatial analyses, the position of each transect was recorded from the middle by a Global Position System device (Garmin eTrex®). We determined the species of ants where possible, or assigned them to morphospecies. Voucher specimens of each ant species collected are held in the AntBase.Net Collection (ABNC) of the Institute of Experimental Ecology of University of Ulm (http://www.antbase.net) and in the private ant collection of the corresponding author.

Data analysis

We prepared a species × sample (occurrence) matrix as an assemblage dataset. Because ants are social, and as a whole colony might occur in one sample, we used an occurrence-based data set instead of an abundance-based data set (see Longino et al., 2002). The resulting assemblage data matrix was Hellinger-transformed to express species occurrences as square-root transformed balanced occurrences in each sampling sites (Legendre & Gallagher, 2001). To identify origins of beta diversity, we applied variation partitioning methods suitable for evaluating contributions of spatial

66 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages and environmental processes to species composition (Borcard et al., 1992; Peres-Neto et al., 2006; Legendre, 2008; but see Smith & Lundholm, 2010).

Turkey Turkmenistan 40°0'0"N

Iraq Iran Afghanistan Caspian Sea

1 38°0'0"N Pakistan 2 Saudi Arabia

3 36°0'0"N 4 34°0'0"N

5 32°0'0"N 6 30°0'0"N

7 8 28°0'0"N

0125 250 Kilometers Persian Gulf 26°0'0"N 44°0'0"E 46°0'0"E 48°0'0"E 50°0'0"E 52°0'0"E 54°0'0"E 56°0'0"E 58°0'0"E 60°0'0"E 62°0'0"E

Figure 1 Map of Iran depicting eight study sites. The numbers indicate the following sampling sites: (1) Mirzabailu, (2) Khoshyelagh, (3) Turan, (4) Kavir, (5) Kolahghazi, (6) Dena, (7) Mond, (8) Naiband.

The application of variation partitioning is suitable when both species assemblages and the environment are spatially dependent (Peres-Neto & Legendre, 2010). We used Mantel r and Moran’s I correlograms to inspect the spatial structure of ant assemblages and environmental variables at a range of distances, respectively.

To describe the spatial structure of species assemblages, we used the normalized Mantel statistic, which determines linear relationships between two sets of distance- based data: the species composition matrix and the geographical distance matrix. GPS coordinates of transects were converted to a geographical distance matrix of

Euclidean distances (km) by software SAM 3.1 (Rangel et al., 2006). We used the R language (R Development Core Team, 2007) function “mantel.correlog” in the vegan package (Oksanen et al., 2009) to compute our multivariate Mantel correlogram, with

67 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages the Pearson correlation, and the SAM 3.1 package to calculate values of Moran’s I and to produce correlograms for the selected environmental variables (see below); for both methods we applied the Sturge equation for estimating distance classes, the Bonferroni progressive correction (α = 0.05), and 999 permutations for tests of significance.

Table 1 Descriptions of the eight study sites with location names, names of the respective phytogeographical regions according to Zohary (1973), elevation, mean annual rainfall, mean annual summer rainfall, mean annual air temperature, and mean annual temperature range.

Phytogeographical Rainfall Summer rainfall Mean daily Temp. range Site no. Site name Elevation (m a.s.l.) regions (mm year^-1) mm year^-1) air temp.(C°) (C°) 1 Mirzabailu Irano-Turanian 1217 185.3 21.1 13.1 12.9 2 Khoshyelagh Irano-Turanian 1580 165.4 23.4 12.5 10.6 3 Turan Irano-Turanian 1190 126.9 7.9 16 13.1 4 Kavir Irano-Turanian 1073 122.3 3.3 18.4 14.5 5 Kolah ghazi Irano-Turanian 1724 122.8 2.1 16.2 16.2 6 Dena Irano-Turanian 2728 862.9 3.4 15.2 13.3 7 Mond Nubo-Sindian 17 241.6 0.4 27.1 9.6 8 Naiband Nubo-Sindian 15 236.5 0.2 27.9 10.1

To model variation of species composition of ants along our ecological transect, we used a set of environment variables (see Appendix S1) that described the contemporary climate, habitat productivity, elevation, and habitat heterogeneity. We used the “forward.sel” function of the packfor package (Dray et al., 2007) of R language (R Development Core Team, 2007) to select those environmental variables that contributed significantly to the explanation of ant composition. The advantage of this forward selection function is that two criteria (alpha level and adjusted R2 of full model) are used to stop the variable selection procedure. This procedure returns fewer variables than the full model with a smaller value of adjusted R2. However, these variables are more realistic than the variables of the full model. Moreover, they are corrected for inflated Type I error and for the overestimation of the amount of explained variance, both of which are common problems in the classical forward selection of explanatory variables (Blanchet et al., 2008). Applying redundancy analysis (RDA), we made a global (full model) test of the variables and calculated the adjusted R2 of the global model, which was used as the second criterion in forward selection. Forward selection was applied at the α = 0.05 level and an adjusted R2 of < 0.51 with 999 random permutations. After the variable selection, we inspected the

68 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages collinearity of the selected environment variables with the “vif.cca” function of the “vegan” package. We used only the selected variables in subsequent analyses.

We modeled the spatial structure of ant samples by using principle coordinates of neighbor matrices (PCNM). This method is used to include spatial structure, e.g., spatial autocorrelation, as predictor variables of variation in ecological assemblages in applications of multiple regression or canonical ordination (Borcard & Legendre, 2002). We computed PCNM variables by the “quickPCNM” function of the PCNM package (Legendre et al., 2009) of R language from the species composition matrix and the GPS coordinates of the transects and used the packfor package (α = 0.05, 999 random permutations) to select the best PCNM eigenfunctions by forward selection. The selected PCNMs eigenfunctions were used as explanatory variables in the variation partitioning of the ant species composition data.

We partitioned the proportion of the variation in species composition explained by environmental and spatial variables by RDA. Using the “varpart” function of the vegan package of R language, we calculated the following fractions of total variation of ant composition: total explained variation [E+b+S], environment variation [E], spatial variation [S], the common variation shared by environment and space [b], and unexplained variation [d]. Furthermore, we calculated the fraction of variation in ant composition that can be explained by purely environmental variables [E|S] and the fraction of variation that can be explained purely by spatial variables [S|E]. We used adjusted R2 statistics for variation partitioning as recommended by Peres-Neto et al. (2006). The significances of testable fractions, [E], [S], [E+b+S], [E|S], and [S|E], were tested by means of 999 permutations by the “rda” and “anova.cca” functions in the vegan package.

We used RDA separately for fractions [E] and [S|E] and examined the relationship between ant assemblages and environmental and spatial variables. We extracted the fitted scores for each of the first two canonical axes and mapped them in biplots to visualize the patterns of the species composition in each of these fractions. The extracted fitted scores were also used to run multiple regression analyses, which selected environmental and spatial parameters were used as explanatory variables to check which parameters contributed most to the site scores of these fractions, as judged from their partial coefficients. RDAs were computed by using the vegan

69 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages package. Multiple regression analyses were performed with STATISTICA 8.0 (Statsoft, 2007).

To determine characteristic species of the various sites, we used “indicator species analysis” (Dufrêne & Legendre, 1997) with PC-ORD 5.17 (Mccune & Mefford, 2006). This analysis calculates indicator values for each species by combining the relative abundance of a certain species with its relative frequency of occurrence in the various transects. The estimated values were tested for statistical significance by randomization tests (Monte Carlo, 1000 permutations).

RESULTS

We collected a total of 69 species/morphospecies along the north-south transect. Cataglyphis was the most speciose genus with 12 species, followed by Messor and Monomorium with ten and seven species, respectively. By contrast, we found only one species per genus for Aphaenogaster, Lasius, Paratrechina, Pachycondyla, and Proformica. The most common species were Monomorium indicum kusnezovi

SANTSCHI, 1928 and Cataglyphis niger (ANDRÉ, 1881), which occurred in 30% and 23% of all pitfall traps, respectively. In contrast, five species Aphaenogaster gibbosa

(LATREILLE, 1798), Lasius cf. alienus (FOERSTER, 1850), Messor cf. striaticeps

(ANDRÉ, 1883), Paratrechina sp. 1, and Crematogaster sp. 5 were unique and occurred only in one pitfall trap. Lepisiota semenovi (RUZSKY, 1905) had the widest distribution and occurred at six out of eight sites, followed by Monomorium indicum kusnezovi, which occurred at five sites. Interestingly, 45 species were restricted to one site, indicating that most species were highly localized.

The species composition of ants showed a significant positive spatial autocorrelation at the shortest distance and significant negative spatial autocorrelations at two farther distances (Figure 2), indicating that, similarity decay pattern occurred only at the shortest distance, where the ant assemblages were more similar than they would be by chance. At the farther distances, the ant assemblages were more dissimilar than expected. The causes of this pattern could be either spatial autocorrelation in ant assemblages, attributable to dispersal, or spatial dependence induced by environmental parameters, which themselves were spatially, positively autocorrelated at the shortest distances (Figure 3).

70 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages

0.7 0.6 0.5 0.4 r 0.3 0.2 0.1 Mantel 0.0 -0.1 -0.2 -0.3 0 200 400 600 800 1000 1200 Distance (km)

Figure 2 Mantel correlogram showing spatial correlation (Mantel r) for the ant assemblages. Black circles indicate significant spatial autocorrelation after progressive Bonferroni corrections (α = 0.05, 999 permutations).

Annual rainfall Annual summer rainfall 1.6 Annual temperature range 1.2 0.8

I 0.4 0.0

Moran's Moran's -0.4 -0.8 -1.2 -1.6 0 200 400 600 800 1000 1200 Distance (km)

Figure 3 Correlogram showing spatial correlation (Moran’s I) for three environmental variables entered in the variation partitioning model: annual rainfall, annual summer rainfall and temperature range. Black circles indicate significant spatial autocorrelation after progressive Bonferroni corrections (α = 0.05, 999 permutations).

The forward selection retained three significant environmental variables for the modeling of species composition: annual rainfall (F = 3.15, P < 0.001), summer rainfall (F = 3.67, P < 0.001), and annual temperature range (F = 3.75, P < 0.001). The cumulative adjusted R2 of this selected model was 0.39.

PCNM generation over 16 transects produced nine positive PCNM eigenvalues. Forward selection procedure retained three significant PCNM out of nine positive

71 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages

PCNM vectors: PCNM 3 (P = 0.005), PCNM 5 (P = 0.001), and PCNM 6 (P = 0.029). Adjusted R2 of the selected model was 0.32, which was near to the global model for all PCNMs without any selection, which had an adjusted R2 of 0.34.

Variation partitioning of the ant species composition included three environmental and three spatial (PCNM) explanatory variables. In total, both the spatial and the environmental variables explained 62% of the species composition variation and left 38% unexplained (Table 2). Environmental variables [E] explained 39% of the variation in species composition. Partialling out the negative spatial structure of environmental variables, purely environmental factors, [E|S], explained a larger amount of variation, equal to 45% of species composition (Table 2). Selected PCNM variables, [S], explained 17% of total variation of species composition. Purely spatial variables, [S|E], explained 0.23 of the total variation. All testable fractions were highly significant (all P < 0.003).

Table 2 Partitioning of the variation in ant community composition by using two subsets of data: spatial (PCNMs) and environmental variables. E = environmental, S = spatial, E|S = purely environmental, S|E = purely spatial, b = the common fraction of variation shared by space and environment, and d = undetermined.

Fraction d.f. R 2 Adjusted R2 P value E 3 0.51 0.39 0.001 S 3 0.33 0.17 0.003 E+b+S 6 0.77 0.62 0.001 E|S 3 0.66 0.45 0.001 b 0 - -0.06 Un-testable S|E 3 0.53 0.23 0.001 d - - 0.38 Un-testable

RDA of fraction [E] retained three axes (axis 1: R2 = 0.22, axis 2: R2 = 0.17, and axis 3: R2 = 0.124; all P < 0.001). The first two axes discriminated ant species composition in four groups (Figure 4a). Summer rainfall had the largest partial contribution to axis 1 (R = - 0.97, P < 0.0001), and annual rainfall the largest partial contribution to axis 2 (R = - 0.98, P < 0.0001). Axis 1, which can be interpreted as a summer rainfall gradient, discriminated the two northern steppe sites from other sites. The second axis, which was strongly correlated with annual rainfall, split ant species composition into three assemblages. The first group was the western steppe, which had the highest amount of annual rainfall, 862 mm year-1. The second group was the coastal desert

72 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages with more than 235 mm annual rainfall. The third group was the remaining sites including the northern steppe and the central desert regions, both with lower than 185 mm annual rainfall (Figure 4a).

1.0 a) Northern steppe Central desert

0.5 Kh-B Ka-B Tu-A Ka-A Kh-A Tu-B Ko-A sum e Mi-A mer ng Ko-B Mi-B rainf ra all p. m te 0.0

RDA2 ll nfa rai Coastal desert al nu an Na-A Mo-A -0.5 Na-B Mo-B De-A

Western steppe De-B -1.0 -1.0 -0.5 0.0 0.5 RDA1

1.5 b)

1.0 Mo-B

Mo-A 0.5 Ka-A Mi-A Ka-B p cnm Ko-B 5 m3 Kh-B RDA2 Ko-A n pc 0.0 Mi-B Kh-A De-B De-A 6 Tu-B m n -0.5 c p Tu-A Na-B Na-A -1.0 -1.0 -0.5 0.0 0.5 1.0 RDA1

Figure 4 Redundancy analysis biplots of the first two axes corresponding to a) environmental variation [E] and b) purely spatial variation [S|E]. Abbreviations indicate De: Dena, Ka: Kavir, Kh: Khoshyelagh, Ko: Kolahghazi, Mi: Mirzabailu, Mo: Mond, Na: Naiband and Tu: Turan. The two transects within each ecoregion are shown as A and B.

73 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages

RDA of [S|E] also retained three significant axes (axis 1: R2 = 0.223, axis 2: R2 =0.204, axis 3: R2 = 0.106; all P < 0.002). PCNM 5 and PCNM 6 contributed largely to axis 1 (R = - 0.76, P = 0.0017) and axis 2 (R = - 0.95, P < 0.00001), respectively. Axis 1 separated the Turan site, and axis 2 separated the Nubo and Mond sites (Figure 4b). The indicator analysis determined a large number of species as indicators (57 out of 69 species, all P < 0.01), suggesting strong associations of ants to their habitats. Species with the three highest indicator values for each site are listed in Table 3.

Table 3 Indicator ant species with their indicator values and statistical significance. Only the three species with the highest indicator values for each site have been listed. Abbreviations for sites: MI = Mirzabailu, KH = Khoshyelagh, TU = Turan, KA = Kavir, KO = Kolahghazi, DE = Dena, MO = Mond, NA = Naiband. The means and standard deviations are given together with p-values for the null-hypothesis of no difference between sites (Mccune & Mefford, 2006).

IV from randomized groups Indicator P Species Site value (IV) Mean Standard value Deviation Messor denticulatus SANTSCHI, 1927 MI 77.6 3.1 0.9 0.0001

Tetramorium schneideri EMERY, 1898 MI 58.6 2.7 0.89 0.0002

Tetramorium armatum SANTSCHI, 1927 MI 11.6 2.7 0.9 0.001

Cataglyphis cugiai MENOZZI, 1939 KH 55.8 5 1 0.0002 Crematogaster sp. 1 KH 52.1 4.3 0.98 0.0002

Messor pratennatus ARNOLDI, 1970 KH 39.6 2 0.84 0.0016

Cataglyphis emeryi KARAVAIEV, 1911 TU 83.3 3.3 0.94 0.0002

Cataglyphis cinnamomeus (KARAVAIEV, 1910) TU 69.4 3.7 0.96 0.0002

Cataglyphis cf. foreli (RUZSKY, 1903) TU 48.3 3.3 0.94 0.0002

Cataglyphis lividus (ANDRÉ, 1881) KA 57.7 3.6 0.97 0.0002 Cardiocondyla sp. 3 KA 25 1.8 0.81 0.0002

Lepisiota semenovi (RUZSKY, 1905) KA 19.4 4.1 0.97 0.0002

Messor intermedius SANTSCHI, 1927 KO 53.1 3 0.95 0.0002

Cataglyphis niger (ANDRÉ, 1881) KO 50.1 5.4 1.03 0.0002 Pheidole sp. 2 KO 16.7 1.6 0.85 0.0002

Cataglyphis kurdestanicus PISARSKI, 1965 DE 98.3 3.6 0.96 0.0002

Camponotus aethiops (LATREILLE, 1798) DE 41.7 2.3 0.85 0.0002

Camponotus oasium FOREL, 1890 DE 41.1 3.4 0.92 0.0002

Monomorium dentigerum (ROGER, 1862) MO 82.8 3.2 0.93 0.0002

Cataglyphis ruber (FOREL, 1903) MO 25.4 3.6 0.95 0.0002

Cataglyphis isis (FOREL, 1913) MO 20.7 1.7 0.81 0.0002 Monomorium sp. 7 NA 82.8 3.2 0.93 0.0002 Cataglyphis stigmatus Radchenko&Paknia NA 22.6 2.2 0.87 0.0002

Monomorium cf. destructor (JERDON, 1851) NA 20.7 1.7 0.81 0.0002

74 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages

DISCUSSION

In this study, we have found that the major part of variation among ant species composition in Iranian arid and semi-arid regions is interpretable by spatial and environmental variables. This result is in contrast to two of the most recent studies on tropical forests ants (Mezger & Pfeiffer, 2010; Vasconcelos et al., 2010), which have detected little evidence for the impact of spatial and environmental parameters on the species composition of ants, suggesting that, in tropical forests, the species composition of ants is not mainly governed by niche-based processes.

In our parsimonious model, three spatially structured climate variables, viz., temperature range, annual rainfall, and summer rainfall, explain 39% of the beta diversity of ants. Temperature and rainfall have been named as the two most important parameters that explain geographical variations in alpha diversity for a large number of animal and plant species (Hawkins et al., 2003). In arid and semi-arid areas rainfall is well known as being the dominant controlling factor for biological processes, such as primary production. Moreover, unpredictable variations occur in its amount and occurrence (Noy-Meir, 1973). Ordination analysis of environmental variables in our study showed that ant species in arid areas of Iran also responded to both the amount of yearly rainfall and the patterns of its occurrence (Figure 4a). The RDA ordination discriminated the western steppe assemblage from the remaining sites due to its high amount of annual rainfall. In contrast to other regions, in which the dominant vegetation cover is dwarf scrub plants, this region is characterized by sparse oak forests. As a consequence, two out of three main indicator species in this region are the arboreal species, Camponotus aethiops (LATREILLE, 1798) and

Camponotus oasium FOREL, 1890.

In their long-term experiment in a North American arid grassland, Kaspari and Valone (2002) found that the abundance of granivorous ants was positively influenced by seed production following summer rainfall, but omnivorous ant species were unaffected by rainfall. This is a potential explanation for the discrimination of the ant assemblage of the northern steppe, predicting that the northern steppe should be designated mainly by granivorous species. Results of indicator species supports this interpretation, as a notable number of indicator species at these sites are indeed granivorous ants, such as Messor denticulatus SANTSCHI, 1927, Messor pratennatus

ARNOLDI, 1970, and Tetramorium schneideri EMERY, 1898.

75 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages

In conformity with Pfeiffer et al. (2003), our findings show that the temperature range can explain the beta diversity of ants, as the central desert region is separated from other sites because of its large temperature range. The central desert region has an extreme continental climate with large diurnal and annual temperature fluctuations and cold winters with 30 – 90 days below freezing per year (Breckle, 2002). As ants are characteristically thermophilic, the instability in temperature probably prevents the colonization of certain regions by any ant genera that have dispersed to this region; but they cannot establish their populations, because they possess no adaptations to temperature fluctuations and low minimum temperature. Pfeiffer et al. (2003) have pointed out that low temperature has a significant impact on the community structure of ants in Mongolia. They conclude that the harsh and fluctuating climatic conditions in Mongolia probably act as a major factor against the diversification of certain ant genera. The Iranian central desert region receives a lower amount of annual rainfall than other sites (Figure 4a). As a result, ant species in this region have to cope with low primary productivity, in addition to temperature fluctuations. Interestingly, the indicator species in this habitat, such as Lepisiota semenovi (RUZSKY, 1905) and six Cataglyphis species, are omnivorous and scavengers, which is a common diet strategy in desert ecosystems with low primary productivity (see Rojas & Fragoso, 2000).

The spatial component that is shared between space and environment, fraction [b], reveals the contribution of spatial autocorrelation in environmental processes. By removing this fraction and testing the pure environmental fraction [E|S], it is possible to have control over the spatial structure of the environment variables and as a result to control type I error (Peres-Neto & Legendre, 2010). Our results show that the pure environmental fraction also contributes significantly to explaining the variance of assemblages (Table 2), suggesting that spatial autocorrelation has not affected the impact of environmental variables on species composition. From a statistical point of view, a negative value for fraction [b] indicates that two groups of variables explain the assemblages variation better than the sum of individual effects of these variables (Legendre & Legendre, 1998).

The purely spatial fraction [S|E] mainly demonstrates the “relative contribution” of spatial autocorrelation of ant assemblages in the variation partitioning that is generated by dispersal limitation. The result of the Mantel correlogram, which reveals the “strength” of spatial autocorrelation at different distances (Laliberté, 2008), shows

76 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages a strong positive spatial autocorrelation at the shortest distance. These findings suggest that local assemblages are spatially autocorrelated because of high dispersal to neighboring suitable and similar habitats (Figure 3).

When no access to relevant and extensive environmental data is available, then the “pure spatial” fraction should also be considered as the result of spatially structured unmeasured explanatory variables (Laliberté, 2008). Mapping this fraction by the first two axes of RDA ordination separates three sites, viz., Naiband, Mond, and Turan, from the remaining sites, but in different directions (Figure 4b). Because soil texture varies considerably among sites from sandy to clay soils we suggest that soil texture and soil chemistry are important components of this fraction. For instance, one potential explanation for the separation of the Naiband site from the remaining sites is that it is the sole sand dune area in our study. The Mond site contained clay soil; however, it is the nearest site to Naiband. Several studies have shown that soil texture affects the species composition of ants (Hill et al., 2008). In addition, ant assemblages can also be affected directly or indirectly by the chemistry of the soil, especially by its salt content, as ants use salt as a nutrient or try to avoid it as a parameter of stress (Kaspari et al., 2008). In our case, the central desert region is an inland basin and is characterized by salty conditions and flat playas, which comprise a wide range of salt contents (Breckle, 2002). The Turan site of the central desert is discriminated by axis 1 from other sites, possibly by soil chemistry (Figure 4b).

Of the variation in ant assemblages, 38% remains undetermined. Non-spatially structured environmental variables, such as local disturbance (Hill et al., 2008) and local mosaic-like environmental variations (Pfeiffer et al., 2003), and biological variables, such as intra- and inter-specific competitive interactions (Parr, 2008; Pfeiffer et al., 2008), which have not been measured or considered in this study, might be responsible for this fraction. Another possible explanation for this fraction is “stochastic processes”. In other words, this undetermined portion can be explained as a result of ecological drift, assuming that differences in the species composition of ants are primarily driven by stochasticity in birth, death, and the dispersal of species.

Highly significant values for both pure spatial and pure environment fractions suggest that the ant species composition in arid and semi-arid regions of Iran is controlled by both niche- and dispersal-based processes, which is in agreement with Hubbell’s (2001, page 24) statement that “actual ecological communities are undoubtedly

77 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages governed by both niche-assembly and dispersal- assembly rules along with ecological drift.” Most neutral models predict that species composition changes across space because of the limited dispersal of species. In these models, the limitation in dispersal is linked to species abilities, because the environment is assumed to be homogeneous. However, the regions studied by us are harshly heterogeneous, suggesting that the high variation among communities is attributable to the heterogeneity of the environments. This restricts the dispersion processes of many species to a short vse5brtrw9drvap09">398

Camponotus oasium FOREL, 1890, which is abundant in the western steppe but has been found only in one pitfall trap in the coastal desert.

CONCLUSION

The hypothesis that ant species composition is influenced largely by environmental variables has been supported by our variation partitioning analysis. The relative contribution of environmental components is almost two times larger than that of spatial components suggesting that niche-assembled processes are the prime determinants of the composition of ants in arid and semi-arid areas of Iran. Nevertheless, ant species composition shows significant contribution of spatial structuring, suggesting that dispersal processes continue to influence ant species composition across the studied arid and semi-arid ecoregions.

78 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages

ACKNOWLEDGMENTS We gratefully acknowledge the support of Elisabeth K.V. Kalko. We thank Alexander Radchenko for extensive help and advice with ant species identification and Theresa Jones and Ashwin Chari for professional language correction. We also thank E. Laliberté, F.G. Blanchet, and P.R. Peres-Neto for their helpful comments and hints on our statistical questions, and Dirk Mezger for his comments on early versions of this manuscript. The research permits to work in natural reserves were granted by the Department of the Environment of Iran. O.P. is a fellow of the German Academic Exchange Service (DAAD).

