Diversity
Di
it and
d species
i composition
iti
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
ii To mom, dad, Elham, Elham, Arman and Ashwin
iii iv Contents
1
General introduction
1
Drylands
Biodiversity in drylands
Objectives of this study
Contributions of co-authors
References
1
1
4
10
11
2
Desert versus steppe: Multi-scale spatial patterns in diversity of ant
communities in Iran
15
Abstract
Introduction
Methods
Results
Discussion
Acknowledgments
References
Appendix S1
15
16
18
22
27
32
33
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
v 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
Abstract
105
Introduction
105
Material and Methods
105
Results
106
Discussion
110
Acknowledgments
111
vi 105
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
vii viii 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 macrodetritivores 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 semiarid 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-tobroad 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
0
Caspian Sea
150
300
Km
38° N
1
2
36° N
3
4
5
34° N
Iran
7
11
10 9
6
32° N
8
12
30° N
Central Persian desert basins
13
14
28° N
Persian Gulf
26° N
Elburz Range forest steppe
South Iran Nubo-Sindian desert
Zagros Mountains forest steppe
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.
9 Chapter 1 - General Introduction
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 coauthor 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.
10
Chapter 1 - General Introduction
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, reidentified 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.
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11 Chapter 1 - General Introduction
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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
0
Caspian Sea
100
200
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
28°0'0"N
8
Central Persian desert basins
Elburz Range forest steppe
South Iran Nubo-Sindian desert
Zagros Mountains forest steppe
44°0'0"E
46°0'0"E
48°0'0"E
50°0'0"E
Persian Gulf
52°0'0"E
54°0'0"E
26°0'0"N
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
diversity by its multiplicative definition (eHα × β = eHγ). 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
Number of species
60
50
40
Sobs
Singletons
Doubletons
Chao2
30
20
10
0
0
200
400
600
800
1000
1200
1400
1600
1800
Number of species occurences
b)
Biomes
50
Number of species
40
30
Sobs desert
Singletons desert
Doubletons desert
Chao2 desert
Sobs steppe
Singletons steppe
Doubletons steppe
Chao2 steppe
20
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. Plot or level name
SOCs Sobs
95 % LB
ᾤ ‫ﭜڮ‬
eH
95 % LB 95 % UB
Eobs
Unique
Duplicate
Desert biome
Steppe biome
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
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
Kavir B
119
143
8
10
8
9
8
10
6.14
7.68
5.49
6.94
6.8
8.42
0.77
0.77
0
0
1
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
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
Mirzabailu
Khoshyelagh
211
248
16
15
14
13
17
16
9.2
9.94
8.12
8.95
10.28
10.94
0.58
0.66
2
1
3
1
Kolahghazi
183
10
10
10
7.00
6.18
7.81
0.7
0
0
Dena
248
14
12
15
10.1
9.23
10.96
0.72
1
2
1
Desert
biome
Steppe biome
Locale level
Turan
331
12
11
12
8.82
8.24
9.4
0.74
0
Kavir
262
11
11
11
7.29
6.69
7.91
0.66
0
1
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
459
431
593
361
24
24
17
24
21
23
17
18
26
25
17
30
13.18
17.08
11.39
14.03
12.03
15.89
10.72
12.82
14.32
18.28
12.07
15.26
0.55
0.71
0.67
0.58
3
1
0
5
3
2
0
1
890
954
45
37
43
31
47
43
29.24
19.77
27.63
18.63
30.86
20.92
0.65
0.53
3
5
3
1
69
65
73
38.9
37.25
40.57
0.56
6
4
Ecoregional level
Elburz steppe
Zagros steppe
Central desert
Nubo desert
Biome level
Steppe biome
Desert biome
Regional level
Total transect
24 1844
Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants
a)
14
Alpha species richness
13
12
11
10
9
8
27
28
29
30
31
32
33
34
35
36
37
33
34
35
36
37
Latitude
b)
1.0
Beta species richness
0.8
0.6
0.4
0.2
0.0
27
28
29
30
31
32
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-steppedesert-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.
Species richness
Observed
Expected
Shannon diversity
Observed
Expected
28.6 ***
18.9 ***
8.5
2.4 ***
10.7 ***
4.7
6.1
8.2
10.4
39.6
1.6 3***
1.77***
1.56***
1.18 ***
7.33 ***
1.02
1.04
1.08
1.16
28.55
Steppe
β ecoregion
β locale
β fine
α fine
21 ***
10.07 ***
2.65 ***
11.28 ***
2.5
3.8
6.1
32.6
1.96 ***
1.64 ***
1.19 ***
7.65 ***
1.03
1.06
1.13
23.54
Desert
β ecoregion
β locale
β fine
α fine
17.33 ***
7.16 ***
2.27 ***
10.24 ***
3.6
4.1
4.6
24.6
1.61 ***
1.49 ***
1.17 ***
7.07 ***
1.02
1.05
1.09
16.84
Partition/Scale
Whole region
β biome
β ecoregion
β locale
β fine
α fine
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 βShannon 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.
1.0
50
45
Species richness
Shannon diversity
Evenness
(a) steppe
0.8
35
0.6
30
25
0.4
20
Evenness
Effective number of species
40
15
0.2
10
5
0
Fine
Local
Ecoregional
Biome
0.0
Spatial scales
1.0
40
35
Species richness
Shannon diversity
Evenness
(b) desert
25
0.6
20
0.4
15
Evenness
Effective number of species
0.8
30
10
0.2
5
0
Fine
Local
Ecoregional
Biome
0.0
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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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
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 (ANDRE, 1881)
Cataglyphis nodus BRULLE, 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
36 1a
1b
2a
2b
3a
3b
4a
4b
5a
5b
6a
6b
7a
7b
8a
8b
1
24
10
1
-
1
2
3
6
17
5
19
-
11
1
28
26
2
1
-
19
1
30
24
16
4
14
-
2
17
1
22
25
28
9
-
1
11
28
28
25
1
5
-
6
23
27
12
-
9
10
22
27
17
-
29
10
-
29
5
6
-
17
13
13
2
18
29
1
18
-
8
1
23
30
-
15
5
13
1
5
9
-
1
7
28
-
20
16
6
1
3
-
9
12
11
4
-
Chapter 2 – Desert vs. Steppe: Spatial Patterns of Ants
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)
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. 4
Tetramorium sp. 5
20
2
4
1
1
8
11
-
25
7
7
5
1
7
23
-
7
2
2
8
-
12
3
8
3
3
23
-
1
1
30
5
25
-
1
5
30
3
27
-
2
30
3
16
-
4
2
2
30
20
-
20
2
2
7
6
2
-
17
4
6
30
4
4
-
9
4
3
2
14
-
3
21
8
5
6
18
7
1
1
13
1
30
30
5
1
-
8
2
29
3
1
-
7
10
8
19
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 semiarid 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 scaledependent (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
Iraq
Iran
40°0'0"N
Afghanistan
0
Caspian Sea
150
300
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
8
Central Persian desert basins
Elburz Range forest steppe
South Iran Nubo-Sindian desert
Persian Gulf
Zagros Mountains forest steppe
44°0'0"E
46°0'0"E
48°0'0"E
50°0'0"E
52°0'0"E
54°0'0"E
26°0'0"N
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 72hour 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
no.
1
2
3
4
5
6
7
8
Plot name
Golestan NP
Khoshyelagh WLR
Turan NP
Kavir NP
Kolahghazi NP
Dena PA
Mond PA
Naiband PA
Elevation
(m a.s.l.)
1217
1580
1190
1073
1724
2728
17
15
Ecoregion
Elburz Range forest steppe
Elburz Range forest steppe
Central Persian desert basins
Central Persian desert basins
Zagros Mountains forest steppe
Zagros Mountains forest steppe
South Iran Nubo-Sindian desert
South Iran Nubo-Sindian desert
Precipitation
mm year^-1
185.3
165.4
126.9
122.3
122.8
862.9
241.6
236.5
Mean daily air
temp (C°) year^-1
13.1
12.5
16
18.4
16.2
15.2
27.1
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). Intersite 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).
46 Chapter 3 – Hierarchical Partitioning of Ant Diversity
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
Species number
20
15
10
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 αdiversity (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
48 landscape
showed a complete change in species
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.
Diversity
Species richness
Observed Expected
P
Observed
Shannon diversity
Expected
P
Whole region
α local
β local
β landscape
β ecoregion
γ
10.38
1.26
1.68
3.14
69
39.62
1.26
1.16
1.19
69
< 0.0001
0.565
< 0.0001
< 0.0001
7.33
1.18
1.56
2.88
38.93
29.8
1.14
1.08
1.06
< 0.0001
< 0.0001
< 0.0001
< 0.0001
12
1.29
1.55
24
18.75
1.15
1.11
< 0.0001
< 0.0001
< 0.0001
8.2
1.17
1.37
13.14
12.12
1.05
1.03
< 0.0001
< 0.0001
< 0.0001
9.75
1.23
2
24
20.95
1.09
1.05
< 0.0001
< 0.0001
< 0.0001
7.11
1.22
1.98
17.17
15.64
1.06
1.03
< 0.0001
< 0.0001
< 0.0001
10.5
1.1
1.47
17
15.43
1.08
1.02
< 0.0001
0.146
< 0.0001
7.48
1.08
1.4
11.31
10.9
1.03
1.01
< 0.0001
< 0.0001
< 0.0001
9.5
1.47
1.71
24
17.93
1.16
1.15
< 0.0001
< 0.0001
< 0.0001
6.48
1.32
1.64
14.03
12.7
1.07
1.04
< 0.0001
< 0.0001
< 0.0001
Ecoregions
Elburz
α local
β local
β landscape
γ
Zagros
α local
β local
β landscape
γ
Central desert
α local
β local
β landscape
γ
Nubo
α local
β local
β landscape
γ
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).
49 Chapter 3 – Hierarchical Partitioning of Ant Diversity
100
75
Information Remaining (%)
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).
50 Chapter 3 – Hierarchical Partitioning of Ant Diversity
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;
54 Chapter 3 – Hierarchical Partitioning of Ant Diversity
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).
55 Chapter 3 – Hierarchical Partitioning of Ant Diversity
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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
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)
1
2
3
4
5
6
7
8
+
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
+
+
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 +
+
+
+
+
+
-
+
+
+
+
+
+
-
+
+
-
+
+
-
+
+
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+
+
+
+
+
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+
+
+
+
+
-
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
Afghanistan
Caspian Sea
1
Pakistan
2
Saudi Arabia
36°0'0"N
Iran
38°0'0"N
Turkmenistan
Turkey
Iraq
40°0'0"N
et al., 2006; Legendre, 2008; but see Smith & Lundholm, 2010).
3
32°0'0"N
34°0'0"N
4
5
28°0'0"N
30°0'0"N
6
7
0
44°0'0"E
125 250
Kilometers
46°0'0"E
48°0'0"E
50°0'0"E
26°0'0"N
8
Persian Gulf
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 distancebased 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.
Site no.
Site name
Phytogeographical
regions
Elevation (m a.s.l.)
Rainfall
Summer rainfall
Mean daily
Temp. range
(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
Mantel r
0.4
0.3
0.2
0.1
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
Annual temperature range
1.6
1.2
Moran's I
0.8
0.4
0.0
-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
E
S
E+b+S
E|S
b
S|E
d
d.f.
3
3
6
3
0
3
-
R2
0.51
0.33
0.77
0.66
0.53
-
Adjusted R2
0.39
0.17
0.62
0.45
-0.06
0.23
0.38
P value
0.001
0.003
0.001
0.001
Un-testable
0.001
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)
Central desert
Northern steppe
0.5
Kh-B
Kh-A
RDA2
Mi-A Mi-B
summ
er rain
fall
te
0.0
an
al
nu
ra i
Tu-A
Tu-B
e
ng
. ra
p
m
ll
nfa
Ka-B
Ka-A
Ko-A
Ko-B
Coastal desert
Na-A Mo-A
-0.5
Na-B
Mo-B
De-A
Western steppe
-1.0
-1.0
De-B
-0.5
0.0
0.5
RDA1
1.5
b)
M o-B
1.0
M o-A
RDA2
0.5
Ka-A
Mi-A
Ko-A
Ko-B
Ka-B
pcn
Mi-B
0.0
m5
Kh-A
3
nm
pc
Kh-B
De-B
pc
nm
6
De-A
-0.5
Tu-B
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
Species
Site
Indicator
value (IV)
Mean
Standard
Deviation
P
value
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</key></foreign-Figure 2 and 3) and prevents their
colonization to further habitats with unfavorable conditions. Nevertheless, in this
dominant environmental process, some species can inhabit in sites with unfavorable
conditions, when dispersal is efficient enough. These species might be poor
competitors in these new sites, because of bad environmental conditions, but they can
survive because of the mass effect mechanism, as has been shown by the existence of
this mechanism in the Iranian ant metacommunity. The mass effect perspective
predicts that species, which are not good competitors in a local assemblage, are not
excluded from this assemblage if they are rescued by immigration from assemblages
in which they are dominant. An example of the mass effect mechanism is
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).