REFERENCES: Baselga, A. (2008) Determinants of species richness, endemism and turnover in European longhorn beetles. Ecography, 31, 263-271 Blanchet, F.G., Legendre, P. & Borcard, D. (2008) Forward selection of explanatory variables. Ecology, 89, 2623-2632 Borcard, D. & Legendre, P. (2002) All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices. Ecological Modelling, 153, 51-68 Borcard, D., Legendre, P. & Drapeau, P. (1992) Partialling out the spatial component of ecological variation. Ecology, 73, 1045-1055 Bray, J.R. & Curtis, J.T. (1957) An ordination of the upland forest communities of Southern Wisconsin. Ecological Monographs, 27, 325-349 Breckle, S.-W. (2002) Salt deserts in Iran and Afghanistan. Sabkha Ecosystems (ed. by H.-J. Barth and B. Boer), pp. 109-122. Kluwer Academic Publishers, Dordrecht. Dray, S., Legendre, P. & Blanchet, F.G. (2007) packfor: Forward selection with permutation, version 0.0-7. Available at: http://cran.r-project.org. Dufrêne, M. & Legendre, P. (1997) Species assemblages and indicator species: The need for a flexible asymmetrical approach. Ecological Monographs, 67, 345- 366 Dunn, R.R., Agosti, D., Andersen, A.N., Arnan, X., Bruhl, C.A., Cerd, X., Ellison, A.M., Fisher, B.L., Fitzpatrick, M.C., Gibb, H., Gotelli, N.J., Gove, A.D., Guenard, B., Janda, M., Kaspari, M., Laurent, E.J., Lessard, J.-P., Longino, J.T., Majer, J.D., Menke, S.B., P., M.T., Parr, C.L., Philpott, S.M., Pfeiffer, M., Retana, J., Suarez, A.V., Vasconcelos, H.L., Weiser, M.D. & Sanders, N.J. (2009) Climatic drivers of hemispheric asymmetry in global patterns of ant species richness. Ecology Letters, 12, 324-333 Griffith, D.A. & Peres-Neto, P.R. (2006) Spatial modeling in ecology: the flexibility of eigenfunction spatial analyses Ecology, 87, 2603-2613 Hawkins, B.A., Field, R., Cornell, H.V., Currie, D.J., Guegan, J.-F., Kaufman, D.M., Kerr, J.T., Mittelbach, G.G., Oberdorff, T., O'brien, E.M., Porter, E.E. &

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Turner, J.R.G. (2003) Energy, water, and broad-scale geographic patterns of species richness. Ecology, 84, 3105-3117 Hill, J.G., Summerville, K.S. & Brown, R.L. (2008) Habitat associations of ant species (Hymenoptera: Formicidae) in a heterogeneous Mississippi landscape. Environmental Entomology, 37, 453-463 Hubbell, S.P. (2001) The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, New Jersey, USA. Johnson, R.A. (2001) Biogeography and community structure of North American seed-harvester ants. Annual Review of Entomology, 46, 1-29 Jost, L., Devries, P., Walla, T., Greeney, H., Chao, A. & Ricotta, C. (2010) Partitioning diversity for conservation analyses. Diversity and Distributions, 16, 65-76 Kaspari, M. & Valone, T.J. (2002) On ectotherm abundance in a seasonal environment-studies of desert ant assemblage. Ecology, 83, 2991-2996 Kaspari, M., Yanoviak, S.P. & Dudley, R. (2008) On the biogeography of salt limitation: A study of ant communities. Proceedings of the National Academy of Sciences of the United States of America, 105, 17848-17851 Kaspari, M., Yuan, M. & Alonso, L. (2003) Spatial grain and the causes of regional diversity gradients in ants. American Naturalist, 161, 459-477 Laliberté, E. (2008) Analyzing or explaining beta diversity? Comment. Ecology, 89, 3232-3237 Legendre, P. (2008) Studying beta diversity: ecological variation partitioning by multiple regression and canonical analysis. Journal of Plant Ecology, 1, 3-8 Legendre, P., Borcard, D., Blanchet, F.G. & Dray, S. (2009) PCNM: PCNM spatial eigenfunction and principal coordinate analyses, version 1.9. Available at: http://cran.r-project.org/. Legendre, P. & Gallagher, E. (2001) Ecologically meaningful transformations for ordination of species data. Oecologia, 129, 271-280 Legendre, P. & Legendre, L. (1998) Numerical Ecology. Elsevier, Amsterdam. Longino, J.T., Coddington, J. & Colwell, R.K. (2002) The ant fauna of a tropical rain forest: estimating species richness three different ways. Ecology, 83, 689-702 Mccune, B. & Mefford, M.J. (2006) PC-ORD. Multivariate Analysis of Ecological Data. Version 5.17. In. MjM Software, Gleneden Beach, Oregan, U.S.A Mezger, D. & Pfeiffer, M. (2010) Partitioning the impact of abiotic factors and spatial patterns on species richness and community structure of ground ant assemblages in four Bornean rainforests. Ecography, doi: 10.1111/j.1600- 0587.2010.06518.x Noy-Meir, I. (1973) Desert ecosystems: Environment and producers. Annual Review of Ecology and Systematics, 4, 25-51 Oksanen, J., Kindt, R., Legendre, P., O'hara, B., Simpson, G.L., Peter, S., Stevens, M.H., H. & Wagner, H. (2009) vegan: community ecology package. R package version 1.15-4. (http://cran.r-project.org/). Parr, C.L. (2008) Dominant ants can control assemblage species richness in a South African savanna. Journal of Animal Ecology, 77, 1191-1198 Peres-Neto, P.R. & Legendre, P. (2010) Estimating and controlling for spatial structure in the study of ecological communities. Global Ecology and Biogeography, 19, 174-184 Peres-Neto, P.R., Legendre, P., Dray, S. & Borcard, D. (2006) Variation partitioning of species data matrices: Estimation and comparison of fractions. Ecology, 87, 2614-2625

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Pfeiffer, M., Chimedregzen, L. & Ulykpan, K. (2003) Community organization and species richness of ants (Hymenoptera: Formicidae) in Mongolia along an ecological gradient from steppe to Gobi desert. Journal of Biogeography, 30, 1921-1935 Pfeiffer, M., Tuck, H.C. & Lay, T.C. (2008) Exploring arboreal ant community composition and co-occurrence patterns in plantations of oil palm Elaeis guineensis in Borneo and Peninsular Malaysia. Ecography, 31, 21-32 R Development Core Team (2007) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. (http://www.R-project.org). Rangel, T.F.L.V.B., Diniz-Filho, J.A.F. & Bini, L.M. (2006) Towards an integrated computational tool for spatial analysis in macroecology and biogeography. Global Ecology and Biogeography, 15, 321-327 Ricklefs, R.E. (1987) Community diversity: Relative roles of local and regional processes. Science, 235, 167-171 Rojas, P. & Fragoso, C. (2000) Composition, diversity, and distribution of a Chihuahuan Desert ant community (Mapimi, Me xico). Journal of Arid Environments, 44, 213-227 Sanders, N.J., Lessard, J.P., Fitzpatrick, M.C. & Dunn, R.R. (2007) Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. Global Ecology and Biogeography, 16, 640-649 Smith, T.W. & Lundholm, J.T. (2010) Variation partitioning as a tool to distinguish between niche and neutral processes. Ecography, 9999 Statsoft, I. (2007) STATISTICA (data analysis software system), version 8.0 www.statsoft.com. Vasconcelos, H.L., Vilhena, J.M.S., Facure, K.G. & Albernaz, A.L.K.M. (2010) Patterns of ant species diversity and turnover across 2000 km of Amazonian floodplain forest. Journal of Biogeography, 37, 432-440 Zohary, M. (1973) Geobotanical foundations of the Middle East. Gustav Fischer Verlag, Stuttgart.

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Appendix S1 Measuring of environmental variables Habitat heterogeneity: We recorded habitat horizontal structure at each sample site during May-July 2007 to settle the habitat heterogeneity. We determined the horizontal structure of the habitat by a modified method described by Botes and others (2006). A 1 m2 plastic grid was placed over each pitfall trap, in total 30 grids for each pitfall transect. For measuring the habitat heterogeneity factors we took one photo from top of each grid. We positioned our camera (Canon Powershot IS2) at a 150 cm distance top of the center of each grid, in a vertical angel and shot. With this method we captured each grid and ground covered within it in a one shot. Later we measured three factors: vegetation cover, bare soil, and exposed rock within each grid ® ® from the photos by ADOBE PHOTOSHOP CS3 Extended software. Mean ground vegetation cover, bare soil and exposed rock were calculated for each sampling transect.

Climate variables: We collected climate data from the nearest local weather stations to our study sites that were situated in mean 53 km (SD = 32.1 km, n = 8) from the corresponding study sites. We obtained a group of environmental parameters from these local weather stations to compare climatic variation of the study sites. We used mean daily air temperature, annual temperature range, annual rainfall, summer rainfall and relative humidity. We normalized these variables by log-transformation.

Productivity: We used the normalized difference vegetation index (NDVI) as the surrogate of productivity. Normalized difference vegetation index, based on a ratio of red and near-infrared reflectance, responds to the absorption of red light by chlorophyll and is a measure of greenness. It has been shown that NDVI has strong relationship with net primary productivity (NPP) (Box and others 1989) and this relationship is relatively consistent over different ecosystems (Fairbanks and McGwire, 2004). The NDVI data were collected from the moderate Resolution Imagining Spectroradiometer (MODIS) an instrument aboard NASA’s Terra and Aqua satellites. The NDVI data used in this study were obtained from the Land Processes Distributed Active Archive Center (LP DAAC) (https://lpdaac.usgs.gov/lpdaac/products/modis_products_table) which is a component of NASAs Earth Observing System (EOS). We generated NDVI values at a 500 - m spatial resolution and temporal resolution of 16 days for each site by overlaying site localities on NDVI data layers by ESRI ARCGIS software. There are 23 such layers for

82 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages a whole year (May/July 2006 – May/July 2007). We calculated the yearly mean values of 23 NDVI images and applied them as explanatory variable for each site.

Elevation: Elevation variable was generated by overlying site localities on elevation data layer which was obtained from the Shuttle Radar Topography Mission (SRTM) (http://dds.cr.usgs.gov/srtm/).

References:

Botes, A., McGeoch, M.A., Robertson, H.G., Van Niekerk, A., Davids, H.P. & Chown, S.L. (2006) Ants, altitude and change in the northern Cape Floristic Region. Journal of Biogeography, 33, 71-90. Box, E.O., Holben, B.N. & Kalb, V. (1989) Accuracy of the AVHRR vegetation index as a predictor of biomass, primary productivity and net CO2 flux. Plant Ecology, 80, 71-89. Fairbanks, D.H.K. & McGwire, K.C. (2004) Patterns of floristic richness in vegetation communities of California: regional scale analysis with multi- temporal NDVI. Global Ecology and Biogeography, 13, 221-235

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84

5Productivity, solely, does not explain species richness of ants in a dryland habitat

Omid Paknia & Martin Pfeiffer In preparation for publication

ABSTRACT

A large body of literature proposes that productivity is responsible for the biodiversity gradients in terrestrial habitats at large extents /grains. But at smaller extents/grains other competitive explanatory variables or models diminish or weaken the effect of productivity predictor. Here we present a short extent of geographical gradient of ant species richness in the Central Persian desert ecoregion. Applying information theoretical model selection procedures, we evaluated the support for the productivity and a series of alternative models, to reveal the impact of key variables and validity of models on ant species richness. In total, the central desert yielded 46 species. Results supported nine models as a confidence set of the best models. The univariate productivity model and multivariate tolerance - productivity model, including temperature range and productivity variables, received the highest support. A curvilinear relationship was evident between ant species richness and productivity. In contrast to earlier works, we found a reverse pattern of latitudinal gradient in ant species richness, increasing the species number along latitude. Our results suggest that in our dryland ecosystem, the plant productivity is the most important variable for Chapter 5 - Ant Species Richness in a Dryland Habitat controlling the ant species richness. But this variable, solely, could not explain the spatial gradient in ant species richness. A valid inference should take especially the tolerance-productivity model into account. Our findings also indicate at the small extents competitive variables which co-vary with productivity have a significant role in the determination of species number.

Keywords: ants, desert, ecoregion, Iran, productivity, short extent, small grain, species richness, tolerance.

INTRODUCTION

Spatial gradients in the number of local coexisting species have long attracted community ecologists. At the large extents and grains, such as global or continental extent, an extensive literature suggests that a water –energy combination, indirectly as plant productivity, drive the biodiversity gradients in terrestrial habitats (Currie, 1991; Hawkins et al., 2003), Although it is suggested that biological relatively to water – energy drivers should be applicable everywhere and always (O'brien, 2006), at the smaller grains and extents the observed patterns are diverse and far from a generalization (Field et al., 2009). This extent/grain-dependence results from the relatively short water-energy range that is sampled by studies with small extents (Field et al., 2009), as water – energy has superiority mainly at large extents/grains. Therefore, at the small extent/grains the effect of the water-energy combination on species richness can be weakened or blurred by the effect of a wide range of competitive explanatory models, such as habitat complexity (Jimenez-Valverde & Lobo, 2007), dispersal (Shurin & Allen, 2001) or stress tolerance (Gough et al., 1994).

These scale-dependence differences have been also observed in ants. Ants have been often chosen as a model taxon for investigating patterns of spatial gradients, because they are important components of the most terrestrial ecosystems as herbivores, omnivores or predators (Hölldobler & Wilson, 1990), and their richness varies widely among habitats (Gotelli & Ellison, 2002). At large extent, global or continental, it has been demonstrated that ant species richness is positively associated with productivity

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(Kaspari et al., 2000), but negatively associated with temperature range (Dunn et al., 2009). At the smaller scales, however, there is no such generalization. For example, studies have been conducted in arid or semi-arid lands and linking ant species richness or diversity to productivity or its surrogates (e.g. rainfall) mostly have not shown a positive correlation. Seed harvesting ant diversity was not positively correlated to rainfall in Australia (Morton & Davidson, 1988). Similarly, there was no positive correlation among ants species richness and rainfall in Mongolia (Pfeiffer et al., 2003), and Paraguayan dry Chaco (Delsinne et al., 2010). For animals, deserts and semi-deserts are productivity-controlled ecosystems. In these ecosystems the low precipitation level, high evapotransporation ratios and high temperature limit the primary productivity to very low levels (Noy-Meir, 1973). As a result, productivity which is closely dependent to water is suggested as the main limiting and forcing factor for species richness of animals in warm and dry ecosystems (Hawkins et al., 2003). Thus, the intuitive question will be why the species richness of ants does not follow the expected pattern. The absence or negative relationships of productivity and ant species richness have been linked to other important explanatory variables such as edaphic variability (Morton & Davidson, 1988), co-varying of productivity with climate factors such as temperature (Pfeiffer et al., 2003), and sociality behavior of ant taxa (Delsinne et al., 2010), suggesting that there are possibly alternative variables or models for explaining the geographical variation in ant diversity.

To answer the above question, in the present study, we examined the geographical gradient in species richness of ants in Central Persian desert basins of Iran, across a small latitudinal gradient. An insight into small extents geographical patterns may be gained, by evaluating the support of several competing hypotheses (models) which are thought to be responsible for the geographical variation of species richness. The multi-model inference is a promising approach in this case, as it is rooted in the view that understanding of ecological processes can best be achieved by simultaneously weighing evidence for multiple competitive hypotheses, formulated as appropriate models (Hobbs & Hilborn, 2006). Among the many hypotheses that have been suggested to explain the species richness and diversity gradient, five can be considered as the main competitive models for explaining the ant diversity gradient in semi-cold drylands such as our under study system: These are: The productivity - species richness model that suggests that geographical variation in species richness

87 Chapter 5 - Ant Species Richness in a Dryland Habitat correlates positively with productivity (Currie, 1991). The habitat heterogeneity model, proposing that higher diversity occurs in an area with greater diversity of habitats, because animal species have habitat preferences and structurally complex habitats may provide more niches and diverse ways of exploiting the environmental resources (Tews et al., 2004). The tolerance model, assuming that species richness in a particular area is limited by the number of species that can tolerate the local conditions (Currie et al., 2004). The evolutionary diversification model that suggests evolutionary rates are faster at higher ambient temperatures due to shorter generation times, higher mutation rates, and faster physiological processes (Currie et al., 2004). Taking the geographical context of the study (dryland) and the specific life history of ants into consideration, the plant-ant interaction model is another alternative model for explaining species richness of ants. This model suggests that plant species richness in the arid environment, directly affects ant species richness. In arid lands, plant - ant interaction can be mainly viewed either as an antagonistic interaction in which plants are food sources of ants, e.g. harvester ants, or as a mutualistic interaction (Gómez et al., 1996; Rico-Gray & Oliveira, 2007). In this study we examine the strength of support for these five univariate models (Table 1), and all possible combinations of these five models (totally 32 models) to evaluate and choose the best model or set of models and to investigate how well this/these model(s) and explanatory variables explain ant species richness and diversity in a vast dryland ecosystem.

METHODS

Study area

We conducted the study in the Central Persian Desert Basins ecoregion, in Iran. This ecoregion covers approximately 600,000 square kilometers of arid steppe and desert regions of central Iran and a small part of Afghanistan. Dominated by large areas of salt deserts, sabkhas and salt marshes, the region shows a great variety of climate conditions (e.g. precipitation and temperature), and topography (Breckle, 2002). The climate of the area is continental with very high annual and daily fluctuations of temperature. The region can be regarded as a transitional zone between the hot deserts of Arabian Peninsula and the cold deserts of Central Asia (Breckle, 2002).

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Figure 1 Location of eight protected areas in Central Persian desert basins ecoregion (shown with grey color). Four sampling sites were chosen within each protected areas.

Data collection

We sampled ant assemblages at 32 sites within eight protected areas from 28 April to 30 May 2008, during the yearly periods of peak ant activity. Within each protected area, we chose four sampling sites which were separated on average by 1108 m (± 696 m SD). At each site, a group of 20 pitfall traps were laid out in a 5 × 4 grid with traps spaced at 10-m intervals. All pitfall traps were left open for a period of four days. We sorted ants from the pitfall collection and identified them to species where possible or assigned to morphospecies. Voucher specimens of each species collected are held in the AntBase.Net Collection (ABNC) of the Institute of Experimental Ecology of the University of Ulm (http://www.antbase.net) and in the private ant collection of the corresponding author.

We measured or extracted five variables which were related to the tested models (Table 1). We extracted climate variables (minimum temperature and annual mean

89 Chapter 5 - Ant Species Richness in a Dryland Habitat temperature) from the Worldclim database at ~ 1 km spatial resolution data layer (Hijmans et al., 2005). We used the Normalized Difference Vegetation (NDVI) data layer obtained from the NASA MODIS algorithm (MOD13Q1) at 250 m spatial resolution and 16 days temporal resolution for a period of May 2007-May 2008 as surrogate of Net Primary Productivity (NPP). The number of plant species was counted along two 40 m transects within each pitfall sampling grid. Horizontal and vertical structure of the habitat were determined by a modified method described by Botes et al. (2006). A 1 m^2 plastic grid was placed over each pitfall trap, in total 20 grids for each pitfall transect. For measuring the habitat heterogeneity factors (horizontal structure), we took one photo from the top of each grid. We positioned our camera (Canon Powershot IS2) in a vertical angel at a 150 cm distance to the top of the center of each grid and shot. With this method we captured the ground cover of each grid in a single shot. Later we measured three factors from the photos by ® ® ADOBE PHOTOSHOP CS3 Extended software: vegetation cover and the partition of bare soil, and exposed rock within each grid and calculated the means for each transect. Habitat complexity (vertical structure) was measured by determining foliage height. Vegetation height was measured at the four corners of a 1 m^2 plastic grid, by placing a 160 cm rod vertically through the vegetation and recording the number of contacts with the foliage at 10 cm height increments (0-10, 11-20,21-30,…,151-160). The total number of contacts within each sampling plot was calculated as a measure of habitat complexity.

Table 1 The five candidate models used in analysis of species richness of ants. See text for details of explanatory variables and models’ explanations.

Model Explanatory variable Explanation Available energy in the environment directly affects ant Productivity NDVI species richness Heterogeneity of habitat facilitates the coexistence of a Heterogeneity Habitat Heterogeneity greater number of species Ant species richness is limited to those species that can Tolerance Minimum temperature tolerate the local condition Plant species richness in the environment directly affects Biotic interaction Plant species richness ant species richness Ambient energy Mean annual temperature Evolutionary rates are faster at higher ambient temperatures

Data analysis

As “observed species richness” is sensitive to the number of individuals collected in samples (Gotelli & Colwell, 2001), and the pitfall trapping method is sensitive to the

90 Chapter 5 - Ant Species Richness in a Dryland Habitat movement heterogeneity among ant species (Melbourne, 1999), we applied non- parametric species richness estimators (Incidence-based Coverage Estimator (ICE), Chao 2, Jackknife 1, Jackknife 2, and Michaelis-Menten) based on Brose and Martinez (2004)’s movement heterogeneity framework for estimating species richness of ants in each site. We used ESTIMATES 7.50 (Colwell, 2005) to calculate estimated species richness (Set).

Based on our five explanatory variables (Table 1), we constructed 32 generalized linear models (GLMs) for ant species richness, with Poisson distribution and logarithmic link function. Prior to applying multimodel inference approach, we inspected overdispersion of models by estimating the variance inflation factor (ĉ) from a global model (Burnham & Anderson, 2002). A value exceeding 1 indicates the presence of overdispersion in models. To identify the best approximate model for the prediction of species richness and diversity of ants, we evaluated the strength of support for alternative models rather than the significance of individual parameters. Following an information theoretic model selection approach suggested by Burnham & Anderson (2002), we employed the Akaike Information Criterion corrected for small sample size (AICc,), to rank models according to their strength of support from the data, with the best model having the lowest AICc value. We used ∆ AICc, the difference between the lowest AICc value, and the AICc value of the other models to select the best sets of models. Models with a ∆ AICc value smaller than 2 are all likely to be a good model and hence they should all be used when making further inference about a system (Richards, 2005). We used also Akaike weights (AICc w) to evaluate the uncertainty of model selection. A model’s Akaike weight is considered as the weight of evidence that the model is the best of those in the model set. In addition, the Akaike weight can be interpreted as an estimate of the chance that the model will be chosen as the best model if the study would be repeated. These weights sum to 1 across all models and a single model with higher than 0.9 Akaike weights, can be regarded as the best model (Burnham & Anderson, 2002). If the Akaike weight of a single model is < 0.9 and competing models give alternative inference, then a 90% confidence set of best models can be selected, by summing the Akaike weights from largest to smallest until the sum of Akaike weight is just ≥ 0.90. The AIC differences (∆) and Akaike weights (w) allow scientific hypotheses, represented by models, to be ranked appropriately. Evidence for the importance of each explanatory variable was

91 Chapter 5 - Ant Species Richness in a Dryland Habitat obtained by summing Akaike weight over those models with a particular variable present (Burnham & Anderson, 2002). We measured goodness of fit of the best model(s) by the likelihood ratio chi-squared statistic, based on comparisons between the maximum values of the log-likelihood function for a minimal model with no covariate terms and the other models (Dobson, 2002).

Because the presence of spatial autocorrelation in the data of an ecological study can affect the AIC-based model selection (Diniz-Filho et al., 2008), we used principal coordinates of neighbour matrices (PCNM) analysis to generate spatial predictors that could be directly integrated into our regression models for taking the effect of space into account (Dray et al., 2006; Diniz-Filho et al., 2008). The generated spatial predictors were used as spatial filters for each model. Environment variables were considered as “floating” explanatory variables, while spatial variables (PCNM) were “fixed” explanatory variables in the model selection process. This means that spatial variables were present in all models for taking the present spatial structure into account, but habitat variables were included into the model depending on the explained hypothesis. The presence of significant spatial autocorrelation in the residuals of GLM models was inspected by randomization tests for Moran’s I. We employed a modified t-test developed by Dutilleul (1993) for partialling out the effect of spatial autocorrelation and testing the significant correlations of the response variable and environmental variables, using SAM software (Rangel et al., 2006). In the results “corrected P” indicates the correction of the P value after factoring out the effect of spatial autocorrelation. Principal coordinates of neighbour matrices were generated by the “PCNM” library (Legendre et al., 2009) of the “R” package (R Development Core Team, 2007). Model construction, model selection and model averaging were computed using the “AICcmodavg” library (Mazerolle, 2010). Moran’s I of residuals of models were calculated by the gearymoran function of the “ade4” library (Chessel et al., 2009). Likelihood ratio tests were carried out by lrtest function of the “lmtest” library (Hothorn et al., 2009). Graphs were depicted by “ggplot2” library (Wickham, 2010).

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RESULTS

In total, we collected 20622 individual ants from pitfall traps at 32 sites. We detected 46 ant species/morphospecies in eleven genera corresponding to 2805 species occurrences. The number of species per site ranged from 0-16. The estimated number of species at the sites ranged from 0-18. Species richness was independent of elevation (rs = 0.1781, corrected P = 0.27), but interestingly, species richness of ants increased roughly significantly with latitude (S: rs= 0.6249, n = 32, corrected P = 0.06; Figure 2).

36 r = 0.6249, n = 32, Corrected P = 0.06

35

34 Latitude

33

32

0246810121416 Species richness

Figure 2 The relationship between ant species richness and latitude in Central Persian desert basin.

Productivity increased with elevation (rs = 0.3816, n = 32, corrected P = 0.066) and latitude (rs = 0.7737, n = 32, corrected P = 0.073), both marginally significant. Species richness of arboreal ants (Crematogaster and Camponotus) correlated significantly with habitat complexity (rs = 0.53, n = 32, corrected P = 0.007). However, species richness of ground living ants was not associated with habitat complexity (rs = 0.49, n = 32, corrected P = 0.113).

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A curvilinear relationship was evident between ant species richness and productivity (Figure 3). Comparing linear model vs. quadratic polynomial model with analysis of deviance, the latter model showed a significant (P = 0.008) decrease of deviance (linear model: D = 39.09, df = 30 vs. quadratic model: D = 32.16, df = 29). As a result, we applied a nonlinear (quadratic) form of productivity variable in our model selection.

Figure 3 Productivity – ant species richness relationship in Central Persian desert basin. Fitted line represents the GLM quadratic polynomial model.

PCNM generation over 32 sample sites produced five PCNM eigenvalues. Forward selection procedure retained two significant PCNM out of five positive PCNM vectors. These two PCNM eigenvalues were used as spatial filters and fixed explanatory variables in all 32 GLM models.

The productivity model was indicated as the best model by both Akaike difference and Akaike weights approaches (Table 2). However, a substantial model selection uncertainty was evident as this model had an Akaike weigh of only 0.29. As result, a

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90% confidence set of models was selected which contained nine models (Table 2). The second best model was tolerance - productivity, included the variables productivity and temperature range. This model had a small ∆ AICc (0.47), indicating a strong support for a multimodel inference. Other seven last models showed approximately small and similar evidences of support. In the third best model annual temperature range was substituted with temperature range. The ambient energy - productivity, however, received small support with a relatively large ∆ AICc (2.73), and small Akaike weigh of only 0.07. In the fourth best model productivity, temperature range and annual temperature range appeared in a model together. The predictor habitat complexity and biotic interactions appeared for the first time in the fifth and sixth models, respectively (Table 2). Among five univariate models, only the productivity model came to this confidence set and was ranked as the first best model. Both Akaike difference and Akaike weigh approach cast substantial doubt concerning the usefulness of the other four univariate models, as all had ∆ AICc > 9.63 and AICc w = 0. Variance inflation factor (ĉ) of global model was 0.89, indicating that our multimodel inference was not affected by overdispersion of the models. All nine best models were highly significant (Table 3).

Averaging the parameter estimates across 32 GLM models, productivity was the most important variable for explaining ant species richness (Table 4). This variable occurred in all nine confidence set models. Temperature range was the second important variables and appeared in four out of nine models. These two predictors showed also high coefficients of regression (Table 3). Moran’s I for residuals of all GLM models were small in numbers (Moran’s I < 1.85) and insignificant, indicating that by using PCNM’s filters, spatial autocorrelation in the residuals of GLM models was completely removed.

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Table 2 Thirty two GLM selected models, sorted by their Akaike weigh. Model name indicates the explanatory variable(s) that has/have been used in each model. K is the number of estimable parameters. All models include two similar PCNM spatial filters. AICc = biased-corrected Akaike Information Criterion (see text for details). ∆ AICc = difference between AIC of the given model and the minimum AIC found for the set of models compared. AICc wt = Akaike Information Criterion weights. Cum wt = cumulative Akaike weights. Moran’s I residual autocorrelation and its P value for each model have been given in last two columns.

Model rank Model K AICc ∆ AICc AICc wt Cum wt Moran’s I P value 1 Productivity 5 167.24 0.00 0.29 0.29 0.032 0.33 2 Tolerance - Productivity 6 167.71 0.47 0.23 0.52 - 0.002 0.40 3 Ambient energy - Productivity 7 169.97 2.73 0.07 0.59 - 0.07 0.55 4 Ambient energy - Tolerance - Productivity 6 170.33 3.09 0.06 0.65 0.031 0.31 5 Complexity - Productivity 6 170.34 3.11 0.06 0.71 0.046 0.30 6 Tolerance - Biotic interaction - Productivity 7 170.36 3.12 0.06 0.77 - 0.02 0.47 7 Tolerance - Complexity - Productivity 7 170.41 3.17 0.06 0.83 0.007 0.42 8 Biotic interaction - Productivity 6 170.58 3.34 0.05 0.88 0.034 0.34 9 Ambient energy - Complexity - Productivity 7 173.09 5.85 0.02 0.90 0.055 0.29

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Table 3 The change in the likelihood from a model of ant species richness with no terms to that nine best GLM models containing different environmental variables. The statistical significance of the change in likelihood was determined by a Chi-squared test.

Model Log-likelihood Df Chi-squared P value No terms added -93.177 Productivity -77.701 4 30.952 <0.001 Tolerance - Productivity -75.938 5 34.477 <0.001 Ambient energy - Productivity -77.305 5 31.745 <0.001 Ambient energy - Tolerance - Productivity -75.831 6 34.693 <0.001 Complexity - Productivity -77.492 5 31.37 <0.001 Tolerance - Biotic interaction - Productivity -75.847 6 34.659 <0.001 Tolerance - Complexity - Productivity -75.871 6 34.611 <0.001 Biotic interaction - Productivity -77.610 5 31.134 <0.001 Ambient energy - Complexity - Productivity -77.211 6 31.932 <0.001

Table 4 Parameter estimates averaged across 32 GLM models, using biased-corrected Akaike Weights (AICcw). Two PCNM spatial filters have not been shown here as they have been considered as fixed explanatory variables in all 32 models. Importance = Relative importance of variables based on sum of Akaike weights of the models that include the variable. Coefficient = the model-averaged coefficient based on all models including the parameter of interest. Unconditional SE = the unconditional standard error for the model-averaged coefficient and its 95% intervals.

Variable Importance Coefficient Unconditional SE 95% Lower 95% Upper NDVI 2.75 0.57 0.63 3.85 0.98 NDVI2 - 0.83 0.35 - 1.53 - 0.13 Temperature range 0.44 - 0.17 0.03 - 0.31 - 0.09 Mean annual temperature 0.39 0.1 0.06 0.01 0.25 Plant species richness 0.27 0.02 0.03 - 0.04 0.09 Complexity 0.16 0.01 0.01 - 0.02 0.01

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DISCUSSION

Productivity has been widely identified as principal driver of animal diversity at large extents/grains (Currie, 1991). In accord with these findings, we found that productivity has a decisive role in the geographical variation of species richness of ants in Central desert (Table 2), although, our samples were taken at small spatial grains and a short extent. Evaluating the support for 32 models, the key finding of this study is that the productivity model is more supported than the other evaluated hypotheses by our data on ant species richness in Central Persian deserts. Although, this model, solely, could not explain the geographical gradient in ant species richness, and there are other competitive models which are likely to become the best model, if the study would be replicated. We found especially strong supportive evidence for the tolerance-productivity model, suggesting that a valid inference on geographical pattern of ant species richness in our productivity-controlled ecosystem is possible by taking this model into account. This outcome, however, is not surprising, as ants are well known thermophiles and many species cannot survive or establish constant large populations in cold instable temperatures (Hölldobler & Wilson, 1990). Many ant species either do not possess any adaptation to cold temperature, therefore they will be extinct very soon after arriving from warmer regions, or they can establish their colonies, but their access to energy is limited by temperature fluctuations for these species (Dunn et al., 2010). Foraging for food in a preferable temperature is a possible solution, however, adds to competing exclusion pressure which is well known in ants (Andersen & Patel, 1994; Andersen, 1995; but see Cerdá et al., 1998). As a result, these species go to local extinction because of the direct effect of temperature stress on their colonization or their energy-harvesting ability or they are possibly excluded from the region by dominant ants in a competitive exclusion process. In an optimistic assumption, these species survive in rarity with small abundances. Our further analysis confirms the last conclusion (data not shown), as we found also a strong effect of temperature range on ant diversity (Shannon diversity), indicating that tolerance is not only responsible for the presence or absence of species, but also for their abundances. This result also shed some light on the question why ant species richness is not fully and lonely predicted by productivity gradient at small extents/grains, showing that other powerful models enforce diversity of ants along with the productivity model.

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The fact that other possible co-explanatory factors, besides productivity influence species richness patterns, has been pointed out by several authors (Morton & Davidson, 1988; Pfeiffer et al., 2003; Delsinne et al., 2010), but has not been tested and demonstrated explicitly. It is also noteworthy that most studies that failed to demonstrate an association among productivity and ant species richness have employed rainfall as a surrogate of productivity (e.g. Pfeiffer et al., 2003; Delsinne et al., 2010). Although in arid lands productivity is tightly associated with rainfall, rainfall should be viewed only as a potential surrogate of plant productivity, because, plant productivity in drylands can also be limited by nutrient. Nitrogen is key limiting nutrient in most deserts (Jones & Shachak, 1990). As result, instead of rainfall, NPP and its proxies such as actual evapotranspiration (AET) and NDVI may be more suitable productivity metrics for the study of species-energy relationships (Evans et al., 2005; Clarke & Gaston, 2006). The positive effect of ambient energy on ant species richness at small grains has been reported widely across altitudinal and latitudinal gradients (Kaspari et al., 2003; Sanders et al., 2007; Kumschick et al., 2009), connecting it to higher evolutionary and metabolic rates or thermophilic behaviour of ants. As we sampled ants in a semi-cold desert region, mean annual temperature showed relatively high support, suggesting that warmer sites support more ant species. Both the Biotic interaction hypothesis and the Heterogeneity hypothesis showed similarly low support. In our study both, plant species richness and heterogeneity, are correlated with productivity (heterogeneity rs = 0.71, n = 32, corrected P = 0.051; plant species richness rs = 0.65, n = 32, corrected P = 0.04). A correlation of these three predictors has been observed also at the global scale (Kreft & Jetz, 2007). Despite of their collinearity, it is suggested that these three variables affect species richness of animal taxa in different ways (Clarke & Gaston, 2006). The classic and well known proposed mechanism for the interaction of productivity and species richness, suggests that productivity increases the number of species through increasing the number of individuals (Currie, 1991). The test of this mechanism in different taxa and different habitats has shown mix results (Mcglynn et al., 2010 and references therein). It has been suggested that the greater productivity also can be interpreted as enhanced diversity of food plants (plant species richness), which allows for the evolution of a wider range of specialist herbivores, and in turns increases the number of species in other trophic levels, e.g. omnivores or scavengers. Haddad et al.

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(2009) demonstrated that arthropod species richness decreases greatly due to the long term loss of plant species richness. Productivity can also be translated into habitat complexity which may allow a greater number of species utilizing these habitats (Evans et al., 2005; Clarke & Gaston, 2006). Our multi-model inference is consistent with this modified mechanism, suggesting that biotic interaction and habitat complexity also play a, however small, role in species richness of ants. Further studies are needed to disentangle the direct and indirect effects of these three variables on ant species richness. Interestingly, habitat complexity showed a positive correlation with the species richness of arboreal ants, indicating that habitat complexity prepares more niches for co-occurring of ant species. Contrary to broadly observed and accepted pattern of latitudinal gradient - species richness, ant species richness increased across our short latitudinal survey (Figure 2). This pattern mirrored the steep productivity gradient that exists across the latitudinal gradient. Studying pattern of ant species richness in a temperate forest and bogs ecosystem, Gotelli and Ellison (2002) found that the usual pattern of decreasing of ant species richness across latitude occurs along their short transect, concluding that this general pattern does not occur only at large global and continental extents and because of differences between tropical and temperature communities, but also in short extents. Our result, however, corroborates with Hillebrand’s (2004) argument, suggesting that at small latitudinal extents there is no systematic changes in latitudinal gradients of species richness. Absence of systematic changes in short extents of latitudinal gradient should also be linked to the not superiority of equator-to-pole gradient of water – energy combination at smaller extents, where other factors act strongly beside it. In conclusion, our findings are consistent with comprehensive studies at global and continental extents, suggesting that productivity is responsible for the spatial gradient in species richness. The work presented here, however, highlights the role of other competitive environmental variables and competitive models in determination of species richness at spatial extents/grains. The competitive variables/models most likely vary among different ecosystems. As a result, more studies conducting in diverse ecosystems are necessary to distinguish them.

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ACKNOWLEDGMENTS The research permits to work in natural reserves were kindly granted by the Department of the Environment of Iran. We thank Alexander Radchenko for extensive help and advice with ant species identification. The support of Elisabeth K.V. Kalko (Head of the Experimental Ecology Institute, Ulm University) made this work possible. O.P. is a fellow of the German Academic Exchange Service (DAAD).

REFERENCES

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Rangel, T.F.L.V.B., Diniz-Filho, J.A.F. & Bini, L.M. (2006) Towards an integrated computational tool for spatial analysis in macroecology and biogeography. Global Ecology & Biogeography, 15, 321-327 Richards, S.A. (2005) Testing ecological theory using the information-theoretic approach: examples and cautionary results. Ecology, 86, 2805-2814 Rico-Gray, V. & Oliveira, P.S. (2007) The ecology and evolution of ant-plant interactions. The University of Chicago Press, Chicago. Sanders, N.J., Lessard, J.P., Fitzpatrick, M.C. & Dunn, R.R. (2007) Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. Global Ecology and Biogeography, 16, 640-649 Shurin, J.B. & Allen, E.G. (2001) Effects of competition, predation, and dispersal on species richness at local and regional scales. The American Naturalist, 158, 624-637 & .Tews, J., Brose, U., Grimm, V., Tielbِrger, K., Wichmann, M.C., Schwager, M Jeltsch, F. (2004) Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures. Journal of Biogeography, 31, 79-92 Wickham, H. (2010) ggplot2: An implementation of the grammar of graphics, version 0.8.8 Available at: http://cran.r-project.org/.

104 Myrmecological News 11 151-159 Vienna, August 2008

A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran

Omid PAKNIA, Alexander RADCHENKO, Helen ALIPANAH & Martin PFEIFFER

Abstract The first checklist of the ants of Iran is presented. The study is based on a comprehensive review of literature and the examination of material from our own collections and from several museums and institutions of different European countries. 110 species belonging to 26 genera of six subfamilies of the Formicidae (Formicinae, Myrmicinae, Ponerinae, Dolichoderinae, Dorylinae and Aenictinae) are recognized from Iran. Most of the reported species were sampled in the north of the country, mostly near human settlements. One subfamily (Dorylinae), two genera (Ponera LATREILLE, 1804 and Dorylus FABRICIUS, 1793), as well as seven species (Aphaenogaster gibbosa (LATREILLE, 1798), A. kurdica RUZSKY, 1905, Crematogaster bogojawlenskii RUZSKY, 1905, Messor minor (ANDRE, 1883), Tapinoma karavaievi EMERY, 1925, Temnothorax parvulus (SCHENCK, 1852) and Tetramorium inerme MAYR, 1877) were recorded for the Iranian fauna for the first time. The most speciose genera were Camponotus MAYR, 1861, Cataglyphis FOERSTER, 1850 and Messor FOREL, 1890 with 19, 14 and 13 species, respectively. Palaearctic zoogeographic elements prevail in Iran, but several Oriental and Afrotropical genera were also found. Many parts of the country are still studied insufficiently or even not studied at all and we suppose that the total species richness in Iran is essentially higher. Key words: Ants, Iran, checklist, new records. Myrmecol. News 11: 151-159 (online 29 June 2008) ISSN 1994-4136 (print), ISSN 1997-3500 (online) Received 29 October 2007; revision received 30 April 2008; accepted 5 May 2008 M.Sc. Omid Paknia (contact author), Dr. Martin Pfeiffer, Institute of Experimental Ecology, University of Ulm, Albert- Einstein Allee 11, D-89069 Ulm, Germany. E-mail: [email protected] Prof. Dr. Alexander Radchenko, Museum and Institute of Zoology, PAS, Wilcza str., 64, 00-679, Warsaw, Poland. M.Sc. Helen Alipanah, Iranian Research Institute of Plant Protection (TRIPP), P.O. Box 19395-1454, Tehran, Iran.

Introduction Much faunistic research on ants has recently been carried and has continued until now. RADCHENKO (1994a, b, 1995, out in several parts of Asia (e.g., East and Southeast Asia: 1996b, 1997a, b), SEIFERT (2003) and TAYLOR (2006) de- KUPYANSKAYA 1990, TERAYAMA 1994, WU & WANG 1995, scribed and recorded several species from Iran. ARDEH CHUNG & MOHAMED 1996, IMAI & al. 2003, OGATA 2005, (1994) recorded 13 ant species from the Karaj Region, Ali- RADCHENKO 2005; India: BHARTI 2002a, b; Central Asia: panah and colleagues studied the ant fauna of Tehran (ALI- TARBINSKY 1976, DLUSSKY & al. 1990, SCHULTZ & al. PANAH & al. 1995, 2000), of the southwest of Iran (Khu- 2006, PFEIFFER & al. 2007); however, in the west and south- zestan province) (ALIPANAH & DEZHAKAM 2000), and of west of Asia only the ant fauna of Saudi Arabia has been several other parts of the country (ALIPANAH 2004). RAD- studied satisfactorily (COLLINGWOOD 1985, COLLINGWOOD CHENKO & ALIPANAH (2004) documented the first Iranian & AGOSTI 1996). The ant fauna of other countries from species of the subfamily Aenictinae. TIRGARI & PAKNIA this region, including Iran, has been investigated only partly. (2004, 2005) and PAKNIA & KAMI (2007) reported 21 spe- Iran situates mainly on the Iranian plateau and covers cies from various areas of the country. an area of 1,623,779 km2. The country is located in the As Iranian myrmecologists have published their reports Palaearctic and naturally divided into six biomes and 14 mostly in local journals or presented them at certain na- ecoregions (OLSON & al 2001). At the same time, its bor- tional scientific congresses, it is difficult for foreign myr- ders are close to the boundaries of the Afrotropical and Ori- mecologists to access this literature. Other problems arise ental regions in the south and south-east of the country. from old records that need revision. For all of these reasons Such position, as well as high environmental diversity, pro- a preliminary checklist of the Iranian ants is a helpful ref- motes the diversity of the Iranian flora and fauna. erence for myrmecologists that are interested in Asian ants. The history of the myrmecological investigations in Iran can be divided into two main periods: the first third of the Material and methods XX century includes works of FOREL (1904a, b), EMERY This study is based on a comprehensive review of litera- (1906), CRAWLEY (1920a, b, 1922) and MENOZZI (1927). ture and the examination of material both collected by two After that, until the 1990ies new data on the Iranian myr- of the authors of this paper (OP & HA), by Hamidreza Haji- mecofauna were very scarce (e.g., ARNOLDI 1977). The qanbar and by many other entomologists as well as on in- second period of ant research in Iran started in the 1990ies vestigated material preserved in the following museums

105 Chapter 6 - Ant Species List of Iran and institutions: Hay Mayans Insect Museum in the Iran- Dorylus sp.: Khuzestan province, Ahvaz, 4.X.1954, leg. ian Research Institute of Plant Protection, Tehran, Iran Sal, 1 worker (HMIM), det. H. Alipanah. (HMIM), Zoological Museum of Gorgan University, Gor- Messor minor (ANDRE, 1883): Hamedan province, Hame- gan, Iran (ZMGU), Institute of Zoology of Ukrainian Na- dan, 25.V.1997, leg. P. Moghadasi, 5 workers (HMIM), tional Academy of Sciences, Kiev, Ukraine (IZK), Zool- det. A. Radchenko. ogical Museum of Moscow State University, Moscow, Rus- Ponera sp.: Golestan province, 20 km East Gorgan, Tus- sia (ZMMU), Zoological Institute of Russian Academy of kestan forest, 28.V.2005, leg. Omid Paknia, 9 workers Sciences, St. Petersburg, Russia (ZISP), Museum and In- (ZMGU), det. A. Radchenko. stitute of Zoology of Polish Academy of Sciences, War- Tapinoma karavaievi EMERY, 1925: Tehran province, Teh- saw, Poland (MIZ), The Natural History Museum, London, ran: 19.X.1992, leg. Shemiranat; Lavasanat, 1600 m, 12. UK (BMNH), Museo Civico di Storia Naturale "Giacomo VII.1994, leg. H. Alipanah, 2 workers (HMIM); Khuze- Doria", Genoa, Italy (MSNG), Istituto di Entomologia, Uni- stan province, Ahwaz, 23.XI.1997, leg. M. Dezhakam, versity di Bologna, Bologna, Italy (IEUB), and Naturhis- 3 workers (HMIM); Khorasan Razavi province, Mash- torisches Museum Basel, Basel, Switzerland (NHMB). had, 23.I.1999, leg. M. Ghasemi, 5 workers (HMIM); Altogether, we examined about 3350 specimens. We Mazandaran, Miankaleh, 14.IV.2005, leg. O. Paknia, 6 verified existence of the voucher specimens for all species workers (ZMGU), det. A. Radchenko. listed in Table 1, except for several species recorded by Temnothorax parvulus (SCHENCK, 1852): Golestan pro- CRAWLEY (1920a, b), MENOZZI (1927) and ANARAKI (1981) vince, Gorgan, Tuskestan forest, 28.V.2005, leg. S. Shal- as we could not locate their vouchers, and the Cardiocon- chian, (ZMGU), det. A. Radchenko. dyla species recently recorded for Iran by SEIFERT (2003). Tetramorium inerme MAYR, 1877: Khorasan Razawi pro- For all species taxonomically described from Iran we as- vince, Sarakhs, 27.IV.2007, leg. H. Hajiqanbar, (IZK), sumed that there are vouchers for them. We avoided to list det. A. Radchenko. unavailable names. While there are no keys specifically for the identifica- Reidentified species tion of the ants of Iran, we used existing keys for adja- Based on the re-examination of voucher specimens (ZMGU, cent regions and taxonomic revisions of different ant gen- HMIM) by A. Radchenko, several previously recorded era (ARNOLDI 1976, 1977, COLLINWOOD 1985, DLUSSKY species are excluded from the Iranian fauna: Camponotus & al 1990, ARAKELYAN 1994, DLUSSKY & RADCHENKO cruentatus (LATREILLE, 1802) and C. micans (NYLANDER, 1994, RADCHENKO 1992a, b, 1994c, d, 1996a, b, 1998, 1856) (TAYLOR 2006) are C. turkestanicus EMERY, 1887 COLLINGWOOD & AGOSTI 1996), as well as unpublished and C. xerxes FOREL, 1904, respectively; Messor galla data and keys for several genera of A. Radchenko. In all (MAYR, 1904) and Camponotus maculatus (FABRICIUS, cases the correct names of the taxa were verified accord- 1782) (TIRGARI & PAKNIA 2004) are M. intermedius SANT- ing to the catalogue of BOLTON & al. (2007) except for SCHI, 1927 and C. armeniacus ARNOLDI, 1967, respec- Tetramorium forte FOREL, 1904 and T. chefketi FOREL, tively; Tapinoma simrothi KRAUSSE, 1911 (PAKNIA & KAMI 1911. The software ArcView program was used to prepare 2007) is T. karavaievi EMERY, 1925; Tetramorium forte a map of all records of ants in Iran. FOREL, 1904 (ARDEH 1994) is a junior synonym of T. chef- keti FOREL, 1911 (see also CSÖSZ & al. 2007); Tetramori- Results um moravicum KRATOCHVÍL, 1941 (ALIPANAH & al. 1995) The proposed preliminary list of Iran comprises 110 ant was determined only as Tetramorium sp. now, but it is clear- species and subspecies from 26 genera of six subfamilies ly not T. moravicum. (Formicinae, Myrmicinae, Ponerinae, Dolichoderinae, Dory- linae and Aenictinae) (Tab. 1). The most speciose subfam- Excluded species ily is that of the Myrmicinae (52 species), while Aenicti- As our colleague Dr. Dobovikov, who checked material nae and Dorylinae include only one species each. The most recorded by FOREL (1904a) in ZISP, did not find the speciose genera in Iran are Camponotus, Cataglyphis and voucher specimens of Messor barbarus (LINNAEUS, 1767) Messor with 19, 14 and 13 species, respectively. By con- and M. capitatus (LATREILLE, 1798), we propose to ex- trast, only one species per genus was also recorded for clude both species from the Iranian list, especially as these Liometopum, Anochetus, Pachycondyla, and Ponera. Nine species are restricted to the Mediterranean Region. species have been taxonomically described from Iran in ear- lier studies (indicated by # in Tab. 1). Included species without vouchers One subfamily (Dorylinae), two genera and seven spe- The vouchers of Camponotus maculatus (FABRICIUS, 1782), cies were recorded for Iran for the first time: Camponotus thoracicus (FABRICIUS, 1804) and Para- Aphaenogaster gibbosa (LATREILLE, 1798): Mazandaran trechina vividula (NYLANDER, 1846) recorded by FOREL province, Miankaleh peninsula, Plangan, in shrubland (1904a) were not found in ZISP. However, as these spe- and grassland habitats, 14.IV.2005, leg. O. Paknia, 3 cies have been recorded from adjacent countries (COL- workers (ZMGU), det. A. Radchenko. LINGWOOD & AGOSTI 1996, DONISTHORPE 1950) and also Aphaenogaster kurdica RUZSKY, 1905: Golestan pro- because the latter one is a tramp species living in the tem- vince, Aliabad, near Kabudval waterfall, in forest habi- perate zones in houses and greenhouses, the presence of tat, 17.VI.2005, leg. O. Paknia, 8 workers (ZMGU), det. these species in Iran appears likely. We included these A. Radchenko. species in our list. Crematogaster bogojawlenskii RUZSKY, 1905: Khorasan The records of Tetramorium caespitum are based on Razawi province, Sarakhs, 27.IV.2007, leg. H. Hajiqan- the old keys and have to be reviewed by use of modern bar, (IZK), det. A. Radchenko. taxonomic approaches (see SCHLICK-STEINER & al. 2006,

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Tab. 1: Preliminary check-list of Iranian ant species. Genera and species within them are listed alphabetically. Listed are the valid names, region names (N = North, NW = Northwest, NE = Northeast, E = East, S = South, SW = Southwest, SE = Southeast, W = West; also see Fig. 1), any former names and junior synonyms and the collections where vouchers are kept. References are marked as follows: a = RADCHENKO & ALIPANAH (2004), b = CRAWLEY (1922), c = ALIPANAH & DEZHAKAM (2000), d = RADCHENKO (1996b, 1997a), e = MENOZZI (1927), f = ALIPANAH & al. (1995), g = TIRGARI & PAKNIA (2004), h = ALIPANAH & al. (2000), I = ALIPANAH (2004), j = FOREL (1904a), k = ARDEH (1994), l = SEIFERT (2003), m = RADCHENKO (1997b), n = KARAVAIEV (1924), o = EMERY (1906), p = PAKNIA & KAMI (2007), q = CRAWLEY (1920b), r = TAHMASEBI & ALIPANAH (2000), s = TAYLOR (2006), t = CRAWLEY (1920a), u = RADCHENKO (1994a), v = ANARAKI (1981), w = AKBARZADEH & al. (2004), x = FOREL (1904b), y = TIRGARI & PAKNIA (2005), z = ARNOLDI (1977). Asterisks at species names refer to those species which have been mentioned in the text, except new records; hatches refer to taxa described from Iran.

Scientific valid name Regions Former name or synonymies, Determinator References and material of Iran used in old literature deposition

Aenictus dlusskyi ARNOLDI, 1968 * N Radchenko a, IZK

Anochetus evansi CRAWLEY, 1922 * # NW Crawley b

Aphaenogaster gibbosa (LATREILLE, 1798) N Radchenko ZMGU

Aphaenogaster kurdica RUZSKY, 1905 N Radchenko ZMGU Aphaenogaster syriaca Emery, 1908 SW Collingwood c, HMIM

Camponotus aethiops (LATREILLE, 1798) N Radchenko d, HMIM

Camponotus armeniacus ARNOLDI, 1967 * S Radchenko d, ZMGU

Camponotus atlantis FOREL, 1890 N Ward, Collingwood k, HMIM or Munsee

Camponotus buddhae FOREL, 1892 N Radchenko d, ZMMU, IZK

Camponotus cecconii EMERY, 1908 N Collingwood h, HMIM

Camponotus fedtschenkoi MAYR, 1877 N Radchenko d, ZMMU, IZK

Camponotus fellah DALLA TORRE, 1893 * S Cook g, ZMGU

Camponotus gestroi EMERY, 1878 N Collingwood h, HMIM

Camponotus kopetdaghensis DLUSSKY & N Radchenko i, HMIM ZABELIN, 1985

Camponotus libanicus ANDRÉ, 1881 N Radchenko h, HMIM

Camponotus maculatus (FABRICIUS, 1782) * E C. maculatus r. cognatus Forel j (SMITH, F. 1858 )

Camponotus oasium FOREL, 1890 SE C. maculatus r. oasium Collingwood, j, c, HMIM Radchenko

Camponotus oertzeni FOREL, 1889 NW Radchenko d, IZK, ZMMU, MIZ

Camponotus sanctus FOREL, 1904 S Radchenko d, HMIM, ZMMU

Camponotus thoracicus (FABRICIUS, 1804) * E C. maculatus r. dichrous Forel j

Camponotus turkestanicus EMERY, 1887 * NE Radchenko d, HMIM, ZMGU

Camponotus turkestanus ANDRÉ, 1882 N Radchenko d, IZK, ZMMU

Camponotus vogti FOREL, 1906 N Collingwood h, k, HMIM

Camponotus xerxes FOREL, 1904 * # N C. maculatus r. xerxes Radchenko d, x, k, ZMGU, ZMMU

Cardiocondyla brachyceps SEIFERT, 2003 # S Seifert l

Cardiocondyla elegans EMERY, 1869 N Seifert, Radchenko l, s, HMIM

Cardiocondyla fajumensis FOREL, 1913 S Seifert l

Cardiocondyla mauritanica FOREL, 1890 S Seifert l

Cardiocondyla persiana SEIFERT, 2003 # S Seifert l

Cardiocondyla sahlbergi FOREL, 1913 NW Seifert l

Cardiocondyla stambuloffii FOREL, 1892 NW Seifert l

Cardiocondyla unicalis SEIFERT, 2003 # W Seifert l

Cataglyphis aenescens (NYLANDER, 1849) N, NE Myrmecocystus cursor r. tancrei Radchenko, Taylor j, p, s, HMIM

Cataglyphis altisquamis (ANDRE, 1881) N Radchenko m, ZMMU

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Cataglyphis bellicosus (KARAVAIEV, 1924) # N Myrmecocystus bicolor ssp. Radchenko n, IZK, ZMMU bellicosus

Cataglyphis bucharicus EMERY, 1925 N Radchenko m, ZMMU

Cataglyphis cuneinodis ARNOLDI, 1964 NW Radchenko m, ZMMU

Cataglyphis emeryi (KARAVAIEV, 1911) N Radchenko m, ZMMU

Cataglyphis foreli (RUZSKY, 1903) NE Myrmecocystus altiquamis r. Radchenko j, ZMGU foreli

Cataglyphis frigidus persicus (EMERY, 1906) # S Myrmecosystus frigidus var. Emery O persica

Cataglyphis lividus (ANDRÉ, 1881) NE, N, S Myrmecocystus albicans r. Collingwood, j, h, k, p, HMIM, ZMGU lividus Radchenko

Cataglyphis niger (ANDRÉ, 1881) N, S Myrmecocystus viaticus r. Collingwood, j, h, k, p, HMIM, ZMGU niger Radchenko

Cataglyphis nigripes ARNOLDI, 1964 W, NW Radchenko m, ZMMU, HMIM

Cataglyphis nodus (BRULLÉ, 1833) N, S Cataglyphis bicolor var. nodus Callingwood, e, h, k, p, ZMGU MENOZZI (1927) Radchenko

Cataglyphis ruber (FOREL, 1903) NW Radchenko m, ZMMU

Cataglyphis setipes (FOREL, 1894) S Radchenko m, HMIM

Crematogaster antaris FOREL, 1894 SW Collingwood C, HMIM

Crematogaster bogojawlenskii RUZSKY, 1905 NE Radchenko IZK

Crematogaster inermis MAYR, 1862 * N Collingwood f, HMIM

Crematogaster schmidti (MAYR, 1853) N C. scutellaris ssp. schmidti Crawley t

Crematogaster sorokini RUZSKY, 1905 W Radchenko i, HMIM

Crematogaster subdentata MAYR, 1877 N Ward, Collingwood k, HMIM or Munsee Dorylus sp. * SW Alipanah HMIM