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81 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages
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/).
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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 multitemporal NDVI. Global Ecology and Biogeography, 13, 221-235
83 Chapter 4 - Environmental-Spatial Drivers of Ant Assemblages
84
5
Productivity, 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
86
Chapter 5 - Ant Species Richness in a Dryland Habitat
(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).
88
Chapter 5 - Ant Species Richness in a Dryland Habitat
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
Productivity
NDVI
Heterogeneity
Habitat Heterogeneity
Tolerance
Minimum temperature
Biotic interaction
Plant species richness
Ambient energy
Mean annual temperature
Explanation
Available energy in the environment directly affects ant
species richness
Heterogeneity of habitat facilitates the coexistence of a
greater number of species
Ant species richness is limited to those species that can
tolerate the local condition
Plant species richness in the environment directly affects
ant species richness
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 nonparametric 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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Chapter 5 - Ant Species Richness in a Dryland Habitat
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
Latitude
35
34
33
32
0
2
4
6
8
10
12
14
16
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).
93 Chapter 5 - Ant Species Richness in a Dryland Habitat
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
94
Chapter 5 - Ant Species Richness in a Dryland Habitat
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.
95 Chapter 5 - Ant Species Richness in a Dryland Habitat
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
1
2
3
4
5
6
7
8
9
96
Model
Productivity
Tolerance - Productivity
Ambient energy - Productivity
Ambient energy - Tolerance - Productivity
Complexity - Productivity
Tolerance - Biotic interaction - Productivity
Tolerance - Complexity - Productivity
Biotic interaction - Productivity
Ambient energy - Complexity - Productivity
K
5
6
7
6
6
7
7
6
7
AICc
167.24
167.71
169.97
170.33
170.34
170.36
170.41
170.58
173.09
∆ AICc
0.00
0.47
2.73
3.09
3.11
3.12
3.17
3.34
5.85
AICc wt
0.29
0.23
0.07
0.06
0.06
0.06
0.06
0.05
0.02
Cum wt
Moran’s I
P value
0.29
0.52
0.59
0.65
0.71
0.77
0.83
0.88
0.90
0.032
- 0.002
- 0.07
0.031
0.046
- 0.02
0.007
0.034
0.055
0.33
0.40
0.55
0.31
0.30
0.47
0.42
0.34
0.29
Chapter 5 - Ant Species Richness in a Dryland Habitat
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
No terms added
Log-likelihood
-93.177
Df
Chi-squared
P value
Productivity
Tolerance - Productivity
Ambient energy - Productivity
Ambient energy - Tolerance - Productivity
Complexity - Productivity
Tolerance - Biotic interaction - Productivity
Tolerance - Complexity - Productivity
Biotic interaction - Productivity
Ambient energy - Complexity - Productivity
-77.701
-75.938
-77.305
-75.831
-77.492
-75.847
-75.871
-77.610
-77.211
4
5
5
6
5
6
6
5
6
30.952
34.477
31.745
34.693
31.37
34.659
34.611
31.134
31.932
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<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
NDVI
NDVI2
Temperature range
Mean annual temperature
Plant species richness
Complexity
Importance
0.98
0.44
0.39
0.27
0.16
Coefficient
2.75
- 0.83
- 0.17
0.1
0.02
0.01
Unconditional SE
0.57
0.35
0.03
0.06
0.03
0.01
95% Lower
0.63
- 1.53
- 0.31
0.01
- 0.04
- 0.02
95% Upper
3.85
- 0.13
- 0.09
0.25
0.09
0.01
97 Chapter 5 - Ant Species Richness in a Dryland Habitat
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.
98
Chapter 5 - Ant Species Richness in a Dryland Habitat
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.
99 Chapter 5 - Ant Species Richness in a Dryland Habitat
(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.
100
Chapter 5 - Ant Species Richness in a Dryland Habitat
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).
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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, AlbertEinstein 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
out in several parts of Asia (e.g., East and Southeast Asia:
KUPYANSKAYA 1990, TERAYAMA 1994, WU & WANG 1995,
CHUNG & MOHAMED 1996, IMAI & al. 2003, OGATA 2005,
RADCHENKO 2005; India: BHARTI 2002a, b; Central Asia:
TARBINSKY 1976, DLUSSKY & al. 1990, SCHULTZ & al.
2006, PFEIFFER & al. 2007); however, in the west and southwest of Asia only the ant fauna of Saudi Arabia has been
studied satisfactorily (COLLINGWOOD 1985, COLLINGWOOD
& AGOSTI 1996). The ant fauna of other countries from
this region, including Iran, has been investigated only partly.
Iran situates mainly on the Iranian plateau and covers
an area of 1,623,779 km2. The country is located in the
Palaearctic and naturally divided into six biomes and 14
ecoregions (OLSON & al 2001). At the same time, its borders are close to the boundaries of the Afrotropical and Oriental regions in the south and south-east of the country.
Such position, as well as high environmental diversity, promotes the diversity of the Iranian flora and fauna.
The history of the myrmecological investigations in Iran
can be divided into two main periods: the first third of the
XX century includes works of FOREL (1904a, b), EMERY
(1906), CRAWLEY (1920a, b, 1922) and MENOZZI (1927).
After that, until the 1990ies new data on the Iranian myrmecofauna were very scarce (e.g., ARNOLDI 1977). The
second period of ant research in Iran started in the 1990ies
and has continued until now. RADCHENKO (1994a, b, 1995,
1996b, 1997a, b), SEIFERT (2003) and TAYLOR (2006) described and recorded several species from Iran. ARDEH
(1994) recorded 13 ant species from the Karaj Region, Alipanah and colleagues studied the ant fauna of Tehran (ALIPANAH & al. 1995, 2000), of the southwest of Iran (Khuzestan province) (ALIPANAH & DEZHAKAM 2000), and of
several other parts of the country (ALIPANAH 2004). RADCHENKO & ALIPANAH (2004) documented the first Iranian
species of the subfamily Aenictinae. TIRGARI & PAKNIA
(2004, 2005) and PAKNIA & KAMI (2007) reported 21 species from various areas of the country.
As Iranian myrmecologists have published their reports
mostly in local journals or presented them at certain national scientific congresses, it is difficult for foreign myrmecologists to access this literature. Other problems arise
from old records that need revision. For all of these reasons
a preliminary checklist of the Iranian ants is a helpful reference for myrmecologists that are interested in Asian ants.
Material and methods
This study is based on a comprehensive review of literature and the examination of material both collected by two
of the authors of this paper (OP & HA), by Hamidreza Hajiqanbar and by many other entomologists as well as on investigated material preserved in the following museums
105
Chapter 6 - Ant Species List of Iran
and institutions: Hay Mayans Insect Museum in the Iranian Research Institute of Plant Protection, Tehran, Iran
(HMIM), Zoological Museum of Gorgan University, Gorgan, Iran (ZMGU), Institute of Zoology of Ukrainian National Academy of Sciences, Kiev, Ukraine (IZK), Zoological Museum of Moscow State University, Moscow, Russia (ZMMU), Zoological Institute of Russian Academy of
Sciences, St. Petersburg, Russia (ZISP), Museum and Institute of Zoology of Polish Academy of Sciences, Warsaw, Poland (MIZ), The Natural History Museum, London,
UK (BMNH), Museo Civico di Storia Naturale "Giacomo
Doria", Genoa, Italy (MSNG), Istituto di Entomologia, University di Bologna, Bologna, Italy (IEUB), and Naturhistorisches Museum Basel, Basel, Switzerland (NHMB).
Altogether, we examined about 3350 specimens. We
verified existence of the voucher specimens for all species
listed in Table 1, except for several species recorded by
CRAWLEY (1920a, b), MENOZZI (1927) and ANARAKI (1981)
as we could not locate their vouchers, and the Cardiocondyla species recently recorded for Iran by SEIFERT (2003).
For all species taxonomically described from Iran we assumed that there are vouchers for them. We avoided to list
unavailable names.
While there are no keys specifically for the identification of the ants of Iran, we used existing keys for adjacent regions and taxonomic revisions of different ant genera (ARNOLDI 1976, 1977, COLLINWOOD 1985, DLUSSKY
& al 1990, ARAKELYAN 1994, DLUSSKY & RADCHENKO
1994, RADCHENKO 1992a, b, 1994c, d, 1996a, b, 1998,
COLLINGWOOD & AGOSTI 1996), as well as unpublished
data and keys for several genera of A. Radchenko. In all
cases the correct names of the taxa were verified according to the catalogue of BOLTON & al. (2007) except for
Tetramorium forte FOREL, 1904 and T. chefketi FOREL,
1911. The software ArcView program was used to prepare
a map of all records of ants in Iran.
Dorylus sp.: Khuzestan province, Ahvaz, 4.X.1954, leg.
Sal, 1 worker (HMIM), det. H. Alipanah.
Messor minor (ANDRE, 1883): Hamedan province, Hamedan, 25.V.1997, leg. P. Moghadasi, 5 workers (HMIM),
det. A. Radchenko.
Ponera sp.: Golestan province, 20 km East Gorgan, Tuskestan forest, 28.V.2005, leg. Omid Paknia, 9 workers
(ZMGU), det. A. Radchenko.
Tapinoma karavaievi EMERY, 1925: Tehran province, Tehran: 19.X.1992, leg. Shemiranat; Lavasanat, 1600 m, 12.
VII.1994, leg. H. Alipanah, 2 workers (HMIM); Khuzestan province, Ahwaz, 23.XI.1997, leg. M. Dezhakam,
3 workers (HMIM); Khorasan Razavi province, Mashhad, 23.I.1999, leg. M. Ghasemi, 5 workers (HMIM);
Mazandaran, Miankaleh, 14.IV.2005, leg. O. Paknia, 6
workers (ZMGU), det. A. Radchenko.
Temnothorax parvulus (SCHENCK, 1852): Golestan province, Gorgan, Tuskestan forest, 28.V.2005, leg. S. Shalchian, (ZMGU), det. A. Radchenko.
Tetramorium inerme MAYR, 1877: Khorasan Razawi province, Sarakhs, 27.IV.2007, leg. H. Hajiqanbar, (IZK),
det. A. Radchenko.
Reidentified species
Based on the re-examination of voucher specimens (ZMGU,
HMIM) by A. Radchenko, several previously recorded
species are excluded from the Iranian fauna: Camponotus
cruentatus (LATREILLE, 1802) and C. micans (NYLANDER,
1856) (TAYLOR 2006) are C. turkestanicus EMERY, 1887
and C. xerxes FOREL, 1904, respectively; Messor galla
(MAYR, 1904) and Camponotus maculatus (FABRICIUS,
1782) (TIRGARI & PAKNIA 2004) are M. intermedius SANTSCHI, 1927 and C. armeniacus ARNOLDI, 1967, respectively; Tapinoma simrothi KRAUSSE, 1911 (PAKNIA & KAMI
2007) is T. karavaievi EMERY, 1925; Tetramorium forte
FOREL, 1904 (ARDEH 1994) is a junior synonym of T. chefketi FOREL, 1911 (see also CSÖSZ & al. 2007); Tetramorium moravicum KRATOCHVÍL, 1941 (ALIPANAH & al. 1995)
was determined only as Tetramorium sp. now, but it is clearly not T. moravicum.
Results
The proposed preliminary list of Iran comprises 110 ant
species and subspecies from 26 genera of six subfamilies
(Formicinae, Myrmicinae, Ponerinae, Dolichoderinae, Dorylinae and Aenictinae) (Tab. 1). The most speciose subfamily is that of the Myrmicinae (52 species), while Aenictinae and Dorylinae include only one species each. The most
speciose genera in Iran are Camponotus, Cataglyphis and
Messor with 19, 14 and 13 species, respectively. By contrast, only one species per genus was also recorded for
Liometopum, Anochetus, Pachycondyla, and Ponera. Nine
species have been taxonomically described from Iran in earlier studies (indicated by # in Tab. 1).