Formica cunicularia LATREILLE, 1798 N Formica glauca RUZSKY, 1896 Collingwood h, HMIM

Formica lusatica SEIFERT, 1997 N Schultz, Seifert p, ZMGU

Formica rufibarbis FABRICIUS, 1793 NW Crawley, Anaraki q, v

Formica sanguinea LATREILLE, 1798 N Collingwood h, HMIM

Lasius alienus (FÖRSTER, 1850) NW Ward, Collingwood q, k, HMIM or Munsee

Lasius brunneus (LATREILLE, 1798) NW Crawley q

Lasius emarginatus (OLIVER, 1792) NW, N L. emarginatus var. nigro- Seifert q, ZMGU emarginatus

Lasius lasioides (EMERY, 1869) N Radchenko p, ZMGU

Lasius neglectus VAN LOON, BOOMSMA & N Schultz, Seifert p, ZMGU ANDRASFALVY, 1990 *

Lasius platythorax SEIFERT, 1991 SW Radchenko i, HMIM

Lasius turcicus SANTSCHI, 1921 N Collingwood, h, HMIM Radchenko

Lepisiota bipartita (SMITH, F., 1861) E, N Acantholepis frauenfeldi r. Collingwood j, HMIM bipartita

Lepisiota dolabellae (FOREL, 1911) N Ward, Collingwood k, HMIM or Munsee

Lepisiota semenovi (RUZSKY, 1905) N Radchenko r, HMIM

Leptothorax acervorum (FABRICIUS, 1793) N Taylor s, ZMGU

Liometopum microcephalum (PANZER, 1798) W Radchenko p, ZMGU

Messor caducus (VICTOR, 1839) N, NW, S Cook f, p, z, ZMMU

Messor concolor SANTSCHI, 1927 N Taylor s, ZMGU

Messor dentatus SANTSCHI, 1927 N Collingwood f, HMIM

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Messor denticulatus SANTSCHI, 1927 NE Radchenko z, HMIM

Messor ebeninus SANTSCHI, 1927 SW, N Collingwood f, HMIM

Messor incorruptus KUZNETSOV-UGAMSKY, W Radchenko i, HMIM 1929

Messor intermedius SANTSCHI, 1927 * S Messor semirufus var. inter- Radchenko e, ZMGU medius

Messor meridionalis (ANDRÉ, 1883) SW, N Ward, Collingwood k, HMIM or Munsee

Messor minor (ANDRÉ, 1883) W Radchenko HMIM

Messor rufotestaceus (FÖRSTER, 1850) S, SW Radchenko, c, ZMGU, HMIM Collingwood

Messor semirufus (ANDRÉ, 1883) SW M. barbarus r. semirufus Crawley t

Messor structor (LATREILLE, 1798) NW, N M. platyceras var. rubella Radchenko t, HMIM

Messor structor platyceras CRAWLEY, 1920 # NW M. platyceras Crawley t

Monomorium abeillei ANDRÉ, 1881 SW Radchenko c, HMIM

Monomorium destructor (JERDON, 1851) * SW Radchenko c, HMIM

Monomorium kusnezovi SANTSCHI, 1928 NE Radchenko i, HMIM

Monomorium nitidiventre EMERY, 1893 S Taylor s, ZMGU

Monomorium pharaonis (LINNAEUS, 1758) * S Radchenko e, ZMGU

Monomorium salomonis (LINNAEUS, 1758) S M. salmonis var. ? Menozzi e

Myrmica bergi RUZSKY, 1902 N, NW M. bergi var. fortior Radchenko t, IZK

Myrmica sabuleti MEINERT, 1861 N Collingwood k, HMIM

Pachycondyla sennaarensis (MAYR, 1862) * S, SE Cook, Paknia y, p, w, ZMGU

Paratrechina flavipes (F. SMITH, 1874) * SW Collingwood c, HMIM

Paratrechina longicornis (LATREILLE, 1802) * N Collingwood, f, p, HMIM, ZMGU Schultz

Paratrechina vividula (NYLANDER, 1846) * SE Prenolepis vividula Forel j

Pheidole pallidula (NYLANDER, 1849) NW Ph. pallidula subsp. arenarum Collingwood t, f, HMIM var. orientalis

Pheidole sinaitica MAYR, 1862 S Cook g, ZMGU

Pheidole teneriffana FOREL, 1893 * N, S Collingwood, Cook f, k, p, ZMGU, HMIM

Plagiolepis pallescens FOREL, 1889 N Collingwood h, HMIM

Plagiolepis taurica SANTSCHI, 1920 N Plagiolepis vindobonensis Ward, Collingwood k, HMIM LOMNICKI, 1925 or Munsee

Polyrhachis lacteipennis F. SMITH, 1858 * S, SE Cook, Radchenko g, HMIM, ZMMU, ZMGU Ponera sp. N Radchenko ZMGU

Solenopsis cf. fugax (LATREILLE, 1798) N Collingwood f, HMIM

Solenopsis cf. latro FOREL, 1894 N Collingwood f, HMIM

Tapinoma erraticum (LATREILLE, 1798) N, S T. erraticum subsp. nigerrimum Radchenko, Taylor t, s, ZISP, ZMGU

Tapinoma karavaievi EMERY, 1925 * N, S Radchenko HMIM, ZMGU

Temnothorax iranicus (RADCHENKO, 1994) # N Leptothorax iranicus Radchenko u, ZMMU, IZK

Temnothorax parvulus (SCHENCK, 1852) N Radchenko ZMGU

Tetramorium cf. caespitum (LINNAEUS, 1758) * N, S Cook t, p, ZMGU

Tetramorium chefketi FOREL, 1911 * N Collingwood f, HMIM

Tetramorium davidi FOREL, 1911 N Collingwood f, HMIM

Tetramorium ferox RUZSKY, 1903 S Cook g, ZMGU

Tetramorium inerme MAYR, 1877 NE Radchenko IZK Tetramorium sp. * N T. moravicum Radchenko HMIM

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Fig. 1: Map of Iran with the localities where ants were collected (black dots).

STEINER & al. 2006, CSÖSZ & al. 2007). However, based on centre of the country that lie in the vast Central Persian distribution maps of Tetramorium ants in the western Pale- desert basin. There are no species records at all from the arctic (see SCHLICK-STEINER & al. 2006) we left this spe- Eastern Iran mountain woodlands, the Kopet Dag wood- cies in the list as Tetramorium cf. caespitum. The situa- lands, Kopet Dag semi desert, Azerbaijan shrub desert and tion is similar with the old records of Lasius species (see steppe, and the desert and semi-desert areas in the centre SEIFERT 1992), that need to be reviewed by modern keys. and the east of Iran that comprise more than 50 % of the Nevertheless this work is beyond the scope of the present area of the country. paper, so we included these species in our preliminary list without revision. Discussion The record of Dorylinae is the first record of this sub- Although the first reports on Iranian ants were published family after the new clarification of this subfamily by BOL- more than 100 years ago, the ant fauna of this country re- TON (2003). mains poorly known. Most of the records are from the north of Iran, but many of these samples were collected in Distribution of records disturbed environments near human settlements that com- Regarding the geographical distribution of the species re- prise only a few percent of the country's surface area. There cords that had been examined in this study, we found a are only a few species reports from the extensive natural mismatching between the area of the locations that had habitats of the north, for example from the Caspian decid- been studied and the respective number of species that uous forests. The latter should have a rich ant fauna due to had been found there: e.g., 34 species of a total of 109 spe- old geological age of that forest region: it has been cov- cies were collected in natural or disturbed habitats of Teh- ered with forests since the late Tertiary period (ZOHARY ran province, though the territory of this province is less 1973). As a result, Tertiarian elements could have survived, than 2 % of the territory of the country. Fifteen species as it is known for the adjacent Talysh and Zuvand dis- were recorded from the Caspian forest region in the north tricts in south-eastern Azerbaijan (ARNOLDI 1930, 1948). of Iran that covers only 4 % of the country. Eight species Among the other regions that have been investigated were found in Zagros Mountains forest-steppe ecoregion only cursorily is the Nubo-Sindian desert and semi desert (about 20 % of entire area of the country). From the south ecoregion in the south and the southeast of the country. of Iran altogether 34 species were reported. In contrast, These areas are particularly interesting faunistically, as they there are only a few records from the wide Elburz Range are close to the boundaries of the Oriental and Afrotrop- forest-steppe and the Eastern Anatoloian Mountains in the ical zoogeographic regions. Four important ecoregions that north and northwest of Iran, or from eastern parts and the comprise a large part of the Irano-Anatolian biodiversity

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110 Chapter 6 - Ant Species List of Iran hotspot have not been studied at all. These are the Eastern Palaearctic ant genera will probably persist in a more com- Anatolian Mountains, Elburz forest-steppe, Kopet Dag prehensive species list to be presented in the future. woodlands and forest-steppe, and Zagros Mountains forest- steppe. For comparison, the well studied myrmecofauna of Acknowledgements the Turkmenistan's part of the Kopet Dag is one of the We thank Hamidreza Hajiqanbar for collecting and send- richest local ant fauna in Central Asia (DLUSSKY & ZABE- ing the material of two ant species, Crematogaster bogo- LIN 1985, DLUSSKY & al. 1990). jawlenskii and Tetramorium inerme that were new to Iran. This scarcity of data, especially from the border re- We thank Dr. Bernhard Seifert for his taxonomic comments, gions, does not permit to make a proper zoogeographical Dr. Cedric A. Collingwood for the identification of many analysis of the Iranian ant fauna at present. However, in specimens and Dr. Dubovikov for checking Forel's vouch- the future, when sufficient material will have been sam- ers in ZISP. We are indebted to John Fellowes, an ano- pled, the Iranian myrmecofauna needs to be compared with nymous referee and the editors for their useful sugges- those of the adjacent regions, e.g., Turkey, Armenia, Az- tions towards the improvement of our paper. We thank erbaijan, Turkmenistan, Afghanistan, and Arabian Penin- Doug Johns and Bryson Voirin for language correction of sula. Similarly, the presence of North African elements in the first and the second versions of the text, respectively. Iran (e.g., Camponotus fellah DALLA TORRE, 1893 or Crem- Zusammenfassung atogaster inermis MAYR, 1862) demonstrates a relation of the Iranian and North African desert faunas that has to be Nach einer umfassenden Durchsicht der Literatur und der confirmed by more intensive sampling. Untersuchung von Material aus eigenen Aufsammlungen The Iranian ant fauna includes eight "tramp" species, sowie aus Museen und Forschungsinstituten verschiede- which have been introduced by humans to many countries ner europäischer Länder, präsentieren wir die erste Arten- and in some cases have gained a worldwide distribution liste der Ameisen des Iran: 110 Arten aus 26 Gattungen und (see also MCGLYNN 1999, PAKNIA 2006, PAKNIA & KAMI sechs Unterfamilien der Formicidae (Formicinae, Myrmi- 2007): Lasius neglectus VAN LOON, BOOMSMA & AN- cinae, Ponerinae, Dolichoderinae, Dorylinae und Aenicti- DRASFALVY, 1990, Monomorium pharaonis (LINNAEUS, nae) wurden bislang gefunden. Die meisten der hier ge- 1758), M. destructor (JERDON, 1851), Pheidole teneriffana listeten Arten wurden im Norden des Landes gesammelt, FOREL, 1893, Paratrechina longicornis (LATREILLE, 1802), zumeist in anthropogen beeinflussten Gebieten in der Nähe P. flavipes (SMITH, F., 1874), P. vividula (NYLANDER, 1846) von Siedlungen. Eine Unterfamilie (Dorylinae), zwei Gat- and Pachycondyla sennaarensis (MAYR, 1862). tungen (Ponera LATREILLE, 1804 und Dorylus FABRICIUS, Almost half of the recorded species in the Iranian 1793), sowie sieben Arten der Formicidae (Aphaenogaster checklist belong to the genera Camponotus, Messor and gibbosa (LATREILLE, 1798), A. kurdica RUZSKY, 1905, Cataglyphis. Similar faunistic patterns are found in other Messor minor (ANDRE, 1883), Tapinoma karavaievi EME- arid Asian regions, e.g., Turkmenistan (DLUSSKY & al. RY, 1925, Temnothorax parvulus (SCHENCK, 1852), Tetra- 1990) and Saudi Arabia (COLLINGWOOD 1985, COLLING- morium inerme MAYR, 1877, und Crematogaster bogojaw- WOOD & AGOSTI 1996). The main reason for the high di- lenskii RUZSKY, 1905) wurden erstmalig für den Iran re- versity of these genera are the environmental conditions in gistriert. Die artenreichsten Gattungen des Iran sind Campo- Iran that comprise mainly arid and semi arid areas, the pre- notus MAYR, 1861 mit 19 Arten sowie Cataglyphis FOERS- ferred habitats of Messor, Cataglyphis and many Campo- TER, 1850 mit 14 und Messor FOREL, 1890 mit 13 Arten. notus species (DLUSSKY 1981, DLUSSKY & al. 1990, HÖLL- Zoogeographisch gesehen dominieren paläarktische Ele- DOBLER & WILSON 1990, ANDERSEN & CLAY 1996, AN- mente im Iran, allerdings wurden auch verschiedene Arten DERSEN & SPAIN 1996). der Orientalis und Afrotropis gefunden. Viele Landesteile The second reason for the dominance of those three gen- wurden bislang kaum oder gar nicht untersucht und der era in ant collections may be artificial: their members are Artenreichtum der Formicidae des gesamten Iran dürfte large and can be easily collected by anyone. In the majo- wesentlich größer sein. rity of the former studies, "direct hand collecting" was the main method. For this reason small-sized and cryptic References ants (e.g., Leptanillinae, Amblyoponinae, Dacetini, etc.) are AGOSTI, D. & ALONSO, L.E. 2000: The ALL Protocol: A stand- under-represented in the investigated material. Furthermore, ard protocol for the collection of ground-dwelling ants. In: social parasites are also missing in the presented species AGOSTI, D., MAJER, J.D., ALONSO, L.E. & SCHULTZ, T.R. list of Iran. To overcome this sampling bias and to establish (Eds.): Ants: standard methods for measuring and monitoring biodiversity. – Smithsonian Institution Press, London, pp. a reliable species list, thorough investigations of ant diver- 204-206. sity in all parts of Iran are urgently needed. They should be conducted by standard sampling methods, like direct col- AKBARZADEH, K., TIRGARI, S. & NATEGHPUR, M. 2004: First re- cord of presence of stinging ants Pachycondyla sennaarensis lecting from ant nests, pitfall traps, bait trapping and litter (Formicidae: Hymenoptera) in Sistan and Baluchestan pro- extraction with Winkler collectors (see AGOSTI & ALONSO vince. – Proceedings of 16th Iran Plant Protection Congress, 2000). Tabriz, p. 290. At last, we have to emphasize that the Iranian ant fauna ALIPANAH, H. 2004: Introduction of one subfamily and six new seems to be one of typically Palaearctic character. If we species of ants (Hym. Formicidae) from Iran. – Proceedings exclude the introduced species mentioned above, native of the 16th Iranian Plant Protection Congress, Tabriz, p. 136. members of only four tropical (Oriental or Afrotropical) ALIPANAH, H. & DEZHAKAM, M. 2000: Introduction of seven genera are found in Iran: Aenictus dlusskyi ARNOLDI, 1968, new species of ants (Hym. Formicidae) from Iran. – Proceed- Anochetus evansi CRAWLEY, 1922, Dorylus sp., and Poly- ings of the 14th Iranian Plant Protection Congress, Esfahan, rhachis lacteipennis SMITH, F., 1858. This dominance of p. 349.

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ALIPANAH, H., KHARAZI-PAKDEL, A. & MOGHADASSI, P. 1995: CRAWLEY, W.C. 1922: Formicidae – a new species and variety. Taxonomical study of Myrmicinae ants in Tehran. – Proceed- – Entomologist's Record and Journal of Variation 34: 85-86. th ings of the 12 Iranian plant protection congress, Karadj, p. 304. CSŐSZ, S., RADCHENKO, A. & SCHULZ, A. 2007: Taxonomic re- ALIPANAH, H., KHARAZI-PAKDEL, A. & MOGHADASSI, P. 2000: vision of the Palaearctic Tetramorium chefketi species com- Taxonomical study of Formicinae ants in Tehran. – Proceed- plex (Hymenoptera: Formicidae). – Zootaxa 1405: 1-38. th ings of the 14 Iranian plant protection congress, Esfahan, p. 350. 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PFEIFFER, M., SCHULTZ, R., RADCHENKO, A., YAMANE, S., WOY- RADCHENKO, A.G. 1997b: A review of ants of the genus Cata- CIECHOWSKI, M., ULYKPAN, A. & SEIFERT, B. 2007: A critical glyphis FOERSTER (Hymenoptera, Formicidae) from Asia. – En- checklist of the ants of Mongolia (Hymenoptera / Formicidae). tomologicheskoye Obozreniye 76: 424-442. (in Russian) – Bonner Zoologische Beiträge B 55: 1-8. RADCHENKO, A.G. 1998: A key to the identification of the ants of RADCHENKO, A. 2005: Monographic revision of the ants (Hyme- the genus Cataglyphis FOERSTER (Hymenoptera, Formicidae) of noptera, Formicidae) of North Korea. – Annales Zoologici 55: Asia. – Entomologicheskoe Obozrenie 77: 502-508. (in Russian) 127-221. RADCHENKO, A.G. & ALIPANAH, H. 2004: The first record of the RADCHENKO, A.G. 1992a: The ants of the genus Tetramorium subfamily Aenictinae (Hymenoptera: Formicidae) from Iran. 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SCHULTZ, R., RADCHENKO, A. & SEIFERT, B. 2006: A critical check- RADCHENKO, A.G. 1994a: New species of ants of the genus Lep- tothorax (Hymenoptera, Formicidae) from the southern and list of the ants of Kyrgyzstan (Hymenoptera: Formicidae). – eastern Palaearctic. – Zhurnal Ukrains'koho Entomolohichnogo Myrmecologische Nachrichten 8: 201-208. Tovarystva 2: 23-34. SEIFERT, B. 1992: A taxonomic revision of the Palaearctic mem- bers of the ant subgenus Lasius s. str. (Hymenoptera, Formi- RADCHENKO, A.G. 1994b: A taxonomic review of the scabrinodis- cidae). – Abhandlungen und Berichte des Naturkundemuseums group of the genus Myrmica LATREILLE (Hymenoptera, Formi- Görlitz 66: 1-67. cidae) of the Central and Eastern Palaearctic. – Zoologiches- kii Zhurnal 73: 75-82. (in Russian; English translation: Ento- SEIFERT, B. 2003: The ant genus Cardiocondyla (Insecta: Hyme- mological Review (Washington) 74: 116-124) noptera: Formicidae) – a taxonomic revision of the C. elegans, C. bulgarica, C. batesii, C. nuda, C. shuckardi, C. stambuloffii, RADCHENKO, A.G. 1994c: A key to species of the genus Myrmica C. wroughtonii, C. emeryi, and C. minutior species groups. – (Hymenoptera, Formicidae) of the central and eastern Palae- Annalen des Naturhistorischen Museums in Wien B 104: 203-338. arctic region. – Zoologicheskii Zhurnal 73: 130-145. (in Rus- sian; English translation: Entomological Review (Washington) STEINER, F.M., SCHLICK-STEINER, B.C. & MODER, K. 2006: Mor- 74: 154-169) phology-based cyber identification engine to identify ants of the Tetramorium caespitum/impurum complex (Hymenoptera: RADCHENKO, A.G. 1994d: A key for the identification of the Formicidae). – Myrmecologische Nachrichten 8: 175-180. genus Leptothorax MAYR (Hymenoptera, Formicidae) of the TAHMASEBI, G. & ALIPANAH, H. 2000: New report of Lepisiota Central and Eastern Palaearctic. – Zoologichesky Zhurnal 73: semenovi (Hym.: Formicidae) in Iran. – Journal of Entomolo- 146-158. (in Russian; English translation: Entomological Re- gical Society of Iran 22: 83-84. (in Persian) view (Washington) 74: 128-142) TARBINSKY, Y.S. 1976: The ants of Kirghizia. – Ilim Press, Frunze, RADCHENKO, A.G. 1995: A taxonomic review of the genus Lep- 217 pp. (in Russian) tothorax (Hymenoptera, Formicidae) of the Central and East- ern Palaearctic. Communication 3. Nylanderi, korbi, nassonovi TAYLOR, B. 2006: Ants of Iran. – , retrieved on 9 Jan- uary 2008. RADCHENKO, A.G. 1996a: The ants of the genus Plagiolepis MAYR (Hymenoptera, Formicidae) of the Central and Southern Palae- TERAYAMA, M. & CHOI, B.M. 1994: Ant faunas of Taiwan, Korea, arctic. – Entomologicheskoe Obozrenie 75: 178-187. (in Russian) and Japan. – Ari 18: 36. (in Japanese) TIRGARI, S. & PAKNIA, O. 2004: Additional records for the Irani- RADCHENKO, A.G. 1996b: Key to the ants of the genus Campo- an Formicidae fauna. – Zoology in the Middle East 32: 115-116. notus from Asian Palaearctic. – Zoologicheskii Zhurnal 75: 1195-1203. (in Russian; English translation: Entomological Re- TIRGARI, S. & PAKNIA, O. 2005: First record of ponerine ant (Pa- view (Washington) 76: 430-437) chycondyla sennaarensis) in Iran and some notes on its ecol- ogy. – Zoology in the Middle East 34: 67-70. RADCHENKO, A.G. 1997a: A review of ants of the subgenera Tan- aemyrmex, Colobopsis, Myrmamblis, Myrmosericus, Orthonoto- WU, J. & WANG, C. 1995: The ants of China. – China Forestry myrmex and Paramyrmamblis of the genus Camponotus MAYR Publishing House, Beijing, 214 pp. (in Chinese) (Hymenoptera, Formicidae) of the Asian part of Palaearctic. ZOHARY, M. 1973: Geobotanical foundations of the Middle East. – Zoologicheskii Zhurnal 76: 806-815. (in Russian) – Gustav Fischer Verlag, Stuttgart, 738 pp.

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113 ASIAN MYRMECOLOGY Volume 3, 29–38, 2010 ISSN 1985-1944 ©OMID PAKNIA,ALEXANDER RADCHENKO AND MARTIN PFEIFFER

New records of ants (Hymenoptera: Formicidae) from Iran

OMID PAKNIA1,ALEXANDER RADCHENKO2 AND MARTIN PFEIFFER1

1University of Ulm, Institute for Experimental Ecology, Albert Einstein Allee 11, 89069 Ulm, Germany, [email protected] and [email protected]

2Museum and Institute of Zoology of Polish Academy of Sciences, Wilcza str. 64, 00-679, Warsaw, Poland, [email protected]

Corresponding author’s email: [email protected]

ABSTRACT. The ant species list of Iran is far from complete. So far, only 110 species belonging to 26 genera have been recorded from Iran. For this study, we collected the majority of ant material in two periods of field work in spring and summer of 2007 and 2008. In total, we checked more than 35,000 specimens, and recorded 32 species and six genera new to Iran: Dolichoderus, Myrmecina, Proformica, Pyramica, Stenamma and Strongylognathus. Our new records from the Central Persian desert basins indicate that the ant fauna of this region prob- ably has more species in common with that of the Central Asian deserts than with the hot subtropical deserts of the Middle East and Arabian Peninsula.

Keywords: ants, faunistic, Dolichoderus, Myrmecina, Proformica, Pyramica, Stenamma, Strongylognathus, Iran

INTRODUCTION The Iranian ant fauna has been poorly investigated. So far, 110 species belonging to 26 genera have Iran is located in the mid-latitude band of arid been recorded from Iran (Paknia et al. 2008). In and semi-arid regions of the Old World, in this paper, we introduce 28 species that are new Southwest Asia. Western and especially for the Iranian fauna. This large number of new southwestern Asia comprises vast arid and semi- records, a short time after publishing a primary arid areas and represents a number of distinctive checklist for the country (Paknia et al. 2008) shows ecoregions that support numerous endemic plants the incompleteness of our myrmecological and animals (Zohary 1973; Olson et al. 2001). knowledge of this region. Biogeographically, this territory represents a transition zone between three regions: Palaearctic, MATERIALS AND METHODS Afrotropical and Oriental. Many of these unique ecoregions have not yet been studied intensively. The majority of ant samples were collected in two Despite more than one hundred years of periods of extensive field work in spring and myrmecology and the publication of a considerable summer, of 2007 and 2008, respectively. The first number of papers on the ant fauna of this region, field study was carried out along a transect from comprehensive faunistic investigations on ants have the north to the south of Iran through arid and semi- not been provided. Additionally, many genera of arid areas in four steppe and desert ecoregions. this region, such as Messor, Cataglyphis, The second block of fieldwork was conducted Camponotus and Monomorium, contain numerous inside the Central Persian desert basins in the taxa of ambiguous status and need modern revision. centre of Iran and in the Caspian Hyrcanian mixed

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30 New records of ants (Hymenoptera, Formicidae) from Iran forests in the north of Iran. Locations of the sample Keys for adjacent regions and taxonomic sites are indicated in Fig. 1. In the first year of revisions of different ant genera used in the present sampling, we employed pitfall trapping and bait study were as follows: Arnoldi (1948, 1977b: trapping for sampling ants, while in the second CentralAsia), Dlussky et al. (1990: Central Asia), year, we used only pitfall trapping for the Radchenko (1992a, 1992b, 1994a, 1994b, 1995a, collection of ants in arid and semi-arid areas. Both 1997a, 1997b: Central Asia), Arakelyan (1994: methods are suitable for rapid survey of open Central Asia), Collingwood and Agosti (1996: habitats, and well represent epigaeic ant species Arabian Peninsula), and Bolton (2000: Dacetini (Agosti & Alonso 2000). In forest areas, pitfall ants). The following ant collections were also used: trapping and Winkler collectors were applied. Museum and Institute of Zoology of Polish Additionally, nest samples were collected during Academy of Sciences, Warsaw, Poland (MIZ); both years of sampling. Moreover, a small number Zoological Museum of Moscow State University, of the samples were collected by hand during Russia (ZMMU); Zoological Institute of Russian different expeditions and in a few cases by Iranian Academy of Sciences, St. Petersburg, Russia students of biology. (ZISP); and Institute of Zoology of Ukrainian Samples were preserved in 96% alcohol, NationalAcademy of Sciences, Kiev, Ukraine (IZK). brought to the laboratory, and cleaned of debris In this paper, we define the following and soil. In total, 35,450 specimens were checked ambiguous geographical terms as follows: and sorted to morphospecies. Most of the Anatolia by its borders in Turkey, based on morphospecies were identified to species level, historical records from Asia Minor (all from within but some remained undetermined and were theAsian part of modern-day Turkey); the Arabian tentatively distinguished by species codes such Peninsula as Kuwait, Oman, Qatar, Saudi Arabia, as “sp. ir-abpari-01”. United Arab Emirates and Yemen; the territory of

Fig. 1. Map of Iran. Triangles show sampling locations of newly recorded ants: 1- Talysh, 2- Astaneh, 3-Abpari, 4- Nur, 5- Babolsar, 6- Ghaemshahr, 7- Gorgan, 8- Khoshyelagh, 9- Golestan National Park, 10- Bane, 11- Saggez, 12- Khojir, 13- Kavir, 14- Turan, 15- Naeen, 16- Siahkooh, 17- Robat, 18- Tabas, 19- Dena, 20- Kerman and 21- Mond.

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“Caucasus” as including Great Caucasus (i.e., Khoshyelagh Wildlife Refuge (36°47´41´´N, Russian North Caucasian republics from the Black 55°28´13´´E), 1534 m asl, 3.VI.2007, leg. Omid Paknia. sea to Caspian sea) as well as Armenia, Azerbaijan and Georgia, while “Transcaucasus” covers only Remarks: C. interjectus is distributed in Central the territory of Armenia,Azerbaijan and Georgia; Asia and Afghanistan (Dlussky et al. 1990; “CentralAsia” as including Kyrgysztan, Tajikistan, Radchenko 1997c) and recorded also from Iraq Turkmenistan and Uzbekistan (i.e., excluding (Wheeler & Mann 1916) and Dagestan in Russia Kazakhstan); and the “Middle East” as consisting (Kuznetsov-Ugamsky 1929). It inhabits mainly of Israel, Jordan, Lebanon, Palestine and Syria. mountain steppes; nests are built in soil, often The territory of Iraq is considered separately. under stones. All determined and undetermined species were deposited in the Antbase.net Collection Camponotus kurdistanicus Emery, 1898 (ABNC) managed by M. Pfeiffer and in the private collection of O. Paknia. Material: 13 , transitional region between Alborz Range forest steppe and Central Persian deserts, RESULTS Khojir National Park (35°38´54´´N, 51°43´44´´E), 1485 m asl, 1.V.2008, leg. Omid Paknia; 2 , Additional to the earlier published checklist Zagros Mountains forest steppe, Kurdistan, Bane (Paknia et al. 2008), we found 32 new ant species (~35°59´N, 45°53´E), ~1557 m asl., summer 2004, for Iran. Twenty-eight were identified to species leg. Shahin Mostafai. level and the other four were identified to genus level and given species codes. Our collecting efforts Remarks: This species was known from Anatolia, also resulted in the addition of six ant genera new Iraq and Azerbaijan (Emery 1898, 1925; Pisarski to Iran: Dolichoderus, Myrmecina, Proformica, 1971; Radchenko 1997c). Pyramica, Stenamma, and Strongylognathus. The new records are listed below. Camponotus libanicus Andre, 1881

Camponotus fallax (Nylander, 1856) Material: 8 , Alborz Range forest steppe, Khoshyelagh Wildlife Refuge (36°48´55´´N, Material: 4 , Caspian Hyrcanian mixed forest 55°31´54´´E), 1701 m asl, 8.VI.2007, leg. Omid Paknia. ecoregion, Gorgan, urban area (~36°50´23´´N, 54°26´19´´E), ~142 m asl, 1.IV.2006, leg. Omid Paknia. Remarks: C. libanicus is distributed in the Middle East countries and Anatolia (Radchenko 1997c). Remarks: the species is distributed in Europe This species inhabits dry and semi-dry areas. Nests (northwards to southern Sweden), the Caucasus, are built in soil. Anatolia and northwestern Kazakhstan, and also reported from the southern part of western Camponotus shaqualavensis Pisarski, 1971 Siberia. Iran is at the southeasternmost edge of the known distribution of this species, that Material: 4 , Nubo Sindian deserts, Kerman, urban inhabits mainly light and dry deciduous and area (~30°16´N, 57°04´E), ~1760 m asl, 26.V.2004, mixed forests, and often occurs in old parks and leg. Shiva Sadeghirad. orchards. It nests in dead parts of living trees or in wooden constructions (Radchenko 1997c; Remarks: C. shaqualavensis was described from Iraq Czechowski et al. 2002). (Pisarski 1971) and additionally was recorded from Turkey (Aktaç 1977; see also Radchenko 1997b). Camponotus interjectus Mayr, 1877 Camponotus staryi Pisarski, 1971 Material: 18 , Alborz Range forest steppe, Golestan National Park (37°20´56´´N, 56°14´46´´E), Material: 23 , Zagros Mountains forest steppe, 1256 m asl, 30.V.2007; 9 ,Alborz Range forest steppe, Dena Protected area (30°54´N, 51°25´E), 2683 m asl, 6.VII.2007, leg. Omid Paknia.

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32 New records of ants (Hymenoptera, Formicidae) from Iran

Remarks: C. staryi was described from Iraq Remarks: C. kurdistanicus was described from Iraq (Pisarski 1971). This is the first record of this (Pisarski 1965) and recorded additionally from species outside its type locality. Anatolia (Aktaç 1977; Radchenko 1997a).

Cataglyphis bergianus Arnoldi, 1964 Dolichoderus quadripunctatus (Linnaeus, 1771)

Material: 5 , Central Persian deserts, Tabas Material: 1 , Caspian Hyrcanian mixed forests, (33°36´53´´N, 57°05´36´´E), 1083 m asl, 16.V.2008, Talysh (37°42´19´´N, 48°53´14´´E), 66 m asl, leg. Omid Paknia. 6.VII.2008, leg. Omid Paknia.

Remarks: C. bergianus was known from Central Remarks: The genus Dolichoderus Lund comprises Asia, southern Kazakhstan and Afghanistan more than 130 species, distributed in all (Pisarski 1967, 1970; Radchenko 1997a). zoogeographic realms except Africa and Madagascar. Only one species, D. quadripunctatus, Cataglyphis cinnamomeus (Karavaiev, 1910) is known from the western Palaearctic. Its range covers central and southern Europe, central and Material: 3 , Central Persian deserts, Siahkooh southern parts of eastern Europe, the Caucasus, National Park (32°35´57´´N, 54°13´56´´E), 995 m Anatolia, southwest Siberia, and the Tien-Shan range asl, 25.V.2008, leg. Omid Paknia; 4 , Central in Central Asia and China. This genus is new to Iran. Persian deserts, Turan National Park (35°58´N, 56°04´E), 1191 m asl, 16.VI.2007, leg. Omid Paknia; Messor sp. (cf. M. oertzeni Forel, 1910) 6 , Central Persian deserts, South of Naeen (32°43´11´´N, 53°16´42´´E), 1372 m asl, 27.V.2008, Material: 3 , Zagros Mountains forest steppe, leg. Omid Paknia. Kurdistan, Saggez (~36°14´N, 46°15´E), ~1481 m asl, 2004, leg. Shahin Mostafai; 4 , Tehran, Remarks: Outside of Iran, C. cinnamomeus was urban area (~35°41´N, 51°25´E), ~1162 m asl, recorded from Central Asia, southern Kazakhstan 16.VII.2005, leg. Nasim Vakhideh; 5 , Mashad, (Karavaiev 1910) and Afghanistan (Pisarski 1967; urban area (~36°17´N, 59°35´E), ~985 m asl, leg. Radchenko 1997a). This species is the most Nayereh Ghafarian. thermophilous among Central Asian Cataglyphis, inhabiting mainly stony and clayey deserts. In Remarks: M. oertzeni is known from the Balkans mountain areas, it lives in dry parts of river valleys. and Anatolia (Agosti & Collingwood 1987; Atanassov & Dlussky 2002). It belongs to the Cataglyphis cugiai Menozzi, 1939 structor species-group, but reliable determination of many species from this group is impossible Material: 18 ,Alborz Range forest steppe, Golestan before a taxonomic revision is provided. National Park (37°20´57´´N, 56°14´44´´E), 1258 m asl, 30.V.2007, leg. Omid Paknia. Messor excursionis Ruzsky, 1905

Remarks: This species was known only from the Material: 21 , Central Persian deserts, Kavir northwest of India, Karakorum region (Menozzi National Park (34°45´14´´N, 52°10´07´´E), 1092 m 1939; Radchenko 1997a). asl, 7.VI.2008, leg. Omid Paknia.

Cataglyphis kurdistanicus Pisarski, 1965 Remarks: M. excursionis was recorded from Central Asia, Afghanistan and Mongolia Material: 54 , Zagros Mountains forest steppe, (Karavaiev 1910; Stärcke 1935; Pisarski 1967; Dena Protected Area (30°54´45´´N, 51°24´39´´E), Arnoldi 1970, 1977a; Dlussky et al. 1990; Pfeiffer 2695 m asl, 7.VII.2007, leg. Omid Paknia; 4 , Zagros et al. 2006). Mountains forest steppe, Kurdistan, Bane (~35°59´N, 45°53´E), ~1557 m asl, summer 2004, leg. Shahin Mostafai.

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Messor perantennatus Arnoldi, 1970 Remarks: M. variabilis is distributed in plains and foothills of Central Asia, where it inhabits different Material: 9 , Alborz Range forest steppe, kinds of deserts (Arnoldi 1970, 1977b; Dlussky et Khoshyelagh Wildlife Refuge (36°47´43´´N, al. 1990). 55°28´12´´E), 1536 m asl, 4.VI.2007, leg. Omid Paknia. Monomorium barbatulum Mayr, 1877

Remarks: This species was known only from Material: 3 , Central Persian deserts, Robat Posht Turkmenistan (Arnoldi 1970; Dlussky et al. 1990). Badam (32°57´22´´N, 55°32´09´´E), 1406 m asl, 17.V.2008, leg. Omid Paknia; 6 , Central Persian Messor subgracilinodis Arnoldi, 1970 deserts, Tabas (33°36´57´´N, 57°05´35´´E), 1074 m asl, 15.V.2008, leg. Omid Paknia. Material: 8 , Central Persian deserts, Turan National Park (35°57´48´´N, 56°05´35´´E), 1131 Remarks: M. barbatulum has a wide distribution m asl, 12.VI.2007, leg. Omid Paknia; 7 , Central from the southeast of Europe (Astrakhan Province Persian deserts, Tabas (33°36´44´´N, 57°05´45´´E), of Russia) to Central Asia and Afghanistan 1083 m asl, 16.V.2008, leg. Omid Paknia; 3 , (Pisarski 1967; Tarbinsky 1976; Dlussky et al. Central Persian deserts, Siahkooh National Park 1990) and the Arabian Peninsula (Collingwood & (32°35´55´´N, 54°13´59´´E), 996 m asl, 25.V.2008, Agosti 1996). leg. Omid Paknia; 5 , Central Persian deserts, Robat Posht Badam (32°57´26´´N, 55°32´08´´E) Monomorium dentigerum (Roger, 1862) 1401 m asl, 20.V.2008, leg. Omid Paknia. Material: 13 , Nobu Sindian deserts, Mond Remarks: M. subgracilinodis was recorded from Protected Area (28°07´08´´N, 51°23´50´´E), 9 m Turkmenistan (Arnoldi 1970, 1977b; Dlussky et asl, 14.VII.2007, leg. Omid Paknia. al. 1990). Remarks: M. dentigerum can be found all over Messor turcmenochorassanicus Arnoldi, 1977 the Middle East (Radchenko 1997d) and the Arabian Peninsula including Yemen (Collingwood Material: 65 , Zagros Mountains forest steppe, & Agosti 1996). Dena Protected Area (30°54´45´´N, 51°24´39´´E), 2627 m asl, 6.VII.2007, leg. Omid Paknia; 7 , Monomorium perplexum Radchenko, 1997 Caspian Hyrcanian mixed forests Nur, urban area (~36°33´N, 51°52´E), ~3 m asl, 17.III.2005, leg. Material: 24 , transitional region of Alborz Range Masud Tavakoli. forest steppe and Central Persian deserts, Khojir National Park (35°39´11´´N, 51°43´37´´E), 1462 Remarks: Messor turcmenochorassanicus was m asl, 1.V.2008, leg. Omid Paknia. recorded only from the Kopetdag Mountains in Turkmenistan (Arnoldi 1977b; Dlussky et al. 1990). Remarks: Outside of Iran, the species is distributed in the Transcaucasus,Anatolia, the Aegean Islands Messor variabilis Kuznetsov-Ugamsky, 1927 and Greece (Radchenko 1997d).

Material: 4 , Alborz Range forest steppe, Monomorium ruzskyi Dlussky et Zabelin, 1985 Golestan National Park (37°20´57´´N, 56°14´44´´E), 1254 m asl, 28.V.2007, leg. Omid Material: 3 , transitional region of Alborz Range Paknia; 6 , Central Persian deserts, Kavir forest steppe and Central Persian deserts, Khojir National Park (34°45´47´´N, 52°10´24´´E), 1051 National Park (35°38´56´´N, 51°43´45´´E), 1466 m m asl, 23.VI.2007, leg. Omid Paknia. asl, 3.V.2008, leg. Omid Paknia.

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34 New records of ants (Hymenoptera, Formicidae) from Iran

Remarks: M. ruzskyi was recorded from the Siberia and Kazakhstan to Tuva in Siberia (Dlussky Transcaucasus, Turkmenistan and Uzbekistan 1969), the Central Asian Mountains (Dlussky et (Dlussky & Zabelin 1985; Arakelyan 1994). al. 1990). This genus is new to Iran.

Myrmecina sp. ir-abpari-01 and ir-ghaemshahr-01 Pyramica sp. ir-golestan-01 and ir-ghaemshahr- 01 (argiola-group) Material: 8 , Caspian Hyrcanian mixed forests, Ghaemshahr (36°22´16´´N, 52°50´53´´E), 155 m Material: 1 , Caspian Hyrcanian mixed forests, asl, 4.VI.2008, leg. Omid Paknia; 4 , Caspian Ghaemshahr (36°22´16´´N, 52°50´53´´E), 155 m Hyrcanian mixed forests, Abpari (36°30´05´´N, asl., 4.VI.2008, leg. Omid Paknia. ; 1 , Caspian 51°55´58´´E), 308 m asl, 22.VI.2008, leg. Omid Hyrcanian mixed forests, Golestan National Park Paknia. (37°24´04´´N, 55°48´04´´E), 520 m asl, 11.VI.2008, leg. Omid Paknia. Remarks: The genus Myrmecina Curtis consists of about 40 described species distributed in the Remarks: In the last ten years, the systematics of Holarctic, South and Southeast Asia,Australia and the tribe Dacetini, to which Pyramica Roger SouthAmerica. It is absent in the Afrotropical and belongs, was cardinally changed. First of all, Malagasy Regions. The highest speciosity occurs Bolton (1999) revived the name Pyramica from in the Oriental Region. In the western Palaearctic, synonymy and proposed to consider it as a senior there are four known species distributed from the synonym of more than 20 generic names. A year Iberian Peninsula and Algeria to the Transcaucasus later, he published a huge taxonomic revision of and the Middle East. This genus is new to Iran. this tribe, describing several hundred new species (Bolton 2000). As a result, Pyramica is now one Myrmica specioides Bondroit, 1918 of the world’s biggest ant genera that includes more than 300 species; its range encompasses the Material: 8 , Caspian Hyrcanian mixed forests, whole world, with the overwhelming majority of Babolsar, in urban area (36°42´30´´N, 52°38´04´´E), species distributed in the tropics. This genus is 10 m asl, 18.IX.2002, leg. Omid Paknia. new to Iran. Several Pyramica species were recorded Remarks: This species has a very wide distribution from the Mediterranean region of Europe and Africa, that includes Europe (northwards to the south of the Middle East, Anatolia and the Transcaucasus, England and Denmark), the Caucasus, Anatolia, while it is not yet known from Central Asia (e.g., Turkmenistan, southwest Siberia and northern Arakelyan & Dlussky 1991;Arakelyan 1994; Bolton Kazakhstan, eastwards to the Altai Mountains et al. 2006; Radchenko 2007). (Radchenko & Elmes 2004; Radchenko 1994b ); it It is necessary to note that in the latest was introduced to North America (Jansen & revision of the tribe Dacetini (Baroni Urbani & Radchenko 2009). De Andrade 2007), the name Pyramica is considered a junior synonym of Strumigenys, but Proformica epinotalis Kusnetsov-Ugamsky, in this paper, we retain Pyramica until there is a 1927 definitive opinion on this question.

Material: 19 ,Alborz Range forest steppe, Golestan Stenamma sp. ir-talysh-01 National Park (37°20´54´´N, 56°14´42´´E), 1261 m asl, 29.V.2007, leg. Omid Paknia; 11 ,Alborz Range Material: 1 , Caspian Hyrcanian mixed forests, forest steppe, Khoshyelagh Wildlife Refuge Talysh (37°40´44´´N, 48°48´27´´E), 869 m asl, (36°46´24´´N, 55°22´42´´E), 1707 m asl, 6.VI.2007, leg. 7.VII.2008, leg. Omid Paknia. Omid Paknia. Remarks: Stenamma is a small genus, comprising Remarks: This species is distributed from Romania about 50 species distributed mainly in the in the west through southeast Europe, southwest Holarctic Region, but some of which penetrated

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Omid Paknia, Alexander Radchenko, and Martin Pfeiffer 35

into Central America, India and Pakistan Tetramorium armatum Santschi, 1927 (Himalayan region), and Southeast Asia. The highest species richness of this genus occurs in Material: 14 ,Alborz Range forest steppe, Golestan the rather warm and humid deciduous Holarctic National Park (37°20´36´´N, 56°14´57´´E), 1226 m asl, forests, many of which can be considered relict 28.V.2007, leg. Omid Paknia; 7 , Central Persian habitats. Stenamma is also quite speciose in deserts, Kavir National Park (34°45´47´´N, Caucasian and Anatolia forests (Arnoldi 1975; 52°10´24´´E), 1045 m asl, 21.VI.2007, leg. Omid DuBois 1998). This genus is new to Iran. Paknia; 6 , Central Persian deserts, Turan National Park (35°58´22´´N, 56°04´43´´E), 1174 m asl, Strongylognathus sp. ir-astaneh-01 15.VI.2007, leg. Omid Paknia.

Material: 3 , Caspian Hyrcanian mixed forests, Remarks: T. armatum is morphologically close to Astaneh, rural area (~37°15´N, 49°56´E), ~0 m asl, T. inerme Mayr, but in contrast to the latter, which summer 2004, leg. Faeze Mohammaddoost. lives in the Central Asian plains, it has been found predominantly in mountains, of Central Asia, the Remarks: The genus Strongylognathus is new to Transcaucasus, Afghanistan and Mongolia Iran. It is one of few genera endemic to the (Dlussky et al. 1990; Radchenko 1992b). Palaearctic Region. It comprises about 25 species, centred in the western Palaearctic (the range being Tetramorium schneideri Emery, 1898 Europe, northwest Africa, Anatolia, the Middle East, Kazakhstan, Central Asia and southwest Material: 21 ,Alborz Range forest steppe, Golestan Siberia), and only three species are known from National Park (37°20´36´´N, 56°14´57´´E), 1226 the eastern Palaearctic (China, Korea and Japan) m asl, 28.V.2007, leg. Omid Paknia. (e.g., Radchenko 1985, 1991; Radchenko 1995b; Wei et al. 2001; Japanese Ant Database Group Remarks: T. schneideri was recorded in the plains 2003; Radchenko 2005). All Strongylognathus and foothills of Central Asia and Afghanistan species are permanent social parasites in nests of (Dlussky et al. 1990; Radchenko 1992b). the Tetramorium Mayr species. Tetramorium striativentre Mayr, 1877 Temnothorax anodonta (Arnoldi, 1977) Material: 16 , Central Persian deserts, Daranjir Material: 5 , Caspian Hyrcanian mixed forests, Protected Area (32°26´18´´N, 55°01´21´´E), 1227 Golestan National Park (37°23´53´´N, 55°48´01´´E), m asl, 19.V.2008, leg. Omid Paknia; 2 , Central 500 m asl, 11.VI.2008, leg. Omid Paknia. Persian deserts, Siahkooh National Park (32°35´55´´N, 54°13´59´´E), 998 m asl, 25.V.2008, Remarks: Outside Iran, T. anodonta was recorded leg. Omid Paknia; 26 , Central Persian deserts from Armenia (Arnoldi 1977a; Arakelyan 1994; (32°57´23´´N, 55°32´08´´E), 1392 m asl, Radchenko 1995a) . 17.V.2008, leg. Omid Paknia; 4 , Central Persian deserts, Tabas (33°36´44´´N, 57°05´46´´E), 1083 Temnothorax nadigi (Kutter, 1925) m asl, 14.V.2008, leg. Omid Paknia; 22 , Central Persian deserts, South of Naeen (32°43´11´´N, Material: 3 , Caspian Hyrcanian mixed forests, 53°16´39´´E), 1372 m, asl, 26.V.2008, leg. Omid Abpari (36°30´07´´N, 51°55´58´´E), 315 m asl, Paknia; 23 , transitional region of Alborz Range 28.VI.2008, leg. Omid Paknia. forest steppe and Central Persian deserts, Khojir National Park (35°38´56´´N, 51°43´45´´E), 1466 Remarks: Known from southern Europe, Anatolia, m asl, 29.IV.2008, leg. Omid Paknia; 32 , Central the Transcaucasus and Turkmenistan (including Persian deserts, Kavir National Park (34°45´40´´N, its junior synonyms Leptothorax caucasicus 52°10´19´´E), 1057 m asl, 7.V.2008, leg. Omid Paknia; Arnoldi and L. hasardaghi Dlussky (Radchenko 39 , Central Persian deserts, Turan National Park 1995a; Czechowski et al. 2002). (35°58´22´´N, 56°04´43´´E), 1176 m asl, 17.VI.2007, leg. Omid Paknia.

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36 New records of ants (Hymenoptera, Formicidae) from Iran

Remarks: Inhabits mainly mountains of Central Asia Apart from Proformica, all the newly and Afghanistan, whereas the morphologically recorded genera (Dolichoderus, Myrmecina, similar T. schneideri lives in plains (Dlussky et al. Pyramica, Stenamma and Strongylognathus) 1990; Radchenko 1992b). were collected from this region. As the natural history (fauna and flora) of the Caspian Hyrcanian DISCUSSION mixed forests is close to those of the Caucasian and Anatolian regions, the ant fauna of the region Iran’s ant species list is far from complete. Iran is most likely related to them. Interestingly, is a vast country with a total area of 1.6 million specimens of three ant genera, Dolichoderus, square kilometres. Although arid and semi-arid Myrmecina and Pyramica were extracted by areas cover nearly half of the country, Iran also Winkler collector from the leaf litter of Caspian includes high mountains with alpine areas Hyrcanian mixed forests. This finding requires us (Noroozi et al. 2008), broadleaf forest in the to make further efforts to explore the litter layer southern coastal plains of the Caspian forests, in such forest ecosystems. Future intensive studies and steppe forests in the north and west (Zohary in the Caspian Hyrcanian area, and in the east, 1973). Thus, the 138 recorded species for the ant southeast and southwest of the country, are most fauna of Iran is still far from the true number. likely to reveal new records or even new species Additionally, our sampling sites were limited to for science. the north and the centre of Iran and no samples In closing, we emphasise the need for were collected from the east, the southeast or the fundamental studies in this part of Asia and invite southwest of the country (Fig. 1). As a result, it all myrmecologists, especially those from Asia, is not yet possible to draw complete and correct to take part in this job. biogeographic conclusions about the overall ant fauna of Iran. Nevertheless, the recorded species ACKNOWLEDGEMENTS give clues about how the ant fauna is biogeographically structured in the areas we have We thank all students named in this paper for the substantially sampled, like the north and centre contribution of specimens. Also, we are grateful to of Iran. the staff of the Department of Environment of Iran Many species on this list were recorded for the assistance and help that allowed this study from the north and centre of the Central Persian to be conducted. We also wish to thank Dr desert basins (Fig. 1), where the highest sampling Yoshimura, Dr Eguchi and one anonymous reviewer effort was applied. The deserts and semi-deserts who made very valuable and helpful suggestions. of Iran represent a transitional zone between the We are indebted to John Fellowes for his useful hot and cold deserts of Asia (Breckle 2002). The suggestions and for language correction of the text. Central Persian desert basins, except in the Financial support for the PhD study of O. Paknia in southeast of the region, are characterised by cold Germany, by the German Academic Exchange winters and an annual average occurrence of 60 Service (DAAD), is gratefully acknowledged. frost days, and thus can be likened to the belt of Central Asian deserts. Our new records from the REFERENCES Central Persian desert basins indicate that the ant fauna of this region has more species in common Agosti D and Alonso L, 2000. The ALL protocol: a with that of the Central Asian deserts (e.g., C. standard protocol for the collection of ground- bergianus, M. subgracilinodis, M. variabilis and dwelling ants. In: Agosti D, Majer J, Alonso L, and M. ruzsky) than with those of the hot subtropical Schultz T (eds), Ants: Standard Methods for Measuring and Monitoring Biodiversity, 204–6. deserts of the Middle East and Arabian Peninsula. Washington, D.C: Smithsonian Institution Press. The Caspian Hyrcanian mixed forests are Agosti D and Collingwood CA, 1987. A provisional list of geologically old and preserve the last remnants of the Balkan ants (Hym. Formicidae) and a key to the primary temperate deciduous broadleaved forests worker caste. I. Synonymic list. Mitteilungen der worldwide, existing there since the late Tertiary Schweizerischen Entomologischen Gessellschaft or period (Zohary 1973; Ramezani et al. 2008). Bulletin de la Societe Entomologique Suisse. 60:51–62.