One subfamily (Dorylinae), two genera and seven species were recorded for Iran for the first time:
Aphaenogaster gibbosa (LATREILLE, 1798): Mazandaran
province, Miankaleh peninsula, Plangan, in shrubland
and grassland habitats, 14.IV.2005, leg. O. Paknia, 3
workers (ZMGU), det. A. Radchenko.
Aphaenogaster kurdica RUZSKY, 1905: Golestan province, Aliabad, near Kabudval waterfall, in forest habitat, 17.VI.2005, leg. O. Paknia, 8 workers (ZMGU), det.
A. Radchenko.
Crematogaster bogojawlenskii RUZSKY, 1905: Khorasan
Razawi province, Sarakhs, 27.IV.2007, leg. H. Hajiqanbar, (IZK), det. A. Radchenko.
Excluded species
As our colleague Dr. Dobovikov, who checked material
recorded by FOREL (1904a) in ZISP, did not find the
voucher specimens of Messor barbarus (LINNAEUS, 1767)
and M. capitatus (LATREILLE, 1798), we propose to exclude both species from the Iranian list, especially as these
species are restricted to the Mediterranean Region.
Included species without vouchers
The vouchers of Camponotus maculatus (FABRICIUS, 1782),
Camponotus thoracicus (FABRICIUS, 1804) and Paratrechina vividula (NYLANDER, 1846) recorded by FOREL
(1904a) were not found in ZISP. However, as these species have been recorded from adjacent countries (COLLINGWOOD & AGOSTI 1996, DONISTHORPE 1950) and also
because the latter one is a tramp species living in the temperate zones in houses and greenhouses, the presence of
these species in Iran appears likely. We included these
species in our list.
The records of Tetramorium caespitum are based on
the old keys and have to be reviewed by use of modern
taxonomic approaches (see SCHLICK-STEINER & al. 2006,
152
106
Chapter 6 - Ant Species List of Iran
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
of Iran
Aenictus dlusskyi ARNOLDI, 1968 *
Anochetus evansi CRAWLEY, 1922 * #
Former name or synonymies,
used in old literature
Determinator
References and material
deposition
N
Radchenko
a, IZK
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
or Munsee
k, HMIM
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 &
ZABELIN, 1985
N
Radchenko
i, HMIM
Camponotus libanicus ANDRÉ, 1881
N
Radchenko
h, HMIM
Camponotus maculatus (FABRICIUS, 1782) *
E
C. maculatus r. cognatus
(SMITH, F. 1858 )
Forel
j
Camponotus oasium FOREL, 1890
SE
C. maculatus r. oasium
Collingwood,
Radchenko
j, c, HMIM
Camponotus oertzeni FOREL, 1889
NW
Radchenko
d, IZK, ZMMU, MIZ
Camponotus sanctus FOREL, 1904
S
Radchenko
d, HMIM, ZMMU
Camponotus thoracicus (FABRICIUS, 1804) *
E
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
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
Radchenko, Taylor
j, p, s, HMIM
Cataglyphis altisquamis (ANDRE, 1881)
N
Radchenko
m, ZMMU
C. maculatus r. dichrous
C. maculatus r. xerxes
Myrmecocystus cursor r. tancrei
153
107
Chapter 6 - Ant Species List of Iran
Cataglyphis bellicosus (KARAVAIEV, 1924) #
N
Cataglyphis bucharicus EMERY, 1925
Myrmecocystus bicolor ssp.
bellicosus
Radchenko
n, IZK, ZMMU
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.
foreli
Radchenko
j, ZMGU
Cataglyphis frigidus persicus (EMERY, 1906) #
S
Myrmecosystus frigidus var.
persica
Emery
O
Cataglyphis lividus (ANDRÉ, 1881)
NE, N, S
Myrmecocystus albicans r.
lividus
Collingwood,
Radchenko
j, h, k, p, HMIM, ZMGU
Cataglyphis niger (ANDRÉ, 1881)
N, S
Myrmecocystus viaticus r.
niger
Collingwood,
Radchenko
j, h, k, p, HMIM, ZMGU
Cataglyphis nigripes ARNOLDI, 1964
W, NW
Radchenko
m, ZMMU, HMIM
Cataglyphis nodus (BRULLÉ, 1833)
N, S
Callingwood,
Radchenko
e, h, k, p, ZMGU
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
Crawley
t
Crematogaster sorokini RUZSKY, 1905
W
Radchenko
i, HMIM
Crematogaster subdentata MAYR, 1877
N
Ward, Collingwood
or Munsee
k, HMIM
Dorylus sp. *
SW
Alipanah
HMIM
Formica cunicularia LATREILLE, 1798
N
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
or Munsee
q, k, HMIM
Lasius brunneus (LATREILLE, 1798)
NW
Crawley
q
Lasius emarginatus (OLIVER, 1792)
NW, N
Seifert
q, ZMGU
Lasius lasioides (EMERY, 1869)
N
Radchenko
p, ZMGU
Lasius neglectus VAN LOON, BOOMSMA &
ANDRASFALVY, 1990 *
N
Schultz, Seifert
p, ZMGU
Lasius platythorax SEIFERT, 1991
SW
Radchenko
i, HMIM
Lasius turcicus SANTSCHI, 1921
N
Collingwood,
Radchenko
h, HMIM
Lepisiota bipartita (SMITH, F., 1861)
E, N
Collingwood
j, HMIM
Lepisiota dolabellae (FOREL, 1911)
N
Ward, Collingwood
or Munsee
k, HMIM
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
Cataglyphis bicolor var. nodus
MENOZZI (1927)
C. scutellaris ssp. schmidti
Formica glauca RUZSKY, 1896
L. emarginatus var. nigroemarginatus
Acantholepis frauenfeldi r.
bipartita
154
108
Chapter 6 - Ant Species List of Iran
Messor denticulatus SANTSCHI, 1927
NE
Radchenko
z, HMIM
Messor ebeninus SANTSCHI, 1927
SW, N
Collingwood
f, HMIM
Messor incorruptus KUZNETSOV-UGAMSKY,
1929
W
Radchenko
i, HMIM
Messor intermedius SANTSCHI, 1927 *
S
Radchenko
e, ZMGU
Messor meridionalis (ANDRÉ, 1883)
SW, N
Ward, Collingwood
or Munsee
k, HMIM
Messor minor (ANDRÉ, 1883)
W
Radchenko
HMIM
Messor rufotestaceus (FÖRSTER, 1850)
S, SW
Radchenko,
Collingwood
c, ZMGU, HMIM
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,
Schultz
f, p, HMIM, ZMGU
Paratrechina vividula (NYLANDER, 1846) *
SE
Prenolepis vividula
Forel
j
Pheidole pallidula (NYLANDER, 1849)
NW
Ph. pallidula subsp. arenarum
var. orientalis
Collingwood
t, f, HMIM
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
Ward, Collingwood
or Munsee
k, HMIM
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
Radchenko, Taylor
t, s, ZISP, ZMGU
Tapinoma karavaievi EMERY, 1925 *
N, S
Radchenko
HMIM, ZMGU
Temnothorax iranicus (RADCHENKO, 1994) #
N
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
Radchenko
HMIM
Messor semirufus var. intermedius
Plagiolepis vindobonensis
LOMNICKI, 1925
T. erraticum subsp. nigerrimum
Leptothorax iranicus
T. moravicum
155
109
Chapter 6 - Ant Species List of Iran
Fig. 1: Map of Iran with the localities where ants were collected (black dots).
STEINER & al. 2006, CSÖSZ & al. 2007). However, based on
distribution maps of Tetramorium ants in the western Palearctic (see SCHLICK-STEINER & al. 2006) we left this species in the list as Tetramorium cf. caespitum. The situation is similar with the old records of Lasius species (see
SEIFERT 1992), that need to be reviewed by modern keys.
Nevertheless this work is beyond the scope of the present
paper, so we included these species in our preliminary list
without revision.
The record of Dorylinae is the first record of this subfamily after the new clarification of this subfamily by BOLTON (2003).
centre of the country that lie in the vast Central Persian
desert basin. There are no species records at all from the
Eastern Iran mountain woodlands, the Kopet Dag woodlands, Kopet Dag semi desert, Azerbaijan shrub desert and
steppe, and the desert and semi-desert areas in the centre
and the east of Iran that comprise more than 50 % of the
area of the country.
Discussion
Although the first reports on Iranian ants were published
more than 100 years ago, the ant fauna of this country remains poorly known. Most of the records are from the
north of Iran, but many of these samples were collected in
disturbed environments near human settlements that comprise only a few percent of the country's surface area. There
are only a few species reports from the extensive natural
habitats of the north, for example from the Caspian deciduous forests. The latter should have a rich ant fauna due to
old geological age of that forest region: it has been covered with forests since the late Tertiary period (ZOHARY
1973). As a result, Tertiarian elements could have survived,
as it is known for the adjacent Talysh and Zuvand districts in south-eastern Azerbaijan (ARNOLDI 1930, 1948).
Among the other regions that have been investigated
only cursorily is the Nubo-Sindian desert and semi desert
ecoregion in the south and the southeast of the country.
These areas are particularly interesting faunistically, as they
are close to the boundaries of the Oriental and Afrotropical zoogeographic regions. Four important ecoregions that
comprise a large part of the Irano-Anatolian biodiversity
Distribution of records
Regarding the geographical distribution of the species records that had been examined in this study, we found a
mismatching between the area of the locations that had
been studied and the respective number of species that
had been found there: e.g., 34 species of a total of 109 species were collected in natural or disturbed habitats of Tehran province, though the territory of this province is less
than 2 % of the territory of the country. Fifteen species
were recorded from the Caspian forest region in the north
of Iran that covers only 4 % of the country. Eight species
were found in Zagros Mountains forest-steppe ecoregion
(about 20 % of entire area of the country). From the south
of Iran altogether 34 species were reported. In contrast,
there are only a few records from the wide Elburz Range
forest-steppe and the Eastern Anatoloian Mountains in the
north and northwest of Iran, or from eastern parts and the
156
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Chapter 6 - Ant Species List of Iran
hotspot have not been studied at all. These are the Eastern
Anatolian Mountains, Elburz forest-steppe, Kopet Dag
woodlands and forest-steppe, and Zagros Mountains foreststeppe. For comparison, the well studied myrmecofauna of
the Turkmenistan's part of the Kopet Dag is one of the
richest local ant fauna in Central Asia (DLUSSKY & ZABELIN 1985, DLUSSKY & al. 1990).
This scarcity of data, especially from the border regions, does not permit to make a proper zoogeographical
analysis of the Iranian ant fauna at present. However, in
the future, when sufficient material will have been sampled, the Iranian myrmecofauna needs to be compared with
those of the adjacent regions, e.g., Turkey, Armenia, Azerbaijan, Turkmenistan, Afghanistan, and Arabian Peninsula. Similarly, the presence of North African elements in
Iran (e.g., Camponotus fellah DALLA TORRE, 1893 or Crematogaster inermis MAYR, 1862) demonstrates a relation of
the Iranian and North African desert faunas that has to be
confirmed by more intensive sampling.
The Iranian ant fauna includes eight "tramp" species,
which have been introduced by humans to many countries
and in some cases have gained a worldwide distribution
(see also MCGLYNN 1999, PAKNIA 2006, PAKNIA & KAMI
2007): Lasius neglectus VAN LOON, BOOMSMA & ANDRASFALVY, 1990, Monomorium pharaonis (LINNAEUS,
1758), M. destructor (JERDON, 1851), Pheidole teneriffana
FOREL, 1893, Paratrechina longicornis (LATREILLE, 1802),
P. flavipes (SMITH, F., 1874), P. vividula (NYLANDER, 1846)
and Pachycondyla sennaarensis (MAYR, 1862).
Almost half of the recorded species in the Iranian
checklist belong to the genera Camponotus, Messor and
Cataglyphis. Similar faunistic patterns are found in other
arid Asian regions, e.g., Turkmenistan (DLUSSKY & al.
1990) and Saudi Arabia (COLLINGWOOD 1985, COLLINGWOOD & AGOSTI 1996). The main reason for the high diversity of these genera are the environmental conditions in
Iran that comprise mainly arid and semi arid areas, the preferred habitats of Messor, Cataglyphis and many Camponotus species (DLUSSKY 1981, DLUSSKY & al. 1990, HÖLLDOBLER & WILSON 1990, ANDERSEN & CLAY 1996, ANDERSEN & SPAIN 1996).