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Aktaç N, 1977. Studies on the myrmecofauna of Turkey Collingwood CA and Agosti D, 1996. Formicidae I. Ants of Siirt, Bodrum and Trabzon. Istanbul (Insecta: Hymenoptera) of Saudi Arabia (Part 2). Universitesi Fen Fakultesi Mecmuasi. Seri B Fauna Saudi Arabia 15:300–85. 41:115–35. Czechowski W, Radchenko A and Czechowska W, Arakelyan GR, 1994. Fauna of the Republic of Armenia. 2002. The Ants (Hymenoptera: Formicidae) of Hymenopterous Insects. Ants (Formicidae). [in Poland. Museum and Institute of Zoology PAS, Russian]. Gitutium, Erevan, p. 153. Warsaw, p200. Arakelyan GR and Dlussky GM, 1991. Dacetine ants Dlussky GM, 1969. Ants of the genus Proformica (Hymenoptera: Formicidae) of the USSR. [in Russian]. Ruzs. of the USSR and contiguous countries Zoologicheskii Zhurnal 70:149–52. (Hymenoptera: Formicidae). [in Russian]. Arnoldi KV, 1948. Ants of Talysh and the Diabar Zoologicheskii Zhurnal 48:218–232. depression. Their importance for the characterization Dlussky GM, Soyunov OS and Zabelin SI, 1990. Ants of communities of terrestrial invertebrates and for of Turkmenistan. Ashkhabad: Ylym Press, p273. historical analysis of the fauna. [in Russian]. Trudy Dlussky GM and Zabelin SI, 1985. Ant fauna Zoologicheskogo Instituta Akademii Nauk SSSR (Hymenoptera: Formicidae) of the River Sumbar 7(2):206–62. Basin (south-west Kopetdag). [In Russian]. In: Arnoldi KV, 1970. New species and races of the ant Nechaevaya NT (ed), The Vegetation and Animal genus Messor (Hymenoptera: Formicidae). [in World of Western Kopetdag. [In Russian] Ylym, Russian]. Zoologicheskii Zhurnal 49:72–88. Ashkhabad. 208–46. Arnoldi KV, 1975. A review of the species of the genus DuBois MB, 1998. A revision of the ant genus Stenamma (Hymenoptera: Formicidae) of the USSR Stenamma in the Palaearctic and Oriental regions and description of new species. [in Russian]. (Hymenoptera: Formicidae: Myrmicinae). Zoologicheskii Zhurnal 54:1819–829. Sociobiology 32:193–403. Arnoldi KV, 1977a. New and little known ant species Emery C, 1898. Beiträge zur Kenntniss der of the genus Leptothorax Mayr (Hymenoptera: palaearktischen Ameisen. Öfversigt af Finska Formicidae) in the European USSR and the Vetenskaps-Societetens Förhandlingar 20:124–51. Caucasus. [in Russian]. Entomologicheskoye Emery C, 1925. I Camponotus (Myrmentoma) paleartici Obozreniye 56:198–204. del gruppo lateralis. Rendiconti delle Sessioni della Arnoldi KV, 1977b. Review of the harvester ants of the Reale Accademia delle Scienze dell’Istituto di genus Messor (Hymenoptera: Formicidae) in the Bologna Classe di Scienze Fisische 29:62–72. fauna of the USSR. [in Russian]. Zoologicheskii Jansen G and Radchenko A, 2009. Myrmica specioides Zhurnal 56:1637–48. Bondroit—a new invasive ant species in the USA? Atanassov N and Dlussky GM, 2002. Fauna Bulgaria Biological Invasions 11:253–56. 22. Hymenoptera, Formicidae [in Bulgarian]. Japanese Ant Database Group, 2003. Ant Image Bulgarian Academy of Sciences, Sofia, p. 310. Database 2003. Japanese Ant Database Group, Baroni Urbani C and De Andrade ML, 2007. The ant Sendai. tribe Dacetini: limits and constituent genera, with Karavaiev V, 1910. Nachtrag zu meinen “Ameisen aus descriptions of new species (Hymenoptera: Transcaspien und Turkestan”. Russkoye Formicidae). Annali del Museo Civico di Storia Entomologicheskoye Obozreniye 9:268–72. Naturale Giacomo Doria 99:1–191. Kuznetsov-Ugamsky NN, 1929. Die Ameisenfauna Bolton B, 1999. Ant genera of the tribe Dacetonini Daghestans. Zoologischer Anzeiger 83:34–45. (Hymenoptera: Formicidae). Journal of Natural Menozzi C, 1939. Formiche dell’Himalaya e del History 33:1639–89. Karakorum raccolte dalla Spedizione italiana Bolton B, 2000. The ant tribe Dacetini. Memoirs of the comandata da S. A. R. il Duca di Spoleto (1929). American Entomological Institute 65:1–1028. Atti della Società Italiana di Scienze Naturali e Bolton B, Alpert G, Ward PS and Naskrecki P, 2006. del Museo Civile di Storia Naturale, Milano Bolton’s Catalogue of Ants of the World: 1758– 78:285–345. 2005, CD-ROM. Cambridge, MA: Harvard Noroozi J, Akhani H and Breckle SW, 2008. University Press. Biodiversity and phytogeography of the alpine Breckle SW, 2002. Salt deserts in Iran and Afghanistan. flora of Iran. Biodiversity and Conservation In: Barth HJ and Boer B (eds), Sabkha Ecosystems. 17:493–521. Dordrecht, Kluwer Academic Publishers, p368.

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Olson DM, Dinerstein E, Wikramanayake ED et al., Radchenko AG, 1995a. A review of the ant genus 2001. Terrestrial ecoregions of the world: a new Leptothorax (Hymenoptera: Formicidae) of the map of life on earth. Bioscience 51(11):933–38. central and eastern Palearctic. Communication 3. Paknia O, Radchenko A, Alipanah H and Pfeiffer M, Groups nylanderi, korbi, nassonovi, and susamyri. 2008. A preliminary checklist of the ants [In Russian]. Vestnik Zoologii 1995(4):3–11. (Hymenoptera: Formicidae) of Iran. Myrmecological Radchenko AG, 1995b. Strongylognathus potanini sp. News 11:151–59. n. - a new ant species from China. [In Russian]. Pfeiffer M, Schultz R, Radchenko A, et al., 2006. A Zhurnal Ukrains’kogo Entomolohichnogo critical checklist of the ants of Mongolia Tovaristva 1(3-4):57–58. (Hymenoptera: Formicidae). Bonner Zoologische Radchenko AG, 1997a. Review of ants of the genus Beitraege 55(1):1–8. Cataglyphis Foerster (Hymenoptera: Formicidae) Pisarski B, 1965. Les fourmis du genre Cataglyphis of Asia. [In Russian]. Entomologicheskoye Foerst. en Irak (Hymenoptera: Formicidae). Bulletin Obozrenie 76:424–42. de L’Academie Polonaise des Sciences Série des RadchenkoAG, 1997b. Review of ants of the subgenera Sciences Biologiques 13:417–22. Tanaemyrmex,Colobopsis,Myrmamblis,Myrmosericus, Pisarski B, 1967. Fourmis (Hymenoptera: Formicidae) Orthonotomyrmex and Paramyrmamblis of the genus d’Afghanistan récoltées par M. Dr. K. Lindberg. Camponotus (Hymenoptera: Formicidae) in the Asian Annales Zoologici 24:375–425. Palearctic. [In Russian.]. Zoologichesky Zhurnal Pisarski B, 1970. Beiträge zur Kenntnis der Fauna 76:806–15. Afghanistans. Formicidae, Hymenoptera. Casopis Radchenko AG, 1997c. Review of ants of the subgenus Moravského Musea Brne 54 (Suppl.): 305–26. Myrmentoma genus Camponotus (Hymenoptera: Pisarski B, 1971. Les fourmis du genre Camponotus Formicidae) of the Asian Palearctic. [In Russian.]. Mayr (Hymenoptera: Formicidae) d’Iraq. Bulletin Zoologicheskii Zhurnal 76:703–11. de L’Academie Polonaise des Sciences Série des RadchenkoAG, 1997d. Review of the ants of scabriceps Sciences Biologiques 19:727–31. group of the genus Monomorium Mayr Radchenko A, 1985. Ants of the genus Strongylognathus (Hymenoptera: Formicidae). Annales Zoologici (Hymenoptera: Formicidae) in the European part of 46:211–24. the USSR. [In Russian]. Zoologicheskii Zhurnal Radchenko AG and Elmes GW, 2004. Taxonomic notes 64:1514–23. on the scabrinodis-group of Myrmica species Radchenko A, 1991. Ants of the genus Strongylognathus (Hymenoptera: Formicidae) living in Eastern Europe (Hymenoptera: Formicidae) of the USSR fauna. and western Asia, with a description of a new species [In Russian]. Zoologicheskii Zhurnal 70:84–90. from Tien Shan. Proceedings of the Russian Radchenko A, 2005. Monographic revision of the ants Entomological Society 75(1):222–33. (Hymenoptera: Formicidae) of North Korea. Ramezani E, et al., 2008. The late-Holocene vegetation Annales Zoologici 55(2):127–221. history of the Central Caspian (Hyrcanian) forests Radchenko A, 2007. Fauna Europea: Formicidae. In: of northern Iran. Holocene 18(2):307–21. Noyes J (Ed.): Fauna Europea: Hymenoptera: StärckeA, 1935. Zoologie. Formicidae. Wissenschaftliche Apocrita. Fauna Europea version 1.3, http:// Ergebnisse der Niederländischen Expedition in den www.faunaeur.org, accesed on 15 May 2007. Karakorum 1:260–9. Radchenko AG, 1992a. Ants of the genus Tetramonium Tarbinsky YS, 1976. The ants of Kirghizia. [in (Hymenoptera: Formicidae) of the USSR fauna. Russian]. Ilim, Frunze, p217. Report 1. [in Russian]. Zoologicheskii Zhurnal Wei C, Xu ZH and He H, 2001. A new species of 71:39–49. the ant genus Strongylognathus Mayr (Hymenoptera: Radchenko AG, 1992b. Ants of the genus Tetramorium Formicidae) from Shaanxi, China. Entomotaxonomia (Hymenoptera: Formicidae) of the USSR fauna. 23:68–70. Report 2. [In Russian]. Zoologicheskii Zhurnal Wheeler WM and Mann WM, 1916. The ants of the 71:50–58. Phillips Expedition to Palestine during 1914. Radchenko AG, 1994a. A key to species of the Bulletin of the Museum of Comparative Zoology Leptothorax (Hymenoptera: Formicidae) from at Harvard University 60:167–74. central and eastern Palearctic. [in Russian]. Zohary M, 1973. Geobotanical foundations of the Zoologicheskii Zhurnal 73:146–158. Middle East. Gustav Fischer Verlag, Stuttgart, Radchenko AG, 1994b. A key to the ants of the genus p738. Myrmica (Hymenoptera, Formicidae) of the Central and Eastern Palaearctic. [In Russian with English summary]. Zoologicheskii Zhurnal 73:130–45.

123 A N N A L E S Z O O L O G I C I (Warszawa), 2010, 60(1): 69-76

TWO NEW SPECIES OF THE GENUS CATAGLYPHIS FOERSTER, 1850 (HYMENOPTERA: FORMICIDAE) FROM IRAN

ALEXANDER RADCHENKO1 and OMID PAKNIA2

1Museum and Institute of Zoology, Polish Academy of Sciences, 64, Wilcza str., 00-679, Warsaw, Poland; e-mail: [email protected] 2Institute of Experimental Ecology, University of Ulm, Albert-Einstein Allee 11, D-89069 Ulm, Germany; e-mail: [email protected]

Abstract.— Two new species, Cataglyphis stigmatus sp. nov. and C. pubescens sp. nov. are described based on workers from Iran. The first species belongs to the bicolor species- group and clearly differs from all known species of this group by its yellow colour (except of C. lunaticus), but well distinguishes from the latter by the longer scape, by the lower propodeum, which dorsal surface is distinctly longer than the posterior one, by the less abundant standing hairs on the alitrunk and petiole, and especially by the much longer propodeal spiracles. Taxonomic position of C. pubescens is less clear, it shares features of the cursor-, emeryi- and emmae-groups, while differs from all species of these groups by the dense and long depressed pubescence on the head and alitrunk. 

Key words.— Ants, Formicidae, Formicinae, Cataglyphis stigmatus, C. pubescens, new species, Iran.

INTRODUCTION 18 species of the genus Cataglyphis have been recorded for Iran till recently (Paknia et al. 2008, Cataglyphis is one of the keystone ant genera in 2009), but in the newly collected material by one of the arid zones of the Old World. It is distributed mainly in co-authors (O. Paknia) we found specimens that belong Palaearctic, while several species dwell in deserts and to two new species, which are described below. One of semi-deserts of Afrotropical and Oriental Regions (In- them belongs to the bicolor species-group and well dif- dia and Pakistan). More than 100 species are known in fers from all members of this group by its totally yellow this genus till now, and even 2 social parasites were colour, except of C. lunaticus Baroni Urbani describ- described (Agosti 1994, Radchenko 1997b, Bolton et al. ed from Turkey. The second new species most probably 2007). is a member of the cursor species-group and has Members of this genus are large (up to 13 mm) ants, unique characteristics in its dense, rather long and and all of them inhabit open dry habitats (steppes, coarse, silverish pubescence of the body. stony mountain slopes, various types of deserts and semi-deserts, etc.), reaching in mountains up to 3500– 3700 m a.s.l. Formerly genus Cataglyphis has been divided to MATERIAL AND METHODS several subgenera (e.g., see Bolton 1995), but more recently subgeneric division was refused and the genus Material was collected from arid areas of the Cen- was separated into several species groups and species tral and Southern Iran (Fig. 1) during two field trips in complexes within them (Agosti 1990, Radchenko 1997a). spring and summer 2007 and 2008.

PL ISSN 0003-4541 © Fundacja Natura optima dux doi: 10.3161/000345410X499533

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70 A. RADCHENKO and O. PAKNIA

Comparative materials, including type specimens of from the lower margin of eye to the articula- many species were analysed in the course of this work. tion with mandible, These materials are preserved in the following Muse- AL – diagonal length of alitrunk (seen in profile), ums and Institutions: Zoological Museum of Moscow measured from the anterior end of propodeum State University, Russia (ZMMU); Zoological Institute to the posterior margin of propodeal lobes, of Russian Academy of Sciences, St.-Petersburg, Rus- PnW – maximum width of pronotum in dorsal view, sia (ZISP); Institute of Zoology of Ukrainian National PL – maximum length of petiole (in profile), Academy of Sciences, Kiev, Ukraine (IZK; including PW – maximum width of petiole (from above), Karawajew’s collection); The Natural History Museum, PH – maximum height of petiole (in profile), London, UK (BMNH); Museo Civico di Storia Naturale HTL – maximum length of hind tibia, “G. Doria”, Genoa, Italy (MCSN); Museum and Institute PSL – maximum diameter of propodeal spiracle. of Zoology of Polish Academy of Sciences, Warsaw, Indices: Poland (MIZ) and National Museum of Natural History, Cephalic index: CI = HW / HL,

Tehran, Iran (MMTT). Scape indices: SI1 = SL / HL; SI2 = SL / HW, Measurements and indices : Ocular indices: OI1 = OL / HW; OI2 = OL / GL, HL – maximum length of head in dorsal view, meas- Funicular segment indices: FSI1 = FS1 / FS2; FSI2 ured in a straight line from the most anterior = FS1 / (FS2+FS3), point of clypeus to the mid-point of occipital Maxillary palps indices: MPI1 = MP4 / MP5; MPI2 margin, = MP4 / (MP5+MP6), HW – maximum visible width of head in dorsal view, Propodeal spiracle index: PSL / HW, measured above or below of eyes (depending Alitrunk index: AL / PnW. from species), SL – maximum straight-line length of scape from its apex to the articulation with condylar bulb, st rd TAXONOMY FS1,FS2,FS3 – length of 1 to 3 funicular segments of antenna, rd th MP3, MP4, MP5, MP6 – length of 3 to 6 segments of Cataglyphis stigmatus sp. nov. maxillary palps, OL – maximum diameter of eye, Etymology. From the Latin word “stigma” – spi- GL –length of gena (seen in profile), measured racle, to emphasize very long propodeal spiracles.

3

1 2

Figure 1. Map of the type localities of Cataglyphis stigmatus sp. nov. (black dots) and C. pubescens sp. nov. (black square). (1) Mond Protected Area; (2) Naiband National Park; (3) Siahkooh National Park.

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TWO NEW SPECIES OF THE GENUS CATAGLYPHIS FOERSTER FROM IRAN 71

Material examined. Holotype worker, Iran, Ecology. Distribution of this species is probably Province Bushehr, Mond protected area, 28°03’N, limited to the northern coastal plains of the Persian 51°36’E, 6 m a.s.l., 15 July 2007, arid area, leg. Omid Gulf. This region is characterized by hot long summer Paknia, collection code: MND-2128002 (MMTT); para- and mild winter, with mean annual temperature 27C° types: 1 worker from the nest of holotype; 2 workers and 236 mm precipitation. Phyto-geographically it be- from the same site, but collected on bait traps; 3 work- longs to the subtropical region. Both nest samples of ers, Iran, Province Bushehr, Naiband National Park, C. stigmatus were collected in open arid areas. The 27°18’N, 52°48’E, 19 July 2007, 6 m a.s.l., nest sample, nest entrance had a small mound ca. 5 cm height and arid area, leg. Omid Paknia (IZK, MMTT). ca. 15 cm in diameter. Specimens were active at the hott- Description. Workers (Figs 2–7). Species of medi- est time of day, between 10.00 and 16.00. Workers were um size, body length ca. 5–7 mm. Head with almost par- attracted on baits both by tuna fish and sugar syrup. allel sides (below the eyes) and gradually convex occip- Comparative diagnosis. Based on all main diag- ital margin, occipital corners not marked, head length nostic feature (e.g. nodiform petiole, body sculpture, subequal to its width. Anterior clypeal margin convex, maxillary palpes structure, etc.; see also Agosti 1990, without median notch. Clypeal setae distinctly shorter Radchenko 1997a), C. stigmatus clearly belongs to the than length of clypeus and joined near its anterior bicolor species-group. Almost all species of this group margin. Eyes relatively small, their maximum diam- are bicoloured (with reddish head and alitrunk and eter 1.2–1.5 times less than length of genae, situat- black gaster) or black with the only one previously ed distinctly beyond the midlength of head margins. known exception – C. lunaticus which has entirely Ocelli relatively big, forming equilateral triangle. yellow body. Consequently, C. stigmatus obviously dif- Antennae 12-segmented, scape long, distinctly longer fers by colour from all known species of this group, than head length, first funicular segment distinctly except of C. lunaticus. Despite we did not investigate shorter than the length of second and third segments the type specimens of the latter species (it has been together. 3rd and 4th segments of maxillary palpes long, described based on 2 workers from Turkey), the subequal in length, 5th segment 1.5–1.6 times shorter detailed original description, including morphometric than 3rd or 4th ones, 6th segment is the shortest; 3rd seg- data and excellent drawings, provided by Baroni Urba- ment somewhat flattened, with abundant erect hairs on ni (1969) allow us to compare both species. inner margin, length of the longest hairs equal or only C. stigmatus well distinguishes from C. lunaticus a little longer than maximum diameter of the segment; by the longer scape (SI1 > 1.20 vs < 1.10), by the low- 4th segment with similar pilosity, two apical segments er propodeum with the dorsal surface being distinctly with abundant but shorter hairs. Mandibles with long longer than the posterior one (the length of the dorsal apical tooth, somewhat smaller preapical one and surface of propodeum in C. lunaticus is subequal to three small basal teeth. the length of posterior one), by the less abundant Alitrunk long and slender, mesonotum not raised standing hairs on the alitrunk and petiole, by the some- over pronotal level. Propodeum low, gradually arched, what smaller size, and especially by the much longer its dorsal surface distinctly longer than posterior one. propodeal spiracles. We examined size of propodeal Propodeal spiracles elongate-oval, while not distinctly spiracles in more than fifty Cataglyphis species, slit-like, and very long: their length exceeds (or at least including about twenty ones from the bicolor-group, reaches) half of the propodeal height. Petiole obviously but could not found such big spiracles in any of the nodiform, with rounded node dorsum. investigated specimens. Surface of whole body with dense microreticulation, appears dull, although not strongly matt. Body with sparse whitish standing hairs. Occiput with 5–6 Cataglyphis pubescens sp. nov. quite long erect hairs, frons with 3–4, clypeus – with 2 similar hairs. Alitrunk and petiolar node with a few Etymology. From the Latin word “pubescens ” – sparse short hairs. Head and gaster with very sparse pubescent, that means character of the depressed and short decumbent pilosity, distance between hairs pubescence on the head and alitrunk. longer than hairs’ length; surface of alitrunk (except of Material examined. Holotype worker, Iran, mesonotal dorsum) and coxae with dense silverfish province Yazd, Siahkooh National Park, 32°35’55”N, pubescence. Tibiae with depressed whitish short setae 54°13’57”E, 987 m, 23 May 2008, nest sample, arid area, and additionally with a few yellowish bristles on inner leg. Omid Paknia, collection code: SIA 2459009 margin. Antennae with fine, short depressed pubes- (MMTT); paratypes: 6 workers from the nest of holo- cence, without semi-erect hairs. type; 2 workers from the same locality but collected by Whole body yellow to orange-yellow. pitfall traps (IZK, MMTT). Queens and males are unknown. Description. Workers (Figs 8–13). Species of Measurements and indices see in Tables 1 and 2. small size, body length ca. 4 mm. Head length subequal

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72 A. RADCHENKO and O. PAKNIA

2 mm 2

1 mm 3

0.5 mm 4

0.5 mm 6 0.5 mm 5

0.5 mm 7

Figures 2–7. Photos of details of structure of Cataglyphis stigmatus sp. nov. (holotype, worker). (2) Body, lateral view; (3) head, frontal view; (4) antenna; (5) maxillary palps, lateral view; (6) hind tibia; (7) propodeum and petiole, lateral view (photo H.-P. Katzmann).

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TWO NEW SPECIES OF THE GENUS CATAGLYPHIS FOERSTER FROM IRAN 73

to its width; head slightly narrowed anteriorly, with deum subequal to posterior one, both meet at a round- straight (not convex) sides (below the eyes), rounded ed blunt angle. Propodeal spiracles small, slit-like. occipital corners and very weakly convex occipital Petiole squamiform, with distinct, rather thick scale. margin. Anterior clypeal margin almost straight, with- Surface of head and propodeum with fine but dense out median notch. Clypeal setae subequal to clypeal microreticulation, appears dull, promesonotum and length and joined near its anterior margin. Eyes rela- gaster with very fine superficial microreticulation, tively small, their maximum diameter ca 1.05–1.25 appear shiny. times less than length of genae, situated distinctly Body with sparse whitish standing pilosity, while it beyond the midlength of head margins. Ocelli small, is somewhat more abundant than in the most of species forming equilateral triangle. Antennae 12-segmented, of the cursor- and emmae-group. Occiput with more scape relatively short, subequal or only slightly longer than 10 straight erect hairs, frons and clypeus without than head length; first funicular segment relatively such hairs. Alitrunk and coxae with scattered erect long, only slightly shorter than length of second and hairs of different length, petiolar scale with a few short third segments together, remainder segments distinct- hairs. Head (especially temples and occiput), meso- ly longer than broad. 3rd and 4th segments of maxillary pleura, propodeum and coxae with dense pubescence, palpes rather long, subequal in length, 5th segment formed by long, very abundant silverish appressed short, 1.5–1.9 times shorter than 4th ones, 6th segment hairs. Gaster with very sparse and short decumbent only slightly shorter that the 5th one; 3rd segment not hairs. Scape and funiculus with short, quite thick, flattened, oval in cross-section, with not abundant whitish subdecumbent hairs, tibiae with numerous, erect hairs, length of the longest hairs less than twice rather long subdecumbent to suberect setae, and addi- longer than maximum diameter of the segment; 4th-6th tionally with less abundant yellowish bristles on inner segments with abundant but somewhat shorter pilosi- margin. Whole body black. ty. Mandibles with long apical tooth, somewhat smaller Queens and males are unknown. preapical one and three small basal teeth. Measurements and indices see in Tables 1 and 2. Alitrunk relatively short and robust, mesonotum Ecology. This species was collected in the interior not raised over pronotal level. Dorsal surface of propo- region of the Central Persian desert basin. This area is

Table 1. Measurements (in mm) of Cataglyphis stigmatus sp. nov. and C. pubescens sp. nov.

C. stigmatus (n=7) C. pubescens (n=9) Measurements holotype min max mean ± SD holotype min max mean ± SD HL 1.580 1.066 2.075 1.506 0.3001 1.030 0.988 1.238 1.087 0.0804 HW 1.680 1.002 2.025 1.456 0.3066 0.983 0.962 1.317 1.079 0.1090 SL 2.125 1.404 2.703 1.975 0.3681 1.085 1.050 1.333 1.141 0.0784

FS1 0.435 0.247 0.643 0.420 0.1189 0.234 0.217 0.290 0.242 0.0205

FS2 0.280 0.169 0.368 0.258 0.0578 0.121 0.122 0.170 0.137 0.0150

FS3 0.305 0.205 0.388 0.288 0.0521 0.165 0.155 0.200 0.170 0.0147

MP3 0.525 0.351 0.610 0.481 0.0796 0.269 0.243 0.318 0.286 0.0218

MP4 0.525 0.351 0.614 0.479 0.0790 0.295 0.273 0.322 0.293 0.0165

MP5 0.330 0.234 0.392 0.306 0.0449 0.186 0.143 0.200 0.176 0.0191

MP6 0.205 0.162 0.240 0.195 0.0249 0.146 0.113 0.174 0.141 0.0176 OL 0.480 0.326 0.575 0.432 0.0739 0.355 0.351 0.410 0.377 0.0194 GL 0.658 0.416 0.850 0.606 0.1182 0.401 0.390 0.494 0.431 0.0388 AL 2.781 1.885 3.560 2.577 0.4857 1.396 1.349 1.720 1.494 0.1056 PnW 1.162 0.710 1.400 1.045 0.2045 0.712 0.675 0.905 0.760 0.0731 PL 0.450 0.350 0.720 0.492 0.1215 0.340 0.320 0.391 0.350 0.0232 PW 0.317 0.247 0.460 0.328 0.0666 0.304 0.299 0.414 0.340 0.0444 PH 0.340 0.210 0.507 0.357 0.0973 0.345 0.278 0.395 0.356 0.0312 HTL 3.240 1.950 3.950 2.875 0.5728 1.381 1.339 1.610 1.447 0.0935 PSL 0.325 0.215 0.375 0.291 0.0491 0.091 0.078 0.125 0.100 0.0143

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74 A. RADCHENKO and O. PAKNIA

8 1 mm

10 9 0.5 mm

0.5 mm

0.2 mm 12 11 0.2 mm

0.2 mm 13

Figures 8–13. Photos of details of structure of Cataglyphis pubescens sp. nov. (holotype, worker). (8) Body, lateral view; (9) head, frontal view; (10) antenna; (11) hind tibia; (12) clypeus and maxillary palps, lateral view; (13) propodeum and petiole, lateral view (photo H.-P. Katzmann).

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TWO NEW SPECIES OF THE GENUS CATAGLYPHIS FOERSTER FROM IRAN 75

Table 2. Morphometric indices of Cataglyphis stigmatus sp. nov. and C. pubescens sp. nov.

C. stigmatus (n=7) C. pubescens (n=9) Indices holotype min max mean ± SD holotype min max mean ± SD CI 1.063 0.938 1.064 0.991 0.0520 0.954 0.977 1.068 1.005 0.0380

SI1 1.345 1.243 1.352 1.314 0.0309 1.053 0.984 1.077 1.050 0.0326

SI2 1.265 1.265 1.422 1.363 0.0502 1.104 1.005 1.149 1.060 0.0466

OI1 0.286 0.283 0.325 0.299 0.0148 0.361 0.308 0.374 0.350 0.0179

OI2 0.729 0.664 0.839 0.719 0.0544 0.885 0.811 0.949 0.876 0.0480

FSI1 1.554 1.409 1.843 1.616 0.1453 1.934 1.622 1.954 1.777 0.1083

FSI2 0.744 0.660 0.875 0.761 0.0817 0.818 0.712 0.888 0.791 0.0541

MPI1 1.591 1.500 1.680 1.561 0.0519 1.586 1.519 1.936 1.679 0.1557

MPI2 0.981 0.886 1.024 0.954 0.0361 0.889 0.858 1.027 0.931 0.0637 PSI 0.193 0.185 0.220 0.202 0.0116 0.093 0.079 0.111 0.092 0.0099 AI 2.393 2.345 2.654 2.477 0.1018 1.961 1.754 2.183 1.973 0.1343

characterized by hot summer and cold winter with definitively resolved when males will be found. Despite mean annual temperature 19°C, and by the very low this little taxonomic vagueness, C. pubescens clearly annual precipitation – 67 mm only. This territory differs from any knows species of the groups men- belongs to the Irano-Turanian phyto-geographical tioned above by the much more developed, dense region. Most specimens were collected by hand from appressed pubescence on the head and alitrunk. a nest. Nest was built in an open area, having a small entrance without surrounding structures. Comparative diagnosis. C. pubescens shares several features of the emeryi-, cursor- and emmae ACKNOWLEDGEMENTS species-groups of Cataglyphis. Thus, setae on the anterior clypeal margin are very long, subequal to or We are sincerely grateful to curators of the ant col- even somewhat longer than the length of clypeus, simi- lections of all Museums and Institutions mentioned larly to C. emeryi (Karawajew), but unlike the latter above for the assistance during our investigations. We species these setae join close to the anterior clypeal thank Martin Pfeiffer and Elisabeth Kalko for their margin, as in the species of cursor-group (Radchenko kind support of O. Paknia study in Germany, and to 1997a, 1998). The first funicular segment is quite long, Hans-Peter Katzmann (Ulm University, Germany) for about twice longer than the second one and only slight- making photos of the holotype specimens. We are also ly shorter than the second and third segments togeth- grateful to the staff of Department of Environment of er: this is one of the diagnostic features of workers of Iran for the assistance and help that allowed that mate- the emmae-group (according Agosti 1990). On the oth- rial to be collected from natural reserves in Iran. We er hand, worker caste of C. pubescens is not dimorphic are also grateful to X. Espadaler (Universitat Autòno- (the latter is characteristic for the emmae-group ma de Barcelona, Spain) and B. Markó (Cluj Universi- species); additionally, they have distinctly thicker peti- ty, Romania) for the available comments to the manu- olar scale than C. emeryi. In general, workers of the script of this paper. species of all three groups mentioned above are super- ficially quite similar to one another, particularly their whole body is blackish-brown to black, they have peti- REFERENCES ole with distinct scale (i.e. it is not cuneiform or nodi- form), but their males well differ by the structure of genitalia (Agosti 1990; Radchenko 1997a). Moreover, Agosti, D. 1990. Review and reclassification of Cataglyphis (Hymenoptera, Formicidae). Journal of Natural History, workers of C. emmae and C. emeryi move slowly, 24: 1457–1505. rather like Proformica Ruzsky species (C. emmae Agosti, D. 1994. A new inquiline ant (Hymenoptera, Formici- has been originally described as a member of dae) in Cataglyphis and its phylogenetic relationship. Proformica) than Cataglyphis, while C. pubescens Journal of Natural History, 28: 913–919. move very fast, like most of the Cataglyphis species. Baroni Urbani, C. 1969. Una nuova Cataglyphis dei monti The proper taxonomic position of this species can be dell’Anatolia. Fragmenta entomologica, 6(3): 213–222.

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Bolton, B. 1995. A new general catalogue of the ants of the Cataglyphis Foerster (Hymenoptera, Formicidae) from World. Cambridge-London, Harvard University Press, Asia. Entomologicheskoe obozrenie, 76 (2): 424–442 (in 504 pp. Russia; English translation: Entomological Review (Wash- Bolton, B., Alpert, G., Ward, P. S. and P. Naskrecki. 2007. ington), 1997, 77: 684–698). Bolton’s Catalogue of Ants of the World. Harvard Univer- Radchenko, A. G. 1997b. Cataglyphis zakharovi sp. nov. – sity Press, Cambridge-London, CD-Rom. the second socially parasitic species in the genus Cata- Paknia, O., Radchenko, A., Alipanah, H. and M. Pfeiffer. 2008. glyphis Foerster (Hymenoptera, Formicidae). Annales A preliminary checklist of the ants (Hymenoptera: Formi- Zoologici, 46: 207–210. cidae) of Iran. Myrmecological News, 11: 151–159. Radchenko, A. G. 1998. A Key to ants of the genus Cata- Paknia, O. Radchenko, A. and M. Pfeiffer. 2009. New records glyphis Foerster (Hymenoptera, Formicidae) from Asia. of ants (Hymenoptera, Formicidae) from Iran. Asian Myr- Entomologicheskoe obozrenie, 77 (2): 502–508 (in Russia; mecology, 3: 29–38 English translation: Entomological Review (Washington), Radchenko, A. G. 1997a. A review of ants of the genus 1998, 78: 475–480).

Received: December 1, 2009 Accepted: March 9, 2010

131

Synthesis

This dissertation is a try to extend our knowledge about community ecology of ants in arid and semi-arid regions of the Palearctic realm. Conducting extensive field works in 2007 and 2008, I prepared one the most complete ant collections of the Southwest Asia. However, as our sampling sites were limited to the north and the centre of Iran, vast regions in the west and the east of the country have remained unstudied. Faunistically and taxonomically speaking, this collection has contributed largely to our understanding from the fauna of the region (Paknia et al., 2008; Seifert & Schultz, 2009; Paknia et al., 2010; Radchenko & Paknia, 2010). Nevertheless, many unidentified or unrecorded material has remained for upcoming works. Moreover, many specimens have been preserved in a condition that can be used for population genetic and phylogeography studies in future. Particularly, phylogeographical studies in the Southwest Asia could reveal interesting results to biogeographical science, as this region is the meeting point of three realms, Palearctic, Afrotropic and Indomalaya. Our preliminary findings suggest that the Iranian ant fauna has a typically Palearctic character, but a few Afrotropical and Indomalayan elements are present in the fauna of Iran. A concrete conclusion can be made when sufficient material will have been sampled and phylogeographical studies will have been conducted in this region. The provided collection will also contribute to the learning and researching of young Iranian students who are interested in ant studies, as www.antbase.net has started publishing Automontage© photographs of Iranian ant species, thus making the fauna easier accessible for non-experts. Beyond the taxonomical and faunistical contributions, the aim of this thesis was to investigate diversity and species compositions of ants in drylands of Iran and the role of spatial and environmental factors in the occurrences of these patterns. In spite of their low productivity, drylands represents highly diverse faunas, especially for specific taxa such as ants (Shachak et al., 2005). At the regional scale, Australian arid and semi-arid regions consist of 4,500 ant species which is much higher than the

Chapter 9 - Synthesis world average species richness. North American and South African arid regions comprise 300 and 400 species, respectively (Andersen, 2007). There is no such a data for the Southwest Asian arid regions and for Iran. Collinwood and Agosti (1996) have reported 250 species only from Arabian Peninsula. In the present work I reported more than 90 species. As result, it could be predicted that ant diversity in the Southwest Asia is similar or higher than in North American and South African arid regions. At the local scale, Australia, again, shows an extraordinary pattern by occurrences of more than one hundred species per hectare, comparing to the North America and the South Africa with 30-40 species in one hectare. A rough comparison among these regions and species richness of ants in Iran at local scale suggests that local diversity of ants in Iran and possibly in the Southwest Asia is lower than the North America and the South Africa. Local species richness of ants in arid and semi- arid regions of Iran did not exceed ~ 18 species. This result, however, can be due to applying a single sampling method in the present study (pitfall trap). Pitfall trap method is appropriate for short-term investigations of open habitats and accurately represents epigaeic ant species diversity but the sampling coverage might be biased by failing to capture cryptic or litter species, especially in steppe regions, as pitfall trapping is not a suitable method for sampling cryptic and litter ants (Agosti et al., 2000). It is likely that due to this bias, I found no significant difference at local scale among steppe and arid regions. More inventories are needed to disclose the real number of ant species at both regional and local scales in Iran and the Southwest Asia. Compare to the alpha diversity, little is known about the beta diversity of ants in arid regions. Pfeiffer et al. (2003) found low local beta diversity in Mongolia. However, they found distinctive ant composition between steppe and desert. In contrary, harvester ants in Australia covered a smaller size range and showed high beta diversity (Morton & Davidson, 1988). In Iranian arid regions, a large number of the total ant diversity was generated by beta diversity. Results of the multiscale approach analysis along latitudinal gradient revealed that the highest proportion of beta diversity in ant assemblages occurred at the broad scales, among biomes and ecoregions. Significant beta diversity even was observed at finest scale, suggesting that most ant species are highly localized in small patches of habitats. Further analyses revealed that contemporary climate factors including annual rainfall, summer rainfall and temperature range are largely responsible for ant beta diversity in arid and semi-arid regions of Iran at broad scales. Specifically, rainfall which is the main

133 Chapter 9 - Synthesis controlling factor for biodiversity in arid ecosystems influenced ant species compositions by its both spatial and temporal dimensions. This finding is consistent with comparable studies on ants in Mongolia (Pfeiffer et al., 2003) and in the North America (Davidson, 1977; Kaspari & Valone, 2002). Moreover, results in chapter 4 of this study, showed a large impact of temperature range on species composition of ants which corroborates with Pfeiffer et al. (2003) findings in Mongolia, suggesting a remarkable impact of low and instability of temperature on ants, as typical thermophiles, in cold and semi-cold drylands such as Mongolia and Iran. The present study, further, illustrated the negative impact of temperature range on species richness of ants in Central Persian desert. Similar influence was only observed by Dunn et al. (2009) at the global scale. In sum, results in chapters 4 and 5 indicate the role of climate variables, especially rainfall (indirectly as productivity) and temperature range in the occurrences of ant species and their identity across geographical variations. The results of this study shed the first light on our understanding of the conservation status of ants in arid and semi-arid regions of Iran and the Southwest Asia. Overall, the ant assemblages in drylands of Iran are under two obvious threats. Climate change, as the first threat, has received considerable international attention among ecologists and conservationists since the eighty’s (e.g. Wigley et al., 1980; Parmesan, 1996; Thomas et al., 2004), but mainly in other ecosystems such as the tropics (e.g. Pounds et al., 1999; Still et al., 1999) and the Arctic region (e.g. Burek et al., 2008). Very recent studies, however, have provided crucial support for the impact of climate change on dry regions and the role of these regions on climate change. These findings show that dryland ecosystems are highly vulnerable to global climate change, especially to climate warming and changes in rainfall regimes (Wu et al., 2010; Xia et al., 2010). On the other side, they play crucial role in climate change mitigation by high rates of carbon uptake (Rotenberg & Yakir, 2010). A few studies conducted on the climatic trends in the Southwest Asia have demonstrated that aridity has increased during last 30 years (e.g. Kafle & Bruins, 2009). In addition, it has been shown that temperature will increase and rainfall will decrease in vast parts of the region (e.g. Evans, 2009). Such a scenario suggests that ant species richness and species composition will be influenced in uncertain degrees. There would most likely be shifts in distributions for a group of ant species (Thomas, 2010) and local and regional extinctions for those species which could not adapt to new conditions or could not

134 Chapter 9 - Synthesis find suitable refuges. Semi-arid regions are more in danger due to shifts of land use by humans from drier regions to these regions (Evans, 2009). The second threat can be categorized as a specific danger for insects regarding to their highly localized distributions, similar to such pattern that was observed for ants in Iran. In this context, it has been suggested that many insect species might be prone to extinction because they are narrow habitat specialists (Dunn, 2005), and vulnerable even to localized habitat changes such as local disturbance. Broadly speaking, rare species are susceptible to local or regional extinctions. Here, rarity is translated to its three dimensions: low abundance, small distribution and ecological specialisation. The work presented here has explicitly revealed that many ant species are rare as they fit well in one of these categories (small distribution). Future work on ants in Iran must include measuring the abundance and degree of ecological specialisation of the species. Overall, the findings of this study suggest the presence of a rich fauna of ants for arid and semi-arid regions of Iran. Further studies are warranted to explore the complete fauna of these regions. At large scales, species richness and species composition are influenced largely by abiotic factors such as rainfall. But as it has been shown, many ant species have been restricted to local habitats, suggesting there are other factors that act at local scale and determine the occurrences of ant species and their distribution ranges within local habitats. Although by reviewing literature a list of the likely driving factors at local scale can be suggested, further studies are needed to define the correct factors which affect ant diversity and assemblages at local scale in the semi-cold arid regions of Iran.

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REFERENCES: Agosti, D., Majer, J.D., Alonso, L.E. & Schultz, T.R. (2000) Ants: Standard methods for measuring and monitoring biodiversity. Smithsonian Institution Press, London. Andersen, A.N. (2007) Ant diversity in arid Australia: a systematic overview. Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80. (ed. by R.R. Snelling, B.L. Fisher and P.S. Ward), pp. 91-51. Burek, K.A., Gulland, F.M.D. & O'hara, T.M. (2008) Effects of climate change on arctic marine mammal health. Ecological Applications, 18, S126-S134 Collingwood, C.A. & Agosti, D. (1996) Formicidae (Insecta: Hymenoptera) of Saudi Arabia (Part 2). Fauna Saudi Arabia, 15, 300-385 Davidson, D.W. (1977) Species diversity and community organization in desert seed- eating ants. Ecology, 58, 711-724 Dunn, R.R. (2005) Modern insect extinctions, the neglected majority. Conservation Biology, 19, 1030-1036 Dunn, R.R., Agosti, D., Andersen, A.N., Arnan, X., Bruhl, C.A., Cerd, X., Ellison, A.M., Fisher, B.L., Fitzpatrick, M.C., Gibb, H., Gotelli, N.J., Gove, A.D., Guenard, B., Janda, M., Kaspari, M., Laurent, E.J., Lessard, J.-P., Longino, J.T., Majer, J.D., Menke, S.B., P., M.T., Parr, C.L., Philpott, S.M., Pfeiffer, M., Retana, J., Suarez, A.V., Vasconcelos, H.L., Weiser, M.D. & Sanders, N.J. (2009) Climatic drivers of hemispheric asymmetry in global patterns of ant species richness. In: Ecology Letters, pp. 324-333 Evans, J. (2009) 21st century climate change in the Middle East. Climatic Change, 92, 417-432 Kafle, H. & Bruins, H. (2009) Climatic trends in Israel 1970–2002: warmer and increasing aridity inland. Climatic Change, 96, 63-77 Kaspari, M. & Valone, T.J. (2002) On ectotherm abundance in a seasonal environment-studies of desert ant assemblage. Ecology, 83, 2991-2996 Morton, S.R. & Davidson, D.W. (1988) Comparative structure of harvester ant communities in arid Australia and North America. Ecological Monographs, 58, 19-38 Paknia, O., Radchenko, A., Alipanah, H. & Pfeiffer, M. (2008) A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News, 11, 151-159 Paknia, O., Radchenko, A. & Pfeiffer, M. (2010) New records of ants (Hymenoptera: Formicidae) from Iran. Asian Myrmecology, 3, 29-38 Parmesan, C. (1996) Climate and species' range. Nature, 382, 765-766 Pfeiffer, M., Chimedregzen, L. & Ulykpan, K. (2003) Community organization and species richness of ants (Hymenoptera: Formicidae) in Mongolia along an ecological gradient from steppe to Gobi desert. Journal of Biogeography, 30, 1921-1935 Pounds, J.A., Fogden, M.P.L. & Campbell, J.H. (1999) Biological response to climate change on a tropical mountain. Nature, 398, 611-615 Radchenko, A. & Paknia, O. (2010) Two new species of the genus Cataglyphis Foerster, 1850 (Hymenoptera: Formicidae) from Iran. Annales Zoologici (Warsaw), 60, 69-76 Rotenberg, E. & Yakir, D. (2010) Contribution of semi-arid forests to the climate system. Science, 327, 451-454

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Seifert, B. & Schultz, R. (2009) A taxonomic revision of the Formica rufibarbis FABRICIUS, 1793 group (Hymeno-ptera: Formicidae). Myrmecological News, 12, 255-272 Shachak, M., Gosz, J.R., Steward, T.A.P. & Perevolotsky, A. (2005) Biodiversity in drylands toward a unified framework. Oxford University Press, New York. Still, C.J., Foster, P.N. & Schneider, S.H. (1999) Simulating the effects of climate change on tropical montane cloud forests. Nature, 398, 608-610 Thomas, C.D. (2010) Climate, climate change and range boundaries. Diversity and Distributions, 16, 488-495 Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M., Beaumont, L.J., Collingham, Y.C., Erasmus, B.F.N., De Siqueira, M.F., Grainger, A., Hannah, L., Hughes, L., Huntley, B., Van Jaarsveld, A.S., Midgley, G.F., Miles, L., Ortega-Huerta, M.A., Townsend Peterson, A., Phillips, O.L. & Williams, S.E. (2004) Extinction risk from climate change. Nature, 427, 145-148 Wigley, T.M.L., Jones, P.D. & Kelly, P.M. (1980) Scenario for a warm, high-CO2 world. Nature, 283, 17-21 Wu, S., Yin, Y., Zhao, D., Huang, M., Shao, X. & Dai, E. (2010) Impact of future climate change on terrestrial ecosystems in China. International journal of climatology, 30, 866-873 Xia, Y., Moore, D.I., Collins, S.L. & Muldavin, E.H. (2010) Aboveground production and species richness of annuals in Chihuahuan Desert grassland and shrubland plant communities. Journal of Arid Environments, 74, 378-385

137

Summary

Arid, semi-arid and sub-humid regions cover more than 47% of the Earth’s land surface. In spite of their low primary productivity, they represent a high diversity of species, especially in several animals and plants taxa such as ants, beetles and annual plants. Drylands are nowadays experiencing a fast rate of changes, particularly due to climate change and disturbance by humans. Understanding these changes in drylands requires the increase of general knowledge about the current status of biodiversity, its distribution along environmental gradients and those factors which have the highest impacts on it. Although, there is a large body of literature on diversity of animals and plants in the Australian and the North American drylands, such knowledge is rare in other arid regions, such as the Palearctic’s. Ants, as one of most successful taxa have colonized all arid biomes and are a dominant component of invertebrate assemblages in deserts and grasslands around the world, in which they are also functionally important elements with a variety of ecological roles. In many arid and semi-arid regions of the Palearctic realm ants have remained poorly known in spite of their fundamental ecological function in these areas.

The general objective of this dissertation was to broaden the understanding of the impact of environmental and spatial factors on species richness, diversity and assemblage organization of ants in arid and semi-arid regions in Iran. To do so, two field works were performed in 2007 and 2008 in Iran. In the first field work, I sampled ants along a latitudinal gradient (ca. 1200 km) across four main ecoregions. In the second field work, ants were collected from Central Persian desert basins.

My thesis included four stages: The first step included faunistical and taxonomical studies on the poorly known ant fauna of Iran (chapters 6, 7 & 8),

Summary which laid the basis for the further studies on the ecology of ant communities. In the second step, I described and analyzed the diversity pattern of ants along a latitudinal gradient and in two main biomes of Iranian arid regions: steppe and desert (chapter 2). In the third step, ant diversity and composition was analyzed by a hierarchical partitioning approach, from a conservational perspective. Assuming that the WWF’s ecoregional map accurately reflects the biogeographical units of the region, I explored whether the highest beta diversity of ants occurs at the ecoregional scale or at lower scales (chapter 3). In the fourth step, I analyzed the impact of environmental and spatial factors on ant species richness and species composition across the latitudinal gradient and in the Central Persian desert ecoregion (chapters 4 and 5).

When exploring the diversity pattern of ants across a latitudinal gradient, my results revealed that alpha species richness increased along the gradient, but beta diversity did not follow any specific trend. Multiscale analysis of the data showed a constant increase in species richness and diversity of ants across fine- to-broad scales. Evenness, however, decreased across this hierarchical trend. The steppe biome had higher numbers of species than the desert biome, but only at the highest scale. The two biomes represented two distinctive ant species compositions. Further analyses of hierarchical partitioning showed that the WWF’s ecoregions correctly represent the biogeographic components of ants in Iran and are an appropriate scale for conserving of ant diversity, as the highest species turnover was observed in this level. Four distinctive ant species compositions were found across four ecoregions. In one case, the boundaries of these four compositions, however, showed a deviation from the depicted boundaries by the WWF’s map, suggesting that the ecoregional boundaries should be considered as a subject for further tests of their robustness.

Analyses of the environmental and spatial impact on ant species composition and species richness across the latitudinal gradient and within the Central Persian desert ecoregion showed that contemporary climate factors such as rainfall and temperature range and productivity have large influence on ant diversity and assemblage organizations. My faunistical and taxonomical studies have resulted in reporting one subfamily, eight genera and 39 species new to Iran and

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Summary describing two species new to science. Additionally, I provided the first species- list for Iranian ants.

In conclusion, arid and semi-arid regions of Iran consist of a relatively rich fauna of ants. More than 90 ant species were collected within 14 natural reserves during two sample years. Although, local (alpha) species richness of ants in the study regions was relatively low and did not exceed ~ 18 species, my analysis showed that there is high significant beta diversity across the all spatial scales, suggesting that many ant species have restricted distributions within the arid and semi-arid regions. This pattern suggests that arid regions are not simply vast homogenous areas; in contrast these regions exhibit high environmental heterogeneity which limit the distribution of many species to a small habitat and produce patchy diversity patterns. The highly localized distributions of many ant species make them vulnerable to extinctions by local or regional disturbances. Furthermore, as their diversity and assemblage organization is highly related to climate factors, the prospective predicted climate changes at the regional scale such as increase of aridity and temperature and higher variability in rainfall will most likely have large impact on ants. They would most likely result in shifts in distributions for a group of ant species and local and regional extinctions for those species which could not adapt to the new conditions or could not find suitable refuges.

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Zusammenfassung

Aride, semi-aride und humide Regionen bedecken mehr als 47% der Erdoberfläche. Trotz ihrer niedrigen Produktivität zeigen sie eine hohe Artenvielfalt, vor allem in mehreren Tier – und Pflanzentaxa, wie Ameisen, Käfern und annuelle Pflanzen. Trockengebiete unterliegen derzeit raschen Veränderungen, besonders durch den Klimawandel und anthropogene Beeinträchtigung. Zum Verständnis dieser Veränderungen, ist es notwendig mehr über den gegenwärtigen Status der Biodiversität und ihre Verteilung entlang von Umweltgradienten wissen, sowie über die Faktoren, die diese Verteilung bestimmen.

Obwohl zahlreiche Veröffentlichungen zur Diversität von Tier-und Pflanzenarten in den Australischen und Nordamerikanischen Trockengebieten erschienen sind, ist unser Wissen über die anderen ariden und semi-ariden Regionen der Erde, zum Beispiel die der Paläarktis, gering. Die Ameisen haben, als eines der erfolgreichsten Taxa, alle ariden Biome besiedelt und sind eine dominante Komponente in den Invertebraten-gemeinschaften der Wüsten und Grasländer der Erde, dort stellen sie funktionell bedeutsame Elemente dar, die eine Vielzahl ökologischer Rollen besetzen. Trotz ihrer wichtigen ökologischen Funktion ist in vielen ariden und semi-ariden Gebieten der Paläarktis wenig über die dortigen Ameisen bekannt.

Ziel dieser Dissertation war es den Einfluss von Umwelt- und räumlichen Faktoren auf den Artreichtum, die Diversität und die Organisation von Ameisengemeinschaften in ariden und semi-ariden Gebieten des Iran zu ergründen. Zu diesem Zweck wurden 2007 und 2008 im Iran Feldarbeiten durchgeführt. In der ersten Periode sammelte ich Ameisen entlang eines Breitengradgradienten (ca. 1200 km) in vier bedeutenden Ökoregionen. Während der zweiten Periode wurden Ameisen im Zentralen Persischen Wüstenbecken gesammelt.

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Meine Arbeit umfasste vier Phasen: Die erste Phase beinhaltete die Erstellung von faunistischen und taxonomischen Studien über die kaum bekannte Ameisenfauna des Iran als Grundlage für die weitere ökologische Forschung an diesen Gemeinschaften (Kapitel 6, 7 und 8). Im zweiten Schritt beschrieb und analysierte ich die Diversitätsmuster von Ameisen entlang eines Breitengradgradienten und in den zwei wichtigsten Biomen der Iranischen Trockengebiete: Steppe und Wüste (Kapitel 2). Im dritten Schritt, wurde die Diversität und Gemeinschaftstruktur mit Hilfe eines mathematischen Ansatzes zur hierarchischen Partitionierung untersucht, dabei spielten Naturschutzaspekte eine Rolle. Unter der Annahme, dass die WWF Karte der Ökoregionen die biogeographischen Einheiten der Region genau widerspiegelt, untersuchte ich, ob die höchste Betadiversität auf der Ebene der Ökoregion, oder auf tieferen Skalen auftritt (Kapitel 3). Im vierten Schritt untersuchte ich den Einfluss von Umwelt- und räumlichen Faktoren auf den Artenreichtum und die Artzusammensetzung der Ameisen in der zentralen Persischen Wüste und entlang des Breitengradgradienten (Kapitel 4 und 5).

Meine Ergebnisse zur Diversität der Ameisen entlang des Breitengradgradienten zeigen, dass der Alpha-Artreichtum entlang des Gradienten zunahm, aber die Betadiversität keinem spezifischen Trend folgte. Multiscale-Analysen der Daten zeigten eine konstante Zunahme von Artreichtum und Diversität der Ameisen mit zunehmender räumlicher Skala. Gegenläufig nahm die Evenness auf größerer Skala ab. Die Steppenbiome hatten eine größere Anzahl von Arten als die Wüstenbiome, aber nur auf der höchsten Ebene der Analyse. Die beiden Biome repräsentierten zwei unterschiedliche Artengemeinschaften. Die genauere Analyse der hierarchischen Partitionierung zeigte, dass die WWF Ökoregionen die biogeographischen Komponenten des Iran gut repräsentieren und eine geeignete Skala darstellen, um Maßnahmen zum Schutz der Ameisendiversität zu treffen, denn der größte Artenwechsel ergab sich auf dieser Ebene.

Für jede der vier Ökoregionen wurde eine distinkte Artengemeinschaft gefunden. In einem Fall zeigten die Grenzen dieser Artengemeinschaften Abweichungen von den Grenzen der WWF Karte, daher sollten die Grenzen der Ökoregionen in weiteren Untersuchungen auf ihre Robustheit geprüft werden.

Die Analysen zum Einfluss der Umwelt und der räumlichen Konstellation auf Artenzusammensetzung und -reichtum entlang des Breitengradgradienten und

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Summary innerhalb der zentralen persischen Wüstenregion zeigten, dass Klimafaktoren, wie z.B. Produktivität oder ihre potentiellen Surrogate (Niederschlag), sowie die Variabilität der Umgebungstemperatur, großen Einfluss auf die Diversität der Ameisen und die Organisation ihrer Gemeinschaften hat. Eine Unterfamilie, acht Gattungen und 39 Arten wurden im Laufe meiner faunistischen und taxonomischen Studien als neu für den Iran aufgelistet, zudem wurden zwei neue Arten wissenschaftlich beschrieben. Außerdem stellte ich die erste Artenliste für die Ameisen des Iran zusammen.

Zusammenfassend zeigen die ariden und semi-ariden Gebiete des Iran eine relativ reiche Ameisenfauna. Mehr als 90 Arten wurden auf 14 Probeflächen im Laufe von zweie Jahren gesammelt. Obwohl die lokale Alphadiversität von Ameisen in den Untersuchungsgebieten relativ gering war und 18 Arten nie überstieg, zeigte meine Analyse eine signifikante, hohe Betadiversität über alle räumlichen Skalen, was darauf hinweist, dass viele Ameisenarten nur eng begrenzte Gebiete der weitläufigen Trockenregionen besiedeln. Dieses Muster suggeriert, dass aride Gebiete nicht einfach riesige, homogene Areale darstellen, sondern eine hohe Umweltheterogenität aufweisen, die die Verbreitung bestimmter Arten limitiert und zu kleinräumigen Diversitätsmustern führt. Die hochgradig kleinräumige Verteilung vieler Ameisenarten macht sie verwundbar für Extinktionen durch lokale und regionale Störungen. Zudem wird der prognostizierte Klimawandel, der sich auf regionaler Ebene zum Beispiel als Erhöhung der Aridität und Temperatur, sowie in einer größeren Variabilität der Niederschläge zeigt, wahrscheinlich starke Auswirkungen auf die Ameisen haben, deren Diversität und Gemeinschaftsorganisation stark von klimatischen Faktoren geprägt ist. Das wird voraussichtlich dazu führen, dass sich die Verbreitungsgebiete einiger Ameisenarten verändern und Arten, die sich den neuen Bedingungen nicht anpassen können, lokal oder regional aussterben werden.