The second reason for the dominance of those three genera in ant collections may be artificial: their members are
large and can be easily collected by anyone. In the majority of the former studies, "direct hand collecting" was
the main method. For this reason small-sized and cryptic
ants (e.g., Leptanillinae, Amblyoponinae, Dacetini, etc.) are
under-represented in the investigated material. Furthermore,
social parasites are also missing in the presented species
list of Iran. To overcome this sampling bias and to establish
a reliable species list, thorough investigations of ant diversity in all parts of Iran are urgently needed. They should be
conducted by standard sampling methods, like direct collecting from ant nests, pitfall traps, bait trapping and litter
extraction with Winkler collectors (see AGOSTI & ALONSO
2000).
At last, we have to emphasize that the Iranian ant fauna
seems to be one of typically Palaearctic character. If we
exclude the introduced species mentioned above, native
members of only four tropical (Oriental or Afrotropical)
genera are found in Iran: Aenictus dlusskyi ARNOLDI, 1968,
Anochetus evansi CRAWLEY, 1922, Dorylus sp., and Polyrhachis lacteipennis SMITH, F., 1858. This dominance of
Palaearctic ant genera will probably persist in a more comprehensive species list to be presented in the future.
Acknowledgements
We thank Hamidreza Hajiqanbar for collecting and sending the material of two ant species, Crematogaster bogojawlenskii and Tetramorium inerme that were new to Iran.
We thank Dr. Bernhard Seifert for his taxonomic comments,
Dr. Cedric A. Collingwood for the identification of many
specimens and Dr. Dubovikov for checking Forel's vouchers in ZISP. We are indebted to John Fellowes, an anonymous referee and the editors for their useful suggestions towards the improvement of our paper. We thank
Doug Johns and Bryson Voirin for language correction of
the first and the second versions of the text, respectively.
Zusammenfassung
Nach einer umfassenden Durchsicht der Literatur und der
Untersuchung von Material aus eigenen Aufsammlungen
sowie aus Museen und Forschungsinstituten verschiedener europäischer Länder, präsentieren wir die erste Artenliste der Ameisen des Iran: 110 Arten aus 26 Gattungen und
sechs Unterfamilien der Formicidae (Formicinae, Myrmicinae, Ponerinae, Dolichoderinae, Dorylinae und Aenictinae) wurden bislang gefunden. Die meisten der hier gelisteten Arten wurden im Norden des Landes gesammelt,
zumeist in anthropogen beeinflussten Gebieten in der Nähe
von Siedlungen. Eine Unterfamilie (Dorylinae), zwei Gattungen (Ponera LATREILLE, 1804 und Dorylus FABRICIUS,
1793), sowie sieben Arten der Formicidae (Aphaenogaster
gibbosa (LATREILLE, 1798), A. kurdica RUZSKY, 1905,
Messor minor (ANDRE, 1883), Tapinoma karavaievi EMERY, 1925, Temnothorax parvulus (SCHENCK, 1852), Tetramorium inerme MAYR, 1877, und Crematogaster bogojawlenskii RUZSKY, 1905) wurden erstmalig für den Iran registriert. Die artenreichsten Gattungen des Iran sind Camponotus MAYR, 1861 mit 19 Arten sowie Cataglyphis FOERSTER, 1850 mit 14 und Messor FOREL, 1890 mit 13 Arten.
Zoogeographisch gesehen dominieren paläarktische Elemente im Iran, allerdings wurden auch verschiedene Arten
der Orientalis und Afrotropis gefunden. Viele Landesteile
wurden bislang kaum oder gar nicht untersucht und der
Artenreichtum der Formicidae des gesamten Iran dürfte
wesentlich größer sein.
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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
1
University of Ulm, Institute for Experimental Ecology, Albert Einstein Allee 11, 89069 Ulm,
Germany, [email protected] and [email protected]
2
Museum 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 probably 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
Iran is located in the mid-latitude band of arid
and semi-arid regions of the Old World, in
Southwest Asia. Western and especially
southwestern Asia comprises vast arid and semiarid areas and represents a number of distinctive
ecoregions that support numerous endemic plants
and animals (Zohary 1973; Olson et al. 2001).
Biogeographically, this territory represents a
transition zone between three regions: Palaearctic,
Afrotropical and Oriental. Many of these unique
ecoregions have not yet been studied intensively.
Despite more than one hundred years of
myrmecology and the publication of a considerable
number of papers on the ant fauna of this region,
comprehensive faunistic investigations on ants have
not been provided. Additionally, many genera of
this region, such as Messor, Cataglyphis,
Camponotus and Monomorium, contain numerous
taxa of ambiguous status and need modern revision.
114
The Iranian ant fauna has been poorly investigated.
So far, 110 species belonging to 26 genera have
been recorded from Iran (Paknia et al. 2008). In
this paper, we introduce 28 species that are new
for the Iranian fauna. This large number of new
records, a short time after publishing a primary
checklist for the country (Paknia et al. 2008) shows
the incompleteness of our myrmecological
knowledge of this region.
MATERIALS AND METHODS
The majority of ant samples were collected in two
periods of extensive field work in spring and
summer, of 2007 and 2008, respectively. The first
field study was carried out along a transect from
the north to the south of Iran through arid and semiarid areas in four steppe and desert ecoregions.
The second block of fieldwork was conducted
inside the Central Persian desert basins in the
centre of Iran and in the Caspian Hyrcanian mixed
Chapter 7 - New Records of Ants from Iran
30
New records of ants (Hymenoptera, Formicidae) from Iran
forests in the north of Iran. Locations of the sample
sites are indicated in Fig. 1. In the first year of
sampling, we employed pitfall trapping and bait
trapping for sampling ants, while in the second
year, we used only pitfall trapping for the
collection of ants in arid and semi-arid areas. Both
methods are suitable for rapid survey of open
habitats, and well represent epigaeic ant species
(Agosti & Alonso 2000). In forest areas, pitfall
trapping and Winkler collectors were applied.
Additionally, nest samples were collected during
both years of sampling. Moreover, a small number
of the samples were collected by hand during
different expeditions and in a few cases by Iranian
students of biology.
Samples were preserved in 96% alcohol,
brought to the laboratory, and cleaned of debris
and soil. In total, 35,450 specimens were checked
and sorted to morphospecies. Most of the
morphospecies were identified to species level,
but some remained undetermined and were
tentatively distinguished by species codes such
as “sp. ir-abpari-01”.
Keys for adjacent regions and taxonomic
revisions of different ant genera used in the present
study were as follows: Arnoldi (1948, 1977b:
Central Asia), Dlussky et al. (1990: Central Asia),
Radchenko (1992a, 1992b, 1994a, 1994b, 1995a,
1997a, 1997b: Central Asia), Arakelyan (1994:
Central Asia), Collingwood and Agosti (1996:
Arabian Peninsula), and Bolton (2000: Dacetini
ants). The following ant collections were also used:
Museum and Institute of Zoology of Polish
Academy of Sciences, Warsaw, Poland (MIZ);
Zoological Museum of Moscow State University,
Russia (ZMMU); Zoological Institute of Russian
Academy of Sciences, St. Petersburg, Russia
(ZISP); and Institute of Zoology of Ukrainian
NationalAcademy of Sciences, Kiev, Ukraine (IZK).
In this paper, we define the following
ambiguous geographical terms as follows:
Anatolia by its borders in Turkey, based on
historical records from Asia Minor (all from within
the Asian part of modern-day Turkey); the Arabian
Peninsula as Kuwait, Oman, Qatar, Saudi Arabia,
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, 18Tabas, 19- Dena, 20- Kerman and 21- Mond.
115
Chapter 7 - New Records of Ants from Iran
Omid Paknia, Alexander Radchenko, and Martin Pfeiffer
“Caucasus” as including Great Caucasus (i.e.,
Russian North Caucasian republics from the Black
sea to Caspian sea) as well as Armenia, Azerbaijan
and Georgia, while “Transcaucasus” covers only
the territory of Armenia, Azerbaijan and Georgia;
“Central Asia” as including Kyrgysztan, Tajikistan,
Turkmenistan and Uzbekistan (i.e., excluding
Kazakhstan); and the “Middle East” as consisting
of Israel, Jordan, Lebanon, Palestine and Syria.
The territory of Iraq is considered separately.
All determined and undetermined species
were deposited in the Antbase.net Collection
(ABNC) managed by M. Pfeiffer and in the private
collection of O. Paknia.
RESULTS
Additional to the earlier published checklist
(Paknia et al. 2008), we found 32 new ant species
for Iran. Twenty-eight were identified to species
level and the other four were identified to genus
level and given species codes. Our collecting efforts
also resulted in the addition of six ant genera new
to Iran: Dolichoderus, Myrmecina, Proformica,
Pyramica, Stenamma, and Strongylognathus. The
new records are listed below.
Camponotus fallax (Nylander, 1856)
Material: 4 , Caspian Hyrcanian mixed forest
ecoregion, Gorgan, urban area (~36°50´23´´N,
54°26´19´´E), ~142 m asl, 1.IV.2006, leg. Omid Paknia.
Remarks: the species is distributed in Europe
(northwards to southern Sweden), the Caucasus,
Anatolia and northwestern Kazakhstan, and also
reported from the southern part of western
Siberia. Iran is at the southeasternmost edge of
the known distribution of this species, that
inhabits mainly light and dry deciduous and
mixed forests, and often occurs in old parks and
orchards. It nests in dead parts of living trees or
in wooden constructions (Radchenko 1997c;
Czechowski et al. 2002).
Camponotus interjectus Mayr, 1877
31
Khoshyelagh Wildlife Refuge (36°47´41´´N,
55°28´13´´E), 1534 m asl, 3.VI.2007, leg. Omid Paknia.
Remarks: C. interjectus is distributed in Central
Asia and Afghanistan (Dlussky et al. 1990;
Radchenko 1997c) and recorded also from Iraq
(Wheeler & Mann 1916) and Dagestan in Russia
(Kuznetsov-Ugamsky 1929). It inhabits mainly
mountain steppes; nests are built in soil, often
under stones.
Camponotus kurdistanicus Emery, 1898
Material: 13 , transitional region between Alborz
Range forest steppe and Central Persian deserts,
Khojir National Park (35°38´54´´N, 51°43´44´´E),
1485 m asl, 1.V.2008, leg. Omid Paknia; 2 ,
Zagros Mountains forest steppe, Kurdistan, Bane
(~35°59´N, 45°53´E), ~1557 m asl., summer 2004,
leg. Shahin Mostafai.
Remarks: This species was known from Anatolia,
Iraq and Azerbaijan (Emery 1898, 1925; Pisarski
1971; Radchenko 1997c).
Camponotus libanicus Andre, 1881
Material: 8 , Alborz Range forest steppe,
Khoshyelagh Wildlife Refuge (36°48´55´´N,
55°31´54´´E), 1701 m asl, 8.VI.2007, leg. Omid Paknia.
Remarks: C. libanicus is distributed in the Middle
East countries and Anatolia (Radchenko 1997c).
This species inhabits dry and semi-dry areas. Nests
are built in soil.
Camponotus shaqualavensis Pisarski, 1971
Material: 4 , Nubo Sindian deserts, Kerman, urban
area (~30°16´N, 57°04´E), ~1760 m asl, 26.V.2004,
leg. Shiva Sadeghirad.
Remarks: C. shaqualavensis was described from Iraq
(Pisarski 1971) and additionally was recorded from
Turkey (Aktaç 1977; see also Radchenko 1997b).
Camponotus staryi Pisarski, 1971
Material: 18 , Alborz Range forest steppe,
Golestan National Park (37°20´56´´N, 56°14´46´´E),
1256 m asl, 30.V.2007; 9 ,Alborz Range forest steppe,
116
Material: 23 , Zagros Mountains forest steppe,
Dena Protected area (30°54´N, 51°25´E), 2683 m
asl, 6.VII.2007, leg. Omid Paknia.
Chapter 7 - New Records of Ants from Iran
32
New records of ants (Hymenoptera, Formicidae) from Iran
Remarks: C. staryi was described from Iraq
(Pisarski 1971). This is the first record of this
species outside its type locality.
Remarks: C. kurdistanicus was described from Iraq
(Pisarski 1965) and recorded additionally from
Anatolia (Aktaç 1977; Radchenko 1997a).
Cataglyphis bergianus Arnoldi, 1964
Dolichoderus quadripunctatus (Linnaeus, 1771)
Material: 5 , Central Persian deserts, Tabas
(33°36´53´´N, 57°05´36´´E), 1083 m asl, 16.V.2008,
leg. Omid Paknia.
Material: 1 , Caspian Hyrcanian mixed forests,
Talysh (37°42´19´´N, 48°53´14´´E), 66 m asl,
6.VII.2008, leg. Omid Paknia.
Remarks: C. bergianus was known from Central
Asia, southern Kazakhstan and Afghanistan
(Pisarski 1967, 1970; Radchenko 1997a).
Remarks: The genus Dolichoderus Lund comprises
more than 130 species, distributed in all
zoogeographic realms except Africa and
Madagascar. Only one species, D. quadripunctatus,
is known from the western Palaearctic. Its range
covers central and southern Europe, central and
southern parts of eastern Europe, the Caucasus,
Anatolia, southwest Siberia, and the Tien-Shan range
in Central Asia and China. This genus is new to Iran.
Cataglyphis cinnamomeus (Karavaiev, 1910)
Material: 3 , Central Persian deserts, Siahkooh
National Park (32°35´57´´N, 54°13´56´´E), 995 m
asl, 25.V.2008, leg. Omid Paknia; 4 , Central
Persian deserts, Turan National Park (35°58´N,
56°04´E), 1191 m asl, 16.VI.2007, leg. Omid Paknia;
6 , Central Persian deserts, South of Naeen
(32°43´11´´N, 53°16´42´´E), 1372 m asl, 27.V.2008,
leg. Omid Paknia.
Remarks: Outside of Iran, C. cinnamomeus was
recorded from Central Asia, southern Kazakhstan
(Karavaiev 1910) and Afghanistan (Pisarski 1967;
Radchenko 1997a). This species is the most
thermophilous among Central Asian Cataglyphis,
inhabiting mainly stony and clayey deserts. In
mountain areas, it lives in dry parts of river valleys.
Cataglyphis cugiai Menozzi, 1939
Material: 18 ,Alborz Range forest steppe, Golestan
National Park (37°20´57´´N, 56°14´44´´E), 1258
m asl, 30.V.2007, leg. Omid Paknia.
Messor sp. (cf. M. oertzeni Forel, 1910)
Material: 3 , Zagros Mountains forest steppe,
Kurdistan, Saggez (~36°14´N, 46°15´E), ~1481
m asl, 2004, leg. Shahin Mostafai; 4 , Tehran,
urban area (~35°41´N, 51°25´E), ~1162 m asl,
16.VII.2005, leg. Nasim Vakhideh; 5 , Mashad,
urban area (~36°17´N, 59°35´E), ~985 m asl, leg.
Nayereh Ghafarian.
Remarks: M. oertzeni is known from the Balkans
and Anatolia (Agosti & Collingwood 1987;
Atanassov & Dlussky 2002). It belongs to the
structor species-group, but reliable determination
of many species from this group is impossible
before a taxonomic revision is provided.
Messor excursionis Ruzsky, 1905
Remarks: This species was known only from the
northwest of India, Karakorum region (Menozzi
1939; Radchenko 1997a).
Material: 21 , Central Persian deserts, Kavir
National Park (34°45´14´´N, 52°10´07´´E), 1092 m
asl, 7.VI.2008, leg. Omid Paknia.
Cataglyphis kurdistanicus Pisarski, 1965
Remarks: M. excursionis was recorded from
Central Asia, Afghanistan and Mongolia
(Karavaiev 1910; Stärcke 1935; Pisarski 1967;
Arnoldi 1970, 1977a; Dlussky et al. 1990; Pfeiffer
et al. 2006).
Material: 54 , Zagros Mountains forest steppe,
Dena Protected Area (30°54´45´´N, 51°24´39´´E),
2695 m asl, 7.VII.2007, leg. Omid Paknia; 4 , Zagros
Mountains forest steppe, Kurdistan, Bane
(~35°59´N, 45°53´E), ~1557 m asl, summer 2004,
leg. Shahin Mostafai.
117
Chapter 7 - New Records of Ants from Iran
Omid Paknia, Alexander Radchenko, and Martin Pfeiffer
Messor perantennatus Arnoldi, 1970
Material: 9 , Alborz Range forest steppe,
Khoshyelagh Wildlife Refuge (36°47´43´´N,
55°28´12´´E), 1536 m asl, 4.VI.2007, leg.
Omid Paknia.
Remarks: This species was known only from
Turkmenistan (Arnoldi 1970; Dlussky et al. 1990).
Messor subgracilinodis Arnoldi, 1970
Material: 8 , Central Persian deserts, Turan
National Park (35°57´48´´N, 56°05´35´´E), 1131
m asl, 12.VI.2007, leg. Omid Paknia; 7 , Central
Persian deserts, Tabas (33°36´44´´N, 57°05´45´´E),
1083 m asl, 16.V.2008, leg. Omid Paknia; 3 ,
Central Persian deserts, Siahkooh National Park
(32°35´55´´N, 54°13´59´´E), 996 m asl, 25.V.2008,
leg. Omid Paknia; 5 , Central Persian deserts,
Robat Posht Badam (32°57´26´´N, 55°32´08´´E)
1401 m asl, 20.V.2008, leg. Omid Paknia.
Remarks: M. subgracilinodis was recorded from
Turkmenistan (Arnoldi 1970, 1977b; Dlussky et
al. 1990).
Messor turcmenochorassanicus Arnoldi, 1977
Material: 65 , Zagros Mountains forest steppe,
Dena Protected Area (30°54´45´´N, 51°24´39´´E),
2627 m asl, 6.VII.2007, leg. Omid Paknia; 7 ,
Caspian Hyrcanian mixed forests Nur, urban area
(~36°33´N, 51°52´E), ~3 m asl, 17.III.2005, leg.
Masud Tavakoli.
Remarks: Messor turcmenochorassanicus was
recorded only from the Kopetdag Mountains in
Turkmenistan (Arnoldi 1977b; Dlussky et al. 1990).
Messor variabilis Kuznetsov-Ugamsky, 1927
Material: 4 , Alborz Range forest steppe,
Golestan National Park (37°20´57´´N,
56°14´44´´E), 1254 m asl, 28.V.2007, leg. Omid
Paknia; 6 , Central Persian deserts, Kavir
National Park (34°45´47´´N, 52°10´24´´E), 1051
m asl, 23.VI.2007, leg. Omid Paknia.
118
33
Remarks: M. variabilis is distributed in plains and
foothills of Central Asia, where it inhabits different
kinds of deserts (Arnoldi 1970, 1977b; Dlussky et
al. 1990).
Monomorium barbatulum Mayr, 1877
Material: 3 , Central Persian deserts, Robat Posht
Badam (32°57´22´´N, 55°32´09´´E), 1406 m asl,
17.V.2008, leg. Omid Paknia; 6 , Central Persian
deserts, Tabas (33°36´57´´N, 57°05´35´´E), 1074
m asl, 15.V.2008, leg. Omid Paknia.
Remarks: M. barbatulum has a wide distribution
from the southeast of Europe (Astrakhan Province
of Russia) to Central Asia and Afghanistan
(Pisarski 1967; Tarbinsky 1976; Dlussky et al.
1990) and the Arabian Peninsula (Collingwood &
Agosti 1996).
Monomorium dentigerum (Roger, 1862)
Material: 13 , Nobu Sindian deserts, Mond
Protected Area (28°07´08´´N, 51°23´50´´E), 9 m
asl, 14.VII.2007, leg. Omid Paknia.
Remarks: M. dentigerum can be found all over
the Middle East (Radchenko 1997d) and the
Arabian Peninsula including Yemen (Collingwood
& Agosti 1996).
Monomorium perplexum Radchenko, 1997
Material: 24 , transitional region of Alborz Range
forest steppe and Central Persian deserts, Khojir
National Park (35°39´11´´N, 51°43´37´´E), 1462
m asl, 1.V.2008, leg. Omid Paknia.
Remarks: Outside of Iran, the species is distributed
in the Transcaucasus, Anatolia, the Aegean Islands
and Greece (Radchenko 1997d).
Monomorium ruzskyi Dlussky et Zabelin, 1985
Material: 3 , transitional region of Alborz Range
forest steppe and Central Persian deserts, Khojir
National Park (35°38´56´´N, 51°43´45´´E), 1466 m
asl, 3.V.2008, leg. Omid Paknia.
Chapter 7 - New Records of Ants from Iran
34
New records of ants (Hymenoptera, Formicidae) from Iran
Remarks: M. ruzskyi was recorded from the
Transcaucasus, Turkmenistan and Uzbekistan
(Dlussky & Zabelin 1985; Arakelyan 1994).
Siberia and Kazakhstan to Tuva in Siberia (Dlussky
1969), the Central Asian Mountains (Dlussky et
al. 1990). This genus is new to Iran.
Myrmecina sp. ir-abpari-01 and ir-ghaemshahr-01
Pyramica sp. ir-golestan-01 and ir-ghaemshahr01 (argiola-group)
Material: 8 , Caspian Hyrcanian mixed forests,
Ghaemshahr (36°22´16´´N, 52°50´53´´E), 155 m
asl, 4.VI.2008, leg. Omid Paknia; 4 , Caspian
Hyrcanian mixed forests, Abpari (36°30´05´´N,
51°55´58´´E), 308 m asl, 22.VI.2008, leg. Omid
Paknia.
Remarks: The genus Myrmecina Curtis consists
of about 40 described species distributed in the
Holarctic, South and Southeast Asia, Australia and
South America. It is absent in the Afrotropical and
Malagasy Regions. The highest speciosity occurs
in the Oriental Region. In the western Palaearctic,
there are four known species distributed from the
Iberian Peninsula and Algeria to the Transcaucasus
and the Middle East. This genus is new to Iran.
Myrmica specioides Bondroit, 1918
Material: 8 , Caspian Hyrcanian mixed forests,
Babolsar, in urban area (36°42´30´´N, 52°38´04´´E),
10 m asl, 18.IX.2002, leg. Omid Paknia.
Remarks: This species has a very wide distribution
that includes Europe (northwards to the south of
England and Denmark), the Caucasus, Anatolia,
Turkmenistan, southwest Siberia and northern
Kazakhstan, eastwards to the Altai Mountains
(Radchenko & Elmes 2004; Radchenko 1994b ); it
was introduced to North America (Jansen &
Radchenko 2009).
Proformica epinotalis Kusnetsov-Ugamsky,
1927
Material: 19 ,Alborz Range forest steppe, Golestan
National Park (37°20´54´´N, 56°14´42´´E), 1261
m asl, 29.V.2007, leg. Omid Paknia; 11 ,Alborz Range
forest steppe, Khoshyelagh Wildlife Refuge
(36°46´24´´N, 55°22´42´´E), 1707 m asl, 6.VI.2007, leg.
Omid Paknia.
Remarks: This species is distributed from Romania
in the west through southeast Europe, southwest
Material: 1 , Caspian Hyrcanian mixed forests,
Ghaemshahr (36°22´16´´N, 52°50´53´´E), 155 m
asl., 4.VI.2008, leg. Omid Paknia. ; 1 , Caspian
Hyrcanian mixed forests, Golestan National Park
(37°24´04´´N, 55°48´04´´E), 520 m asl,
11.VI.2008, leg. Omid Paknia.
Remarks: In the last ten years, the systematics of
the tribe Dacetini, to which Pyramica Roger
belongs, was cardinally changed. First of all,
Bolton (1999) revived the name Pyramica from
synonymy and proposed to consider it as a senior
synonym of more than 20 generic names. A year
later, he published a huge taxonomic revision of
this tribe, describing several hundred new species
(Bolton 2000). As a result, Pyramica is now one
of the world’s biggest ant genera that includes
more than 300 species; its range encompasses the
whole world, with the overwhelming majority of
species distributed in the tropics. This genus is
new to Iran.
Several Pyramica species were recorded
from the Mediterranean region of Europe and Africa,
the Middle East, Anatolia and the Transcaucasus,
while it is not yet known from Central Asia (e.g.,
Arakelyan & Dlussky 1991;Arakelyan 1994; Bolton
et al. 2006; Radchenko 2007).
It is necessary to note that in the latest
revision of the tribe Dacetini (Baroni Urbani &
De Andrade 2007), the name Pyramica is
considered a junior synonym of Strumigenys, but
in this paper, we retain Pyramica until there is a
definitive opinion on this question.
Stenamma sp. ir-talysh-01
Material: 1 , Caspian Hyrcanian mixed forests,
Talysh (37°40´44´´N, 48°48´27´´E), 869 m asl,
7.VII.2008, leg. Omid Paknia.
Remarks: Stenamma is a small genus, comprising
about 50 species distributed mainly in the
Holarctic Region, but some of which penetrated
119
Chapter 7 - New Records of Ants from Iran
Omid Paknia, Alexander Radchenko, and Martin Pfeiffer
into Central America, India and Pakistan
(Himalayan region), and Southeast Asia. The
highest species richness of this genus occurs in
the rather warm and humid deciduous Holarctic
forests, many of which can be considered relict
habitats. Stenamma is also quite speciose in
Caucasian and Anatolia forests (Arnoldi 1975;
DuBois 1998). This genus is new to Iran.
Strongylognathus sp. ir-astaneh-01
Material: 3 , Caspian Hyrcanian mixed forests,
Astaneh, rural area (~37°15´N, 49°56´E), ~0 m asl,
summer 2004, leg. Faeze Mohammaddoost.
Remarks: The genus Strongylognathus is new to
Iran. It is one of few genera endemic to the
Palaearctic Region. It comprises about 25 species,
centred in the western Palaearctic (the range being
Europe, northwest Africa, Anatolia, the Middle
East, Kazakhstan, Central Asia and southwest
Siberia), and only three species are known from
the eastern Palaearctic (China, Korea and Japan)
(e.g., Radchenko 1985, 1991; Radchenko 1995b;
Wei et al. 2001; Japanese Ant Database Group
2003; Radchenko 2005). All Strongylognathus
species are permanent social parasites in nests of
the Tetramorium Mayr species.
35
Tetramorium armatum Santschi, 1927
Material: 14 ,Alborz Range forest steppe, Golestan
National Park (37°20´36´´N, 56°14´57´´E), 1226 m asl,
28.V.2007, leg. Omid Paknia; 7 , Central Persian
deserts, Kavir National Park (34°45´47´´N,
52°10´24´´E), 1045 m asl, 21.VI.2007, leg. Omid
Paknia; 6 , Central Persian deserts, Turan National
Park (35°58´22´´N, 56°04´43´´E), 1174 m asl,
15.VI.2007, leg. Omid Paknia.
Remarks: T. armatum is morphologically close to
T. inerme Mayr, but in contrast to the latter, which
lives in the Central Asian plains, it has been found
predominantly in mountains, of Central Asia, the
Transcaucasus, Afghanistan and Mongolia
(Dlussky et al. 1990; Radchenko 1992b).
Tetramorium schneideri Emery, 1898
Material: 21 ,Alborz Range forest steppe, Golestan
National Park (37°20´36´´N, 56°14´57´´E), 1226
m asl, 28.V.2007, leg. Omid Paknia.
Remarks: T. schneideri was recorded in the plains
and foothills of Central Asia and Afghanistan
(Dlussky et al. 1990; Radchenko 1992b).
Tetramorium striativentre Mayr, 1877
Temnothorax anodonta (Arnoldi, 1977)
Material: 5 , Caspian Hyrcanian mixed forests,
Golestan National Park (37°23´53´´N, 55°48´01´´E),
500 m asl, 11.VI.2008, leg. Omid Paknia.
Remarks: Outside Iran, T. anodonta was recorded
from Armenia (Arnoldi 1977a; Arakelyan 1994;
Radchenko 1995a) .
Temnothorax nadigi (Kutter, 1925)
Material: 3 , Caspian Hyrcanian mixed forests,
Abpari (36°30´07´´N, 51°55´58´´E), 315 m asl,
28.VI.2008, leg. Omid Paknia.
Remarks: Known from southern Europe, Anatolia,
the Transcaucasus and Turkmenistan (including
its junior synonyms Leptothorax caucasicus
Arnoldi and L. hasardaghi Dlussky (Radchenko
1995a; Czechowski et al. 2002).
120
Material: 16 , Central Persian deserts, Daranjir
Protected Area (32°26´18´´N, 55°01´21´´E), 1227
m asl, 19.V.2008, leg. Omid Paknia; 2 , Central
Persian deserts, Siahkooh National Park
(32°35´55´´N, 54°13´59´´E), 998 m asl, 25.V.2008,
leg. Omid Paknia; 26 , Central Persian deserts
(32°57´23´´N, 55°32´08´´E), 1392 m asl,
17.V.2008, leg. Omid Paknia; 4 , Central Persian
deserts, Tabas (33°36´44´´N, 57°05´46´´E), 1083
m asl, 14.V.2008, leg. Omid Paknia; 22 , Central
Persian deserts, South of Naeen (32°43´11´´N,
53°16´39´´E), 1372 m, asl, 26.V.2008, leg. Omid
Paknia; 23 , transitional region of Alborz Range
forest steppe and Central Persian deserts, Khojir
National Park (35°38´56´´N, 51°43´45´´E), 1466
m asl, 29.IV.2008, leg. Omid Paknia; 32 , Central
Persian deserts, Kavir National Park (34°45´40´´N,
52°10´19´´E), 1057 m asl, 7.V.2008, leg. Omid Paknia;
39 , Central Persian deserts, Turan National Park
(35°58´22´´N, 56°04´43´´E), 1176 m asl, 17.VI.2007,
leg. Omid Paknia.
Chapter 7 - New Records of Ants from Iran
36
New records of ants (Hymenoptera, Formicidae) from Iran
Remarks: Inhabits mainly mountains of Central Asia
and Afghanistan, whereas the morphologically
similar T. schneideri lives in plains (Dlussky et al.
1990; Radchenko 1992b).
DISCUSSION
Iran’s ant species list is far from complete. Iran
is a vast country with a total area of 1.6 million
square kilometres. Although arid and semi-arid
areas cover nearly half of the country, Iran also
includes high mountains with alpine areas
(Noroozi et al. 2008), broadleaf forest in the
southern coastal plains of the Caspian forests,
and steppe forests in the north and west (Zohary
1973). Thus, the 138 recorded species for the ant
fauna of Iran is still far from the true number.
Additionally, our sampling sites were limited to
the north and the centre of Iran and no samples
were collected from the east, the southeast or the
southwest of the country (Fig. 1). As a result, it
is not yet possible to draw complete and correct
biogeographic conclusions about the overall ant
fauna of Iran. Nevertheless, the recorded species
give clues about how the ant fauna is
biogeographically structured in the areas we have
substantially sampled, like the north and centre
of Iran.
Many species on this list were recorded
from the north and centre of the Central Persian
desert basins (Fig. 1), where the highest sampling
effort was applied. The deserts and semi-deserts
of Iran represent a transitional zone between the
hot and cold deserts of Asia (Breckle 2002). The
Central Persian desert basins, except in the
southeast of the region, are characterised by cold
winters and an annual average occurrence of 60
frost days, and thus can be likened to the belt of
Central Asian deserts. Our new records from the
Central Persian desert basins indicate that the ant
fauna of this region has more species in common
with that of the Central Asian deserts (e.g., C.
bergianus, M. subgracilinodis, M. variabilis and
M. ruzsky) than with those of the hot subtropical
deserts of the Middle East and Arabian Peninsula.
The Caspian Hyrcanian mixed forests are
geologically old and preserve the last remnants of
primary temperate deciduous broadleaved forests
worldwide, existing there since the late Tertiary
period (Zohary 1973; Ramezani et al. 2008).
Apart from Proformica, all the newly
recorded genera (Dolichoderus, Myrmecina,
Pyramica, Stenamma and Strongylognathus)
were collected from this region. As the natural
history (fauna and flora) of the Caspian Hyrcanian
mixed forests is close to those of the Caucasian
and Anatolian regions, the ant fauna of the region
is most likely related to them. Interestingly,
specimens of three ant genera, Dolichoderus,
Myrmecina and Pyramica were extracted by
Winkler collector from the leaf litter of Caspian
Hyrcanian mixed forests. This finding requires us
to make further efforts to explore the litter layer
in such forest ecosystems. Future intensive studies
in the Caspian Hyrcanian area, and in the east,
southeast and southwest of the country, are most
likely to reveal new records or even new species
for science.
In closing, we emphasise the need for
fundamental studies in this part of Asia and invite
all myrmecologists, especially those from Asia,
to take part in this job.
ACKNOWLEDGEMENTS
We thank all students named in this paper for the
contribution of specimens. Also, we are grateful to
the staff of the Department of Environment of Iran
for the assistance and help that allowed this study
to be conducted. We also wish to thank Dr
Yoshimura, Dr Eguchi and one anonymous reviewer
who made very valuable and helpful suggestions.
We are indebted to John Fellowes for his useful
suggestions and for language correction of the text.
Financial support for the PhD study of O. Paknia in
Germany, by the German Academic Exchange
Service (DAAD), is gratefully acknowledged.
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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 speciesgroup 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.
18 species of the genus Cataglyphis have been
recorded for Iran till recently (Paknia et al. 2008,
2009), but in the newly collected material by one of the
co-authors (O. Paknia) we found specimens that belong
to two new species, which are described below. One of
them belongs to the bicolor species-group and well differs from all members of this group by its totally yellow
colour, except of C. lunaticus Baroni Urbani described from Turkey. The second new species most probably
is a member of the cursor species-group and has
unique characteristics in its dense, rather long and
coarse, silverish pubescence of the body.
INTRODUCTION
Cataglyphis is one of the keystone ant genera in
arid zones of the Old World. It is distributed mainly in
Palaearctic, while several species dwell in deserts and
semi-deserts of Afrotropical and Oriental Regions (India and Pakistan). More than 100 species are known in
this genus till now, and even 2 social parasites were
described (Agosti 1994, Radchenko 1997b, Bolton et al.
2007).
Members of this genus are large (up to 13 mm) ants,
and all of them inhabit open dry habitats (steppes,
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
several subgenera (e.g., see Bolton 1995), but more
recently subgeneric division was refused and the genus
was separated into several species groups and species
complexes within them (Agosti 1990, Radchenko 1997a).
MATERIAL
AND METHODS
Material was collected from arid areas of the Central and Southern Iran (Fig. 1) during two field trips in
spring and summer 2007 and 2008.
PL ISSN 0003-4541 © Fundacja Natura optima dux
doi: 10.3161/000345410X499533
124
Chapter 8 - Two New Species of Cataglyphis
70
A. RADCHENKO and O. PAKNIA
from the lower margin of eye to the articulation with mandible,
AL – diagonal length of alitrunk (seen in profile),
measured from the anterior end of propodeum
to the posterior margin of propodeal lobes,
PnW – maximum width of pronotum in dorsal view,
PL – maximum length of petiole (in profile),
PW – maximum width of petiole (from above),
PH – maximum height of petiole (in profile),
HTL – maximum length of hind tibia,
PSL – maximum diameter of propodeal spiracle.
Indices:
Cephalic index: CI = HW / HL,
Scape indices: SI1 = SL / HL; SI2 = SL / HW,
Ocular indices: OI1 = OL / HW; OI2 = OL / GL,
Funicular segment indices: FSI1 = FS1 / FS2; FSI2
= FS1 / (FS2+FS3),
Maxillary palps indices: MPI1 = MP4 / MP5; MPI2
= MP4 / (MP5+MP6),
Propodeal spiracle index: PSL / HW,
Alitrunk index: AL / PnW.
Comparative materials, including type specimens of
many species were analysed in the course of this work.
These materials are preserved in the following Museums and Institutions: Zoological Museum of Moscow
State University, Russia (ZMMU); Zoological Institute
of Russian Academy of Sciences, St.-Petersburg, Russia (ZISP); Institute of Zoology of Ukrainian National
Academy of Sciences, Kiev, Ukraine (IZK; including
Karawajew’s collection); The Natural History Museum,
London, UK (BMNH); Museo Civico di Storia Naturale
“G. Doria”, Genoa, Italy (MCSN); Museum and Institute
of Zoology of Polish Academy of Sciences, Warsaw,
Poland (MIZ) and National Museum of Natural History,
Tehran, Iran (MMTT).
Measurements and indices:
HL – maximum length of head in dorsal view, measured in a straight line from the most anterior
point of clypeus to the mid-point of occipital
margin,
HW – maximum visible width of head in dorsal view,
measured above or below of eyes (depending
from species),
SL – maximum straight-line length of scape from its
apex to the articulation with condylar bulb,
FS1, FS2, FS3 – length of 1st to 3rd funicular segments of
antenna,
MP3, MP4, MP5, MP6 – length of 3rd to 6th segments of
maxillary palps,
OL – maximum diameter of eye,
GL – length of gena (seen in profile), measured
TAXONOMY
Cataglyphis stigmatus sp. nov.
Etymology. From the Latin word “stigma” – spiracle, 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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Chapter 8 - Two New Species of Cataglyphis
TWO NEW SPECIES OF THE GENUS CATAGLYPHIS FOERSTER FROM IRAN
71
Ecology. Distribution of this species is probably
limited to the northern coastal plains of the Persian
Gulf. This region is characterized by hot long summer
and mild winter, with mean annual temperature 27C°
and 236 mm precipitation. Phyto-geographically it belongs to the subtropical region. Both nest samples of
C. stigmatus were collected in open arid areas. The
nest entrance had a small mound ca. 5 cm height and
ca. 15 cm in diameter. Specimens were active at the hottest time of day, between 10.00 and 16.00. Workers were
attracted on baits both by tuna fish and sugar syrup.
Comparative diagnosis. Based on all main diagnostic feature (e.g. nodiform petiole, body sculpture,
maxillary palpes structure, etc.; see also Agosti 1990,
Radchenko 1997a), C. stigmatus clearly belongs to the
bicolor species-group. Almost all species of this group
are bicoloured (with reddish head and alitrunk and
black gaster) or black with the only one previously
known exception – C. lunaticus which has entirely
yellow body. Consequently, C. stigmatus obviously differs by colour from all known species of this group,
except of C. lunaticus. Despite we did not investigate
the type specimens of the latter species (it has been
described based on 2 workers from Turkey), the
detailed original description, including morphometric
data and excellent drawings, provided by Baroni Urbani (1969) allow us to compare both species.
C. stigmatus well distinguishes from C. lunaticus
by the longer scape (SI1 > 1.20 vs < 1.10), by the lower propodeum with the dorsal surface being distinctly
longer than the posterior one (the length of the dorsal
surface of propodeum in C. lunaticus is subequal to
the length of posterior one), by the less abundant
standing hairs on the alitrunk and petiole, by the somewhat smaller size, and especially by the much longer
propodeal spiracles. We examined size of propodeal
spiracles in more than fifty Cataglyphis species,
including about twenty ones from the bicolor-group,
but could not found such big spiracles in any of the
investigated specimens.
Material examined. Holotype worker, Iran,
Province Bushehr, Mond protected area, 28°03’N,
51°36’E, 6 m a.s.l., 15 July 2007, arid area, leg. Omid
Paknia, collection code: MND-2128002 (MMTT); paratypes: 1 worker from the nest of holotype; 2 workers
from the same site, but collected on bait traps; 3 workers, Iran, Province Bushehr, Naiband National Park,
27°18’N, 52°48’E, 19 July 2007, 6 m a.s.l., nest sample,
arid area, leg. Omid Paknia (IZK, MMTT).
Description. Workers (Figs 2–7). Species of medium size, body length ca. 5–7 mm. Head with almost parallel sides (below the eyes) and gradually convex occipital margin, occipital corners not marked, head length
subequal to its width. Anterior clypeal margin convex,
without median notch. Clypeal setae distinctly shorter
than length of clypeus and joined near its anterior
margin. Eyes relatively small, their maximum diameter 1.2–1.5 times less than length of genae, situated distinctly beyond the midlength of head margins.
Ocelli relatively big, forming equilateral triangle.
Antennae 12-segmented, scape long, distinctly longer
than head length, first funicular segment distinctly
shorter than the length of second and third segments
together. 3rd and 4th segments of maxillary palpes long,
subequal in length, 5th segment 1.5–1.6 times shorter
than 3rd or 4th ones, 6th segment is the shortest; 3rd segment somewhat flattened, with abundant erect hairs on
inner margin, length of the longest hairs equal or only
a little longer than maximum diameter of the segment;
4th segment with similar pilosity, two apical segments
with abundant but shorter hairs. Mandibles with long
apical tooth, somewhat smaller preapical one and
three small basal teeth.
Alitrunk long and slender, mesonotum not raised
over pronotal level. Propodeum low, gradually arched,
its dorsal surface distinctly longer than posterior one.
Propodeal spiracles elongate-oval, while not distinctly
slit-like, and very long: their length exceeds (or at least
reaches) half of the propodeal height. Petiole obviously
nodiform, with rounded node dorsum.
Surface of whole body with dense microreticulation,
appears dull, although not strongly matt. Body with
sparse whitish standing hairs. Occiput with 5–6
quite long erect hairs, frons with 3–4, clypeus – with 2
similar hairs. Alitrunk and petiolar node with a few
sparse short hairs. Head and gaster with very sparse
and short decumbent pilosity, distance between hairs
longer than hairs’ length; surface of alitrunk (except of
mesonotal dorsum) and coxae with dense silverfish
pubescence. Tibiae with depressed whitish short setae
and additionally with a few yellowish bristles on inner
margin. Antennae with fine, short depressed pubescence, without semi-erect hairs.
Whole body yellow to orange-yellow.
Queens and males are unknown.
Measurements and indices see in Tables 1 and 2.
Cataglyphis pubescens sp. nov.
Etymology. From the Latin word “pubescens ” –
pubescent, that means character of the depressed
pubescence on the head and alitrunk.
Material examined. Holotype worker, Iran,
province Yazd, Siahkooh National Park, 32°35’55”N,
54°13’57”E, 987 m, 23 May 2008, nest sample, arid area,
leg. Omid Paknia, collection code: SIA 2459009
(MMTT); paratypes: 6 workers from the nest of holotype; 2 workers from the same locality but collected by
pitfall traps (IZK, MMTT).
Description. Workers (Figs 8–13). Species of
small size, body length ca. 4 mm. Head length subequal
126
Chapter 8 - Two New Species of Cataglyphis
72
A. RADCHENKO and O. PAKNIA
2 mm
2
1 mm
3
0.5 mm
4
0.5 mm
6
5
0.5 mm
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).
127
Chapter 8 - Two New Species of Cataglyphis
TWO NEW SPECIES OF THE GENUS CATAGLYPHIS FOERSTER FROM IRAN
73
deum subequal to posterior one, both meet at a rounded blunt angle. Propodeal spiracles small, slit-like.
Petiole squamiform, with distinct, rather thick scale.
Surface of head and propodeum with fine but dense
microreticulation, appears dull, promesonotum and
gaster with very fine superficial microreticulation,
appear shiny.
Body with sparse whitish standing pilosity, while it
is somewhat more abundant than in the most of species
of the cursor- and emmae-group. Occiput with more
than 10 straight erect hairs, frons and clypeus without
such hairs. Alitrunk and coxae with scattered erect
hairs of different length, petiolar scale with a few short
hairs. Head (especially temples and occiput), mesopleura, propodeum and coxae with dense pubescence,
formed by long, very abundant silverish appressed
hairs. Gaster with very sparse and short decumbent
hairs. Scape and funiculus with short, quite thick,
whitish subdecumbent hairs, tibiae with numerous,
rather long subdecumbent to suberect setae, and additionally with less abundant yellowish bristles on inner
margin. Whole body black.
Queens and males are unknown.
Measurements and indices see in Tables 1 and 2.
Ecology. This species was collected in the interior
region of the Central Persian desert basin. This area is
to its width; head slightly narrowed anteriorly, with
straight (not convex) sides (below the eyes), rounded
occipital corners and very weakly convex occipital
margin. Anterior clypeal margin almost straight, without median notch. Clypeal setae subequal to clypeal
length and joined near its anterior margin. Eyes relatively small, their maximum diameter ca 1.05–1.25
times less than length of genae, situated distinctly
beyond the midlength of head margins. Ocelli small,
forming equilateral triangle. Antennae 12-segmented,
scape relatively short, subequal or only slightly longer
than head length; first funicular segment relatively
long, only slightly shorter than length of second and
third segments together, remainder segments distinctly longer than broad. 3rd and 4th segments of maxillary
palpes rather long, subequal in length, 5th segment
short, 1.5–1.9 times shorter than 4th ones, 6th segment
only slightly shorter that the 5th one; 3rd segment not
flattened, oval in cross-section, with not abundant
erect hairs, length of the longest hairs less than twice
longer than maximum diameter of the segment; 4th-6th
segments with abundant but somewhat shorter pilosity. Mandibles with long apical tooth, somewhat smaller
preapical one and three small basal teeth.
Alitrunk relatively short and robust, mesonotum
not raised over pronotal level. Dorsal surface of propo-
Table 1. Measurements (in mm) of Cataglyphis stigmatus sp. nov. and C. pubescens sp. nov.
C. stigmatus (n=7)
Measurements
C. pubescens (n=9)
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
128
Chapter 8 - Two New Species of Cataglyphis
74
A. RADCHENKO and O. PAKNIA
8
1 mm
10
9
0.5 mm
0.5 mm
0.2 mm
11
12
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).
129
Chapter 8 - Two New Species of Cataglyphis
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)
Indices
C. pubescens (n=9)
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
mean annual temperature 19°C, and by the very low
annual precipitation – 67 mm only. This territory
belongs to the Irano-Turanian phyto-geographical
region. Most specimens were collected by hand from
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
species-groups of Cataglyphis. Thus, setae on the
anterior clypeal margin are very long, subequal to or
even somewhat longer than the length of clypeus, similarly to C. emeryi (Karawajew), but unlike the latter
species these setae join close to the anterior clypeal
margin, as in the species of cursor-group (Radchenko
1997a, 1998). The first funicular segment is quite long,
about twice longer than the second one and only slightly shorter than the second and third segments together: this is one of the diagnostic features of workers of
the emmae-group (according Agosti 1990). On the other hand, worker caste of C. pubescens is not dimorphic
(the latter is characteristic for the emmae-group
species); additionally, they have distinctly thicker petiolar scale than C. emeryi. In general, workers of the
species of all three groups mentioned above are superficially quite similar to one another, particularly their
whole body is blackish-brown to black, they have petiole with distinct scale (i.e. it is not cuneiform or nodiform), but their males well differ by the structure of
genitalia (Agosti 1990; Radchenko 1997a). Moreover,
workers of C. emmae and C. emeryi move slowly,
rather like Proformica Ruzsky species (C. emmae
has been originally described as a member of
Proformica) than Cataglyphis, while C. pubescens
move very fast, like most of the Cataglyphis species.
The proper taxonomic position of this species can be
definitively resolved when males will be found. Despite
this little taxonomic vagueness, C. pubescens clearly
differs from any knows species of the groups mentioned above by the much more developed, dense
appressed pubescence on the head and alitrunk.
ACKNOWLEDGEMENTS
We are sincerely grateful to curators of the ant collections of all Museums and Institutions mentioned
above for the assistance during our investigations. We
thank Martin Pfeiffer and Elisabeth Kalko for their
kind support of O. Paknia study in Germany, and to
Hans-Peter Katzmann (Ulm University, Germany) for
making photos of the holotype specimens. We are also
grateful to the staff of Department of Environment of
Iran for the assistance and help that allowed that material to be collected from natural reserves in Iran. We
are also grateful to X. Espadaler (Universitat Autònoma de Barcelona, Spain) and B. Markó (Cluj University, Romania) for the available comments to the manuscript of this paper.
REFERENCES
Agosti, D. 1990. Review and reclassification of Cataglyphis
(Hymenoptera, Formicidae). Journal of Natural History,
24: 1457–1505.
Agosti, D. 1994. A new inquiline ant (Hymenoptera, Formicidae) in Cataglyphis and its phylogenetic relationship.
Journal of Natural History, 28: 913–919.
Baroni Urbani, C. 1969. Una nuova Cataglyphis dei monti
dell’Anatolia. Fragmenta entomologica, 6(3): 213–222.
130
Chapter 8 - Two New Species of Cataglyphis
76
A. RADCHENKO and O. PAKNIA
Cataglyphis Foerster (Hymenoptera, Formicidae) from
Asia. Entomologicheskoe obozrenie, 76 (2): 424–442 (in
Russia; English translation: Entomological Review (Washington), 1997, 77: 684–698).
Radchenko, A. G. 1997b. Cataglyphis zakharovi sp. nov. –
the second socially parasitic species in the genus Cataglyphis Foerster (Hymenoptera, Formicidae). Annales
Zoologici, 46: 207–210.
Radchenko, A. G. 1998. A Key to ants of the genus Cataglyphis Foerster (Hymenoptera, Formicidae) from Asia.
Entomologicheskoe obozrenie, 77 (2): 502–508 (in Russia;
English translation: Entomological Review (Washington),
1998, 78: 475–480).
Bolton, B. 1995. A new general catalogue of the ants of the
World. Cambridge-London, Harvard University Press,
504 pp.
Bolton, B., Alpert, G., Ward, P. S. and P. Naskrecki. 2007.
Bolton’s Catalogue of Ants of the World. Harvard University Press, Cambridge-London, CD-Rom.
Paknia, O., Radchenko, A., Alipanah, H. and M. Pfeiffer. 2008.
A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News, 11: 151–159.
Paknia, O. Radchenko, A. and M. Pfeiffer. 2009. New records
of ants (Hymenoptera, Formicidae) from Iran. Asian Myrmecology, 3: 29–38
Radchenko, A. G. 1997a. A review of ants of the genus
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 semiarid 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.
135
Chapter 9 - Synthesis
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 seedeating 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
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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
136 Chapter 9 - Synthesis
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 fineto-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
139 Summary
describing two species new to science. Additionally, I provided the first specieslist 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.
140 Summary 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.
141 Summary
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
142 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.
143 ‫‪Summary‬‬
‫ﺧﻼ‬
‫ﻨﺎ ﻖ ﺸﮏ‪ ،‬ﻪ ﺸﮏ و ﻪ ﻮب‬
‫و ه‬
‫از ﻞ و ﺖ ﺻﺪ از ﺢ ز ﻦ را ﯽ ﭘﻮﺷﺎ ﻨﺪ‪ .‬ﺑﺎ و ﻮد ﻮ ﯿﺪ او ﻪ ﮐﻢ‪ ،‬ا ﻦ ﻨﺎ ﻖ ﻮع ﻮ ای ﺑﺎﻻ ﯽ را‬
‫ﻮص ﯽ از ﺟﺎ ﺪاران و ﯿﺎ ن ﻣﺎ ﻨﺪ ﻮر ‪ ،‬ﻮﺳﮏ و ﯿﺎ ن ﯾﮏ ﺳﺎ دارا ﯽ ﺑﺎ ﻨﺪ‪ .‬ﻨﺎ ﻖ ﺸﮏ ھﻢ ا ﻮن ﺮات ﺮ‬
‫ﻂ ز ﺖ د ﺖ ا ﺴﺎن را‬
‫ﻮص د ﻮ ﯽ ی آب و ﻮا ﯽ و‬
‫ﺑﺎره ﺮا ﻂ ﻮع ﻮ ای‪ ،‬ﺮﭘا ﻨﺪ ﯽ ﻮ‬
‫ﻮل‬
‫و ه‬
‫ﻋ‬
‫ﯽ ﻨﺪ‪ .‬ک ا ﻦ ﺮات ﻨﺎ ﻖ ﺸﮏ ﯿﺎز ا ا ﺶ دا ﺶ ﻮ ﯽ‬
‫ی ﯽ و ﻨﺎ ﺖ ﻮر ﯽ دارد‬
‫ﻦ ﺮ را ﻮع ﻮ ای دار ﺪ‪ .‬ا‬
‫ﻨﺎ ﻊ اوا ﯽ‬
‫ﻮص ﻮع ﻮ ای ﺟﺎ ﻮران و ﯿﺎ ن ﻨﺎ ﻖ ﺸﮏ ا ﺮا ﯿﺎ و آ ﮑﺎی ﻤﺎ ﯽ و ﻮد دارد‪ ،‬ﻦ اﻃﻼﻋﺎ ﯽ دﯾ ﻨﺎ ﻖ ﻣﺎ ﻨﺪ ﻮا ﯽ ﺸﮏ و ﻪ ﺸﮏ‬
‫ﺴ‬
‫ﻮان از ﻮ ﻖ ﻦ د ﻪ از ﺟﺎ ﺪا ران ﻤﺎ ﯽ ز ﺖ ﻮم ی ﺸﮏ د ﯿﺎ ﺳﺎ ﻦ ﺪه ا ﺪ و ﺖ ﻏﺎ‬
‫ﭘﺎ ﺌﺎر ﯿﮏ ﯾﺎب ﯽ ﺑﺎ ﺪ‪ .‬ﻮر‬
‫ﭼ‬
‫ﺮات را ﻨﺎ ﻖ ﯿﺎﺑﺎ ﯽ و ﺰار‬
‫ﺮار د ﯿﺎ‬
‫ﻮ ﻮع ﮐ ﯽ ا ﻦ رﺳﺎ‬
‫ھ‬
‫ﻋ‬
‫ا ﻦ ﻨﺎ ﻖ ﻦ ا اء ﮫﻢ ﻤﻠ د ا ﻮ ﻮژ ز ‪ A‬ﮕﺎه ﯽ ﺑﺎ ﻨﺪ و ﺶ ی ﮫﻢ‬
‫ﯿﺎر ﮫﻢ ﻮا ﯽ ﺸﮏ‪ ،‬ﻮر‬
‫ا ﻮ ﻮژ را ﮫﺪه دار ﺪ‪ .‬ﻋ ﯽ ر ﻢ ﺶ ا ﻮ ﻮژ‬
‫ﺮش دا ﺶ ﻮ ﻮد‬
‫ﻞ ﯽ د ﻨﺪ‬
‫ا ﻤﺎع‬
‫ﯿﺎری از ﻨﺎ ﻖ ﯿﺎﺑﺎ ﯽ و ﻪ ﯿﺎﺑﺎ ﯽ ﭘﺎ ﺌﺎر ﯿﮏ ﻨﺎ ﻪ ﻣﺎ ﺪه ا ﺪ‪.‬‬
‫ﻮص ﺮ ﻮا ﻞ ﯽ و ﻮر ی ا ﯿﺎ ﯽ روی ﻮع ﻮ ای و ﻨﺎی ﻮ ای و ﺳﺎ ﺘﺎر ا ﻤﺎ ﯽ ﻮر‬
‫ﻋ‬
‫ﻨﺎ ﻖ ﺸﮏ و ﻪ ﺸﮏ ا ان ﻮده ا ﺖ‪ .‬ای ﻖ ا ﻦ ﻮ ﻮع‪ ،‬دو ﻋﻤ ﯿﺎت ا ﯽ ﺳﺎل ی ‪ ١٣٨۶‬و ‪ ١٣٨٧‬ﯽ ا ان ﻤﻞ آ ﺪ‪.‬‬
‫او ﻦ ﻋﻤ ﯿﺎت ا ﯽ ﻮر‬
‫ﻪ ﯿﺎﺑﺎن ی‬
‫ﻮل ا ﯿﺎ ﯽ ﺑﺎ ﺬر از ﮫﺎر ﻪ ز ﯽ )‪ ١٢٠٠‬ﯿ ﻮ ﺮ( ﻊ آوری دﯾﺪ ﺪ‪ .‬دو ﻦ ﻋﻤ ﯿﺎت ا ﯽ‪ ،‬ﻮر از زی ‪-‬‬
‫ی ا ان ﻊ آوری ﺪ ﺪ‪.‬‬
‫‪144 ‬‬
‫‪ ‬‬
‫‪Summary ‬‬
‫رﺳﺎ ﺣﺎ ﺮ ﺷﺎ ﻞ ﮫﺎر ﺣ ﻪ ﻮده ا ﺖ‪ .‬ﺣ ﻪ ﺖ ﺷﺎ ﻞ ﻄﺎ ﻌﺎت ﻮ ﯿﮏ و ﻮ ﻮ ﯿﮏ روی ﻮن ا ان ﻮده ا ﺖ‬
‫ﻮرد ﻄﺎ ﻪ ار‬
‫ﻪ ا ﺖ ) ﻞ ی ‪ ٧ ،۶‬و ‪ (٨‬و ﻮان ﭘﺎ ای ای ﻄﺎ ﻌﺎت ا ﻮ ﻮژ‬
‫ﻮل ا ﯿﺎ ﯽ ﺷﺎ ﻞ دو ز ﻮم ﭘ ﻦ د ﺖ و ﯿﺎﺑﺎن ﺮ ﺢ و‬
‫ﺣ ﻪ دوم ﺳﺎ ﺘﺎر ﺮﭘا ﻨﺪ ﯽ ﻮر‬
‫ﻮ ای ﻮر ﺑﺎ روﯾ د ء ﻨﺪی‬
‫ا ان ﺢ ﻮده و‬
‫ﻪ ای از دﯾﺪﮔﺎه ﻔﺎ ﯽ ﻮرد‬
‫ﯽ وا ﺪ ی ز ﯽ‪ -‬ا ﯿﺎی‬
‫ﺤ‬
‫و ﻞ ار‬
‫ﻞ ی ﻌﺪ ﯽ ﺑﺎ ﺪ‪.‬‬
‫ﺤ‬
‫و ﻞ ﺪ ) ﻞ ‪ .(٢‬ﻮ ﻦ ﺣ ﻪ‪ ،‬ﻮع‬
‫‪ .‬ا ﻦ ﻄﺎ ﻪ‪ ،‬ﺑﺎ ض ا ﻪ ﻪ ﻮا ﯽ ز ﯽ ‪WWF‬‬
‫ﻪ را ارا ﯽ ﻨﺪ‪ ،‬ر ﯽ ﺪ آﯾﺎ‬
‫ﻦ ﺰان ﺮ ﻮ‬
‫ﯿﺎس ﻮا ﯽ ز ﯽ ا ﺠﺎم ﯽ ﺮد و‬
‫ﯿﺎ ﯽ ﻮﭼﮏ ) ﻞ ‪ .(٣‬ﮫﺎر ﻦ ﺣ ﻪ ﺰان ﺮ ﻮا ﻞ ﯽ و ﻮر ی ا ﯿﺎ ﯽ روی ﻮع ﻮ ای و ﻨﺎی ﻮ ای و ﺳﺎ ﺘﺎر ا ﻤﺎ ﯽ‬
‫ﯾﺎ‬
‫ﻮر‬
‫ﻮل ا ﯿﺎ ﯽ و زی ‪ -‬ﻪ ﯿﺎﺑﺎن ی‬
‫ﺮات ﻨﺎی ﻮ ای ﻈﺎم‬
‫ﻪ ای ﯽ ا ا ﺶ ﻮل ا ﯿﺎ ﯽ ا ا ﺶ ﯾﺎ‬
‫ﻪ ای ﮐﺎ ﺶ ﭘﯿﺪا د‪ .‬ز ﻮم ﭘ ﻦ د ﺖ دارای ﻨﺎی ﻮ ای‬
‫ﻪ ای ﺸﺎن داد‬
‫ﻮل ﮫﺎر زی ‪ -‬ﻪ ﻮرد ﻄﺎ ﻪ ﺸﺎ ﺪه ﺪ‪ .‬ا‬
‫ی‬
‫ز ﻮم ﯿﺎﺑﺎن ﻮد‪.‬‬
‫ا ان ﺸﺎن ﯽ د ﺪ و‬
‫ﯾﮏ ﻮرد ز ﻨﺎ ﻪ ﺪه‬
‫ا ﻦ ﮫﺎر ﺳﺎ ﺘﺎر ﺺ ﻮ ای ﻔﺎوت از ز ﺺ ﺪه‬
‫ﻮل ا ﯿﺎ ﯽ و‬
‫ھ‬
‫ﻦ زی ‪ -‬ﻪ ﯿﺎﺑﺎن ی‬
‫ﺤ‬
‫و ﻞا‬
‫ﻪی‬
‫ﯽوا‬
‫ی ا ان ﺸﺎن داد ﺮا ﻂ آب و‬
‫ﻮای ﻌﺎ ﺮ ﻣﺎ ﻨﺪ ﺑﺎر ﺪ ﯽ و ﺮات دﻣﺎ ﯽ و ﻮ ﯿﺪ او ﻪ ﯿﺎ ﯽ دارای ﺮ زﯾﺎدی روی ﻮع ﻮ ای و ﺳﺎ ﺘﺎر ﻮ ای ﻮر‬
‫‪145 ‬‬