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ﺧﻼ

ﻨﺎﻖ ﺸﮏ، ﻪ ﺸﮏ و ﻪ ﻮب از ﻞ و ﺖ ﺻﺪ از ﺢ زﻦ را ﯽ ﭘﻮﺷﺎﻨﺪ. ﺑﺎ وﻮد ﻮﯿﺪ اوﻪ ﮐﻢ، اﻦ ﻨﺎﻖ ﻮع ﻮ ای ﺑﺎﻻﯽ را

وه ﻮص ﯽ از ﺟﺎﺪاران و ﯿﺎن ﻣﺎﻨﺪ ﻮر ، ﻮﺳﮏ و ﯿﺎن ﯾﮏ ﺳﺎ دارا ﯽ ﺑﺎﻨﺪ. ﻨﺎﻖ ﺸﮏ ھﻢ اﻮن ﺮات ﺮ وه

ﻋ ﻮص دﻮﯽ ی آب و ﻮاﯽ و ﻂ زﺖ دﺖ اﺴﺎن را ﯽ ﻨﺪ. ک اﻦ ﺮات ﻨﺎﻖ ﺸﮏ ﯿﺎز ااﺶ داﺶ ﻮﯽ

ﺑﺎره ﺮاﻂ ﻮع ﻮ ای، ﺮﭘاﻨﺪﯽ ﻮ ﻮل ی ﯽ و ﻨﺎﺖ ﻮرﯽ دارد ﻦ ﺮ را ﻮع ﻮ ای دارﺪ. ا ﻨﺎﻊ اواﯽ

ﻮص ﻮع ﻮ ای ﺟﺎﻮران و ﯿﺎن ﻨﺎﻖ ﺸﮏ اﺮاﯿﺎ و آﮑﺎی ﻤﺎﯽ وﻮد دارد، ﻦ اﻃﻼﻋﺎﯽ دﯾ ﻨﺎﻖ ﻣﺎﻨﺪ ﻮاﯽ ﺸﮏ و ﻪ ﺸﮏ

ﺴ ﭘﺎﺌﺎر ﯿﮏ ﯾﺎب ﯽ ﺑﺎﺪ. ﻮر ﻮان از ﻮﻖ ﻦ دﻪ از ﺟﺎﺪاران ﻤﺎﯽ زﺖ ﻮم ی ﺸﮏ دﯿﺎ ﺳﺎﻦ ﺪه اﺪ و ﺖ ﻏﺎ اﻤﺎع

ﭼ ھ ﻋ ﺮات را ﻨﺎﻖ ﯿﺎﺑﺎﯽ و ﺰار ﺮار دﯿﺎ ﻞ ﯽ دﻨﺪ اﻦ ﻨﺎﻖ ﻦ ااء ﮫﻢ ﻤﻠد اﻮﻮژ زA ﮕﺎه ﯽ ﺑﺎﻨﺪ و ﺶ ی ﮫﻢ

اﻮﻮژ را ﮫﺪه دارﺪ. ﻋﯽ رﻢ ﺶ اﻮﻮژ ﯿﺎر ﮫﻢ ﻮاﯽ ﺸﮏ، ﻮر ﯿﺎری از ﻨﺎﻖ ﯿﺎﺑﺎﯽ و ﻪ ﯿﺎﺑﺎﯽ ﭘﺎﺌﺎرﯿﮏ ﻨﺎﻪ ﻣﺎﺪه اﺪ.

ﻮﻮع ﮐﯽ اﻦ رﺳﺎ ﺮش داﺶ ﻮﻮد ﻮص ﺮ ﻮاﻞ ﯽ و ﻮری اﯿﺎﯽ روی ﻮع ﻮ ای و ﻨﺎی ﻮ ای و ﺳﺎﺘﺎر اﻤﺎﯽ ﻮر

ﻋ ﻨﺎﻖ ﺸﮏ و ﻪ ﺸﮏ اان ﻮده اﺖ. ای ﻖ اﻦ ﻮﻮع، دو ﻋﻤﯿﺎت اﯽ ﺳﺎل ی ١٣٨۶ و ١٣٨٧ ﯽ اان ﻤﻞ آﺪ.

اوﻦ ﻋﻤﯿﺎت اﯽ ﻮر ﻮل اﯿﺎﯽ ﺑﺎ ﺬر از ﮫﺎر ﻪ ز ﯽ (١٢٠٠ ﯿﻮﺮ) ﻊ آوری دﯾﺪﺪ. دوﻦ ﻋﻤﯿﺎت اﯽ، ﻮر از زی -

ﻪ ﯿﺎﺑﺎن ی ی اان ﻊ آوری ﺪﺪ.

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رﺳﺎ ﺣﺎﺮ ﺷﺎﻞ ﮫﺎر ﺣﻪ ﻮده اﺖ. ﺣﻪ ﺖ ﺷﺎﻞ ﻄﺎﻌﺎت ﻮﯿﮏ و ﻮﻮﯿﮏ روی ﻮن اان ﻮده اﺖ ﻮن ﻮرت ﺰ

ﻮرد ﻄﺎﻪ ار ﻪ اﺖ ( ﻞ ی ۶، ٧ و ٨) و ﻮان ﭘﺎ ای ای ﻄﺎﻌﺎت اﻮﻮژ ای ﺳﺎﺘﺎر اﻤﺎﯽ ﻮر ﻞ ی ﻌﺪ ﯽ ﺑﺎﺪ.

ﺤ ﺣﻪ دوم ﺳﺎﺘﺎر ﺮﭘاﻨﺪﯽ ﻮر ﻮل اﯿﺎﯽ ﺷﺎﻞ دو زﻮم ﭘﻦ دﺖ و ﯿﺎﺑﺎن ﺮ ﺢ و و ﻞ ﺪ ( ﻞ ٢). ﻮﻦ ﺣﻪ، ﻮع

ﺤ WWF ﻮ ای ﻮر ﺑﺎ روﯾد ء ﻨﺪی ﻪ ای از دﯾﺪﮔﺎه ﻔﺎﯽ ﻮرد و ﻞ ار . اﻦ ﻄﺎﻪ، ﺑﺎ ض اﻪ ﻪ ﻮاﯽ ز ﯽ

اان ﺢ ﻮده و ﯽ واﺪی ز ﯽ -اﯿﺎی ﻪ را ارا ﯽ ﻨﺪ، رﯽ ﺪ آﯾﺎ ﻦ ﺰان ﺮ ﻮ ﯿﺎس ﻮاﯽ ز ﯽ اﺠﺎم ﯽ ﺮد و

ﯾﺎ ﯿﺎﯽ ﻮﭼﮏ ( ﻞ ٣). ﮫﺎرﻦ ﺣﻪ ﺰان ﺮ ﻮاﻞ ﯽ و ﻮری اﯿﺎﯽ روی ﻮع ﻮ ای و ﻨﺎی ﻮ ای و ﺳﺎﺘﺎر اﻤﺎﯽ

ﺤ ﻮر ﻮل اﯿﺎﯽ و زی -ﻪ ﯿﺎﺑﺎن ی ی اان ﻮرد رﯽ و و ﻞ ار ( ﻞ ۴ و ۵).

ﺑﺎ رﯽ ﺳﺎﺘﺎر اﻤﺎﯽ ﻮر ﻮل اﯿﺎﯽ ﺘﺎ ﺞ ﺸﺎن داد ﻨﺎی ﻮ ای دا ﻞ ﻪ ای ﯽ ااﺶ ﻮل اﯿﺎﯽ ااﺶ ﯾﺎ اﻣﺎ

ﺤ ﺮات ﻨﺎی ﻮ ای ﻈﺎم ﯽ را ﯽ ﻮد. و ﻞ داده ی ﻨﺪﻻ ای ااﺶ ﻨﺎی ﻮ ای ﻮر را ﻮل ﻻ ی ﻮﭼﮏ رگ

ھ ﺸﺎن داد. ﺷﺎﺺ ﻮاری ﻮل اﻦ ﺳﺎﺘﺎر ﻪ ای ﮐﺎﺶ ﭘﯿﺪا د. زﻮم ﭘﻦ دﺖ دارای ﻨﺎی ﻮ ای ی زﻮم ﯿﺎﺑﺎن ﻮد. و

ﺤ WWF ﻞ ا ﺑﺎ ء ﻨﺪی ﻪ ای ﺸﺎن داد ﻪ ی ﻮاﯽ ز ﯽ ﯽ واﺪی زﺖ-اﯿﺎﯽ ﻮر اان ﺸﺎن ﯽ دﺪ و

ﻨﺎﺐ ای ﻄﺎﻌﺎت ﻔﺎﯽ ﯽ ﺑﺎﺪ د ﻞ آﻧﻪ ﻦ ﺮات ﻨﺎی ﻮ ای اﻦ ﯿﺎس و اﻦ ﻻ رخ داد. ﮫﺎر ﺳﺎﺘﺎر ﺺ ﻮ ای

ﻮل ﮫﺎر زی - ﻪ ﻮرد ﻄﺎﻪ ﺸﺎﺪه ﺪ. ا ﯾﮏ ﻮرد ز ﻨﺎﻪ ﺪه اﻦ ﮫﺎر ﺳﺎﺘﺎر ﺺ ﻮ ای ﻔﺎوت از ز ﺺ ﺪه ﻪ ی

WWF ﺤ ﻮد. اﻦ ﯾﺎﻪ ﺸﺎن ﯽ دﺪ ﺖ و ﺪرت زی ﻮاﯽ ز ﯽ ﺑﺎﯾﺪ ﻮرﺪاﮔﺎ ای ﻮرد ﻄﺎﻪ ار ﺮد. و ﻞ ا ﯽ و ا

ھ اﯿﺎﯽ روی ﻨﺎی ﻮ ای و ﺳﺎﺘﺎر ﻮ ای ﻮر ﻮل اﯿﺎﯽ و ﻦ زی -ﻪ ﯿﺎﺑﺎن ی ی اان ﺸﺎن داد ﺮاﻂ آب و

ﻮای ﻌﺎﺮ ﻣﺎﻨﺪ ﺑﺎرﺪﯽ و ﺮات دﻣﺎﯽ و ﻮﯿﺪ اوﻪ ﯿﺎﯽ دارای ﺮ زﯾﺎدی روی ﻮع ﻮ ای و ﺳﺎﺘﺎر ﻮ ای ﻮر ﻨﺪ. ﺣﺎ ﻞ ﻘﺎت

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ﻮﯿﮏ و ﻮﻮﯿﮏ روی ﻮر ی اان ارش ﯾﮏ زﺧﺎﻮاده، ﺖ ﺲ و ﯽ و ﻮ ﺪﯾﺪ ای اان و دو ﻮ ﺪﯾﺪ ای دﯿﺎ ﻮد. ﻋﻼوه

اﻦ ای اوﻦ ﺑﺎر ﺖ ﻮ ی ﻮر ی اان ﺰ ﺮ دﯾﺪ.

ﺧﺎﻪ، ﻮاﯽ ﺸﮏ و ﻪ ﺸﮏ اان دارای ﻨﺎی ﻮ ای ﺘﺎ ﺑﺎﻻﯽ ﯽ ﺑﺎﻨﺪ. از ﻮد ﻮ ﻮر دا ﻞ ﮫﺎرده ﻪ ﻔﺎﺖ ﺪه ﻮل دو

ﺤ ﺳﺎل ﻊ آوری دﯾﺪ. ا ﻨﺎی ﻮ ای دا ﻞ ﻪ ای ﺘﺎ ﭘﺎ ﻮد و از ﺪه ﺠﺎوز ﻮد، و ﻞ ی اﺠﺎم ﻪ ﺸﺎن داد ﺰان

ﺮات ﻮ ای ﻤﺎﯽ ﯿﺎس ی اﯿﺎﯽ زﯾﺎد و ﯽ دار ﯽ ﺑﺎﺪ ﺸﺎن ﯽ دﺪ از ﻮ ی ﯿﺎری از ﻮر ﻮاﯽ ﺸﮏ و ﻪ ﺸﮏ اان

دارای ﺮﭘاﻨﺪﯽ ﺪود ﯽ ﺑﺎﻨﺪ. اﻦ اﻟﻮی ﺮﭘاﻨﺪﯽ ﻮر ﺸﺎن ﯽ دﺪ ﻨﺎﻖ ﺸﮏ ﮫﺎ ﻨﺎﯽ وﻊ و ﯾﻮاﺖ ﯽ ﺑﺎﻨﺪ. ﺧﻼف آن اﻦ ﻨﺎﻖ

دارای ﺳﺎﺘﺎری ﺠﺎﺲ ﻮده ﺮﭘاﻨﺪﯽ ﯿﺎری از ﻮ را زA ﮕﺎه ی ﻮﭼﮏ ﺪود ﯽ ﻤﺎﯾﺪ و ﺳﺎﺘﺎری ﺗﻪ ﺗﻪ ﻮع ﻮ ای ﻮر ﭘﺪﯾﺪ ﯽ آورد.

ﺮﭘاﻨﺪﯽ ﺪود ﺪه ﯿﺎری از ﻮ ی ﻮر آﮫﺎ را ا ااض ﻪ ای و ﻪ ای آ ﭘﺬ ﯽ ﻤﺎﯾﺪ . ﻋﻼوه اﻦ از آﺠﺎ ﻨﺎی ﻮ ای و ﺳﺎﺘﺎر

ﻮ ای ﻮر ﺰان زﯾﺎدی ﺮاﻂ آب و ﻮاﯽ ﺘﯽ دارد، ﺮات آب و ﻮا ﯽ ﺶﯿﭘ ﺪه ﻪ ﻣﺎﻨﺪ ااﺶ ﺰان ﯽ و دﻣﺎ و ﺮات

ﺮده د رﺰان ﺑﺎرش ﺑﺎران اﻤﺎل زﯾﺎد ﺮ زﯾﺎدی روی ﻮر ﻮاﺪ داﺖ. اﻦ ﺮان اﻤﺎل اوان ، ﺮ ﺮﭘاﻨﺪﯽ ﯽ از ﻮر

ﻮاﺪ ﻮد و ﻮر ﯽ ﻮاﯽ ﺳﺎزﮔﺎری ﺑﺎ ﺮاﻂ ﺪﯾﺪ را ﺪاﻪ ﺑﺎﻨﺪ و ﻮاﻨﺪ ﭘﻨﺎﮕﺎه ﻨﺎﯽ ﯿﺎﻨﺪ ﯿﺎس ﻪ ای و ﻪ ای ض ﻮاﻨﺪ ﺪ.

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General Appendix

Figure 1: Photographs of 8 study sites in Iran. They are from top left to right down: Siahkooh National Park, Turan National Park, Naiband National Park, Saghand National Park, Khoshyelagh Wildlife Refuge, and Golestan National Park.

General Appendix

Figure 2: Color automontage pictures of two new described ants of genus Cataglyphis from Iran: Cataglyphis stigmatus (left) and Cataglyphis pubescens (right). Photographs by Hans-Peter Katzmann © Antbase.net.

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General Appendix

Figure 3: Pictures of the second field work in 2008. From tope left to right down: 1- Plastic quadrate used for measuring habitat heterogeneity in sampling sites, 2- Measuring the vertical complexity of habitat by a plastic rod. 3- Taking soil samples from the sample plots, 4- Direct collecting ants from plants, 5- Measuring plant diversity across 40 m transect in a sample plot, 6- Breakfast under shade of the car.

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Acknowledgments

Foremost, I would like to thank my supervisory panel. I am grateful to Martin Pfeiffer, my advisor, who in 2005, accepted me as a PhD student and gave me the chance to embark for this journey. He helped me to write a research proposal which was successful in being funded by German Academic Exchange Service (DAAD). I am thankful for his supervision, guidance, and discussions on topics of this work which all has greatly assisted my professional development. Special thanks to him for giving me the freedom to pursue my own interests in this dissertation and the friendship he has given me during my work on this dissertation. I would also like to thank Elisabeth Kalko whose support made this work possible. Especially her kindness and friendship has been always supportive for me. I thank Manfred Ayasse, Marco Tschapka and Steven Jansen for accepting to be members of the promotion board. This research would not have been possible without financial support. I acknowledge the financial support from the DAAD. As a scholarship for four years, this support made me able to attend a German language course in Goethe Institute in Dresden for four months and to conduct my PhD at the University of Ulm. Conducting this project in Iran was not possible without my uncle Mohammad who joined me in both of my field works in 2007 and 2008. Driving more than 12000 kilometers on roads and helping me in collecting ants and soil samples were an invaluable assistance. I have been fortunate throughout my thesis to have worked with excellent ant taxonomists: Alexander Raschenko, Bernhard Seifert and Donat Agosti. I joined them in their labs or in field and I have been taught how difficult and how long is the way of learning ant taxonomy and they pushed me to always reach further knowledge. I thank my co-authors, Martin Pfeiffer, Alexander Radchenko, and Helen

Acknowledgments

Alipanah for their invaluable contributions. Martin contributed in all manuscripts by his advice and by his patience for reading earlier versions of all my manuscripts. Alex contributed in three manuscripts, one as a first author, and opened new horizons for me in taxonomy and faunistic studies on ants. Helen shared here great data sets on Iranian ants with me for publishing Iranian ant species list. I want to thank Jakob Fahr for all long discussion which we had on “doing partitioning or not doing partitioning” on the balcony of our institute. Dirk Mezger and Hans-Peter Kazmann were great company in our small “Ameisen-Gruppe”. Dirk gave valuable comments on earlier version of my two manuscripts and was a great company during identification of ants in lab. Hans-Peter made beautiful photos from Iranian ants. I want to thank Matthias Herkt for his helpful answers to my GIS questions. I thank Markus Metz for the GRASS course that he gave to me. I want to thank Konstans Wells for his support and for encouraging me to learn R and modern statistical knowledge. Further he gave his comments on earlier versions of chapter 5. Lou Jost, Thomas Crist, Etienne Laliberté, Guillaume Blanchet, John Fox, Ramon Diaz-Uriarte, Thiago F. Rangel and Pedro Peres-Neto are acknowledged for their answers to my statistical questions. I thank John Fellows, Ashwin Chari, Bryson Voirin, and Parimah Kazemi for their contribution in language correction of the manuscripts. I want to thank editors of various journals, Alan Andersen, Katsuyuki Eguchi, Birgit Schlick-Steiner, and Florian Steiner who improved my papers by their constructive comments. Nathan Sanders, Rob Dunn, John Fellows, Masashi Yashimura, and several anonymous reviewers have strengthened this dissertation by their insightful comments on my manuscripts. I want to thank Himender Bharti, Mostafa Sharaf, Cedric Collingwood, and Dmitry Dubovikoff who helped me in various ways during my taxonomic and faunistic work. At the institute, I would like to thank my room-mate, Kirstin Übernickel, for her friendship and her helps, especially for proofreading of parts of this dissertation. I am grateful to other room-mates during these four years, Tanja Weis-Dootz, Stefan Klose, Joana Fietz, Larissa Albrecht, Ramón Seage Ameneiros (Moncho), and Monika Riepl. They, all, made a great friendly atmosphere around me. Ingrid Dillon was much more helpful than a secretary to me. I am very grateful for the friendship of Robert Hodgkison, Kirsten Jung, Christoph Meyer, Silke Berger, Marion Gschweng, Marco Mello, Inga Geipel, Katrin Heer, Taina Conrad, Sebastian Zschunke, Andrea Weiß, Mirjam Knörnschild, Ralph Simon, Swen Renner, Reisla Oliveira, Clemens

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Acknowledgments Schlindwein, Ulrike Stehle, and Gabriele Wiest. Stefan Jarau, Kirsten Kreuter, and Ann-Marie Rottler were great company during travel to Copenhagen for IUSSI conference. I would like to thank Julia Dolezil, Stefan Böhm, Maria Helbig, Simon Ripperger, and all other people who played soccer with me each Friday evening. They all made the Bio III a lovely place to work and live. I am thankful to Marco Tschapka and Heiko Bellmann for their organization and helps in different ways. I would also like to thank Irmi Pfeiffer for her company during the first field trip in 2007 in Iran and all these years in Ulm. I am grateful to many Iranian colleagues and friends, Mohammade Medadi, Mohammad Montazemi, Hossein Zohuri, Alinaghi Maghsudlu, Alireza Naderi, Marayme Peymani, Mohsen Jafari and Hadi Ashraf, who helped and guided me in various ways, especially for granting research permits to work in natural reserves. I also acknowledge Farhad Khormali who accepted to analysis my soil samples in his lab in Gorgan University. I am thankful to my colleagues in Gorgan University, especially Gholam Reza Haddadchi, Haj Gholi Kami, Hamid Sadeghipour, Mahnaz Aghdasi, Mohammad Bagherie who supported me and encouraged me to start this journey. During two field works in 2007 and 2008, in fourteen ranger’s stations, I had this chance to know many nice people who worked hard and put their life in danger to save nature. Among them I remember well Saeed Azizi who I met him in 2007 in Khoshyelagh wildlife refuge, but coming back to this station in 2008, he had passed away in an accident during his job. I am very thankful to my friends, Alireza Ghasemzade, Hossein Zamiri and his family, Mehdi Ghasemi, Mehdi Pordel, Amir Masudi, Siavash Afzali, Mehdi Sabaghpour, Jafar Jafari, Ali Bagherian, Ghasem Ghasemi, and Davood Ghaderi for keeping me warm and close, and to give me comfort and advice. Last but not least, I am grateful to my wife Elham Kashani, for her love, patience and encouragement and to my family for supporting me throughout my life and as well as throughout my PhD. I deeply appreciate their love and faith.

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Curriculum Vitae OMID PAKNIA University of Ulm, Institute for Experimental Ecology, Albert Einstein Allee 89069, Ulm, Germany Email: [email protected] Phone: +49 731 50-22668 Fax: +49 731 50-22683

Personal Information First, Last Name Omid Paknia Date of Birth 23 July 1977 Place of Birth Tehran, Iran Citizenship Iran Marital status Married

Scientific Education

1991 - 1995 High school at Prof. Hashtroodi high school in Tehran, Iran. 1995 University entrance exam (Natural Science). 1995 - 1999 B. Sc. in Biology, Gorgan University, Gorgan, Iran. 1999 - 2002 M.Sc. in Medical Entomology, Tarbiat Modaress University, Tehran. 2003 - 2006 Research scientist and instructor, Institute of Biology, Gorgan University. Jun.- Oct. 2006 German language course in Goethe Institute in Dresden, Germany. Oct. - Dec. 2006 Ecology module, Ulm University, Ulm, Germany. 2007 - present PhD in Ecology, University of Ulm, Germany.

Awards and Fellowships 1999 First student of M. Sc. entrance exam. 2006 DAAD full PhD fellowship

Scientific Referee Journal of Arid Environments Asian Myrmecology African Journal of Environmental Science and Technology African Journal of Biotechnology Jordanian Journal of Agricultural Sciences Journal of Plant Breeding and Crop Science

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Curriculum Vitae

List of Publications

Paknia O, Pfeiffer M (accepted) Hierarchical partitioning of ant diversity: implications for conservation of biogeographical diversity in arid and semi-arid areas. Diversity and Distributions.

Radchenko A, Paknia O (2010) Two new species of the genus Cataglyphis Foerster, 1850 (Hymenoptera: Formicidae) from Iran. Annales Zoologici 60: 69-76

Paknia O, Radchenko A, Pfeiffer M (2010) New records of ants (Hymenoptera: Formicidae) from Iran. Asian Myrmecology 3:29-38

Hojati, V. Kami, H.G, Golmohammadi, M., Paknia O (2009) Diversity and community structure of ants in desert and steppe regions of Damghan County [Persian]. Iranian Journal of Zoology 1: 3-10.

Paknia O, Radchenko A, Alipanah H & Pfeiffer M (2008) A preliminary checklist of the ant fauna (Hymenoptera: Formicidae) of Iran. Myrmecological News 11: 151- 159

Paknia O, Kami HG (2007) New and additional record for Formicid (Hymenoptera: Insecta) fauna of Iran. Zoology in the Middle East 40: 85-90

Paknia O (2006) Distribution of the introduced ponerine ant Pachycondyla sennaarensis (Hymenoptera: Formicidae) in Iran. Myrmecological News 8: 235-238

Tirgari S, Paknia O (2005) First record of ponerine ant (Pachycondyla sennaarensis) in Iran and some notes on its ecology. Zoology in the Middle East 34: 67-70

Tirgari S, Paknia O (2004) Additional records for the Iranian Formicidae fauna. Zoology in the Middle East 32: 115-116

Conference Presentations

XVIth Congress of International Union for the study of social Insects (IUSSI), Copenhagen, Denmark Paknia O, Pfeiffer M (2010) Determinants of ant community composition along a 1200 km latitudinal gradient in arid and semi-arid regions of Iran. Poster presentation.

First Central European Meeting of International Union for the study of social Insects (IUSSI), Chiemsee, Germany Paknia O, Pfeiffer M (2009) Spatial patterns in diversity of ant communities in Iran along a north-to-south environmental gradient. Oral presentation.

154 Curriculum Vitae

7th ANeT meeting Jakarta, Indonesia Paknia O, Pfeiffer M (2009) A hierarchical analysis of species richness and diversity of ants in Iran. Poster presentation.

6th ANeT meeting, Patiala, India Paknia O, Pfeiffer M (2007) Myrmecological studies along ecological gradients in Iran - questions and methods. Poster presentation. Vakhide N, Paknia O, Habibi H (2007) Description of specific behaviors of acrobat ant Crematogaster sp. in Babolsar climate, Iran. Oral presentation. Vakhide N, Bagherian A, Paknia O (2007) Diurnal and seasonal activity patterns in the acrobat ants Crematogaster sp. in Babolsar, Iran. Oral presentation. Bagherian A, Akbarzadeh M, Paknia O (2007) Diurnal and seasonal activity pattern in invasive ant Lasius neglectus (Formicinae) in South coast Of Caspian Sea, Iran. Oral presentation. Bagherian A, Dehghani, AA. Paknia O (2007) The use of artificial neural networks in ecological analysis: estimating activity pattern in ants (Crematogaster: Formicidae). Oral presentation.

5th ANeT meeting, Kuala Lumpur, Malaysia Paknia O, Kashani, E (2005) Distribution of invasive ants Pachycondyla sennaarensis in Iran. Oral presentation. Paknia O (2005) Ant collection in Zoological Museum of Gorgan University (ZMGU), Iran. Oral presentation.

First Biological Evolution Conference, Mashhad, Iran Paknia O (2005) Brief review of phylogeny of Formicidae subfamilies. Poster presentation.

13th Iranian Biology Conference, Rasht, Iran Paknia O, Kami HGH, Nikmahzar E (2005) New records of Centipedes fauna of Iran. Poster presentation.

First International Iranian Congress on Biological sciences, Karaj, Iran Paknia O, Rahmani Z (2005) Preliminary fauna and abundance of ant subfamilies in Bojnurd County. Poster presentation.

XXII International Congress of Entomology, Brisbane, Queensland, Australia Tirgari S, Akbarzadeh K, Nateghpour M, Paknia O (2004) First report on the presence and medical importance of stinging ant in Southern Iran (Hym: Formicidae: Ponerinae). Oral presentation.

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Curriculum Vitae

16th Congress of plant protection, Tabriz University, Tabriz, Iran Paknia O, Tirgari S (2004) Study of fauna and geographical distribution of stinging ants (Hymenoptera: Formicidae) and their medical importance in Lar, Iran. Poster presentation.

XI Iranian Biology Conference, Urima, Iran. Paknia O, and Kashani E (2003) The study of ants (Formicidae: Hymenoptera) fauna in Lar city. Poster presentation.

Membership International Biogeography Society (IBS) Iranian Medical Entomology Society

Languages Persian, English, German

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Eidesstattliche Erklärung

Hiermit erkläre ich, die vorliegende Dissertationsarbeit selbstständig angefertigt und keine anderen als die in der Arbeit aufgeführten Hilfsmittel verwendet zu haben. Wörtlich oder inhaltlich übernommene Stellen wurden als solche gekennzeichnet.

Ulm, 13.10.2010

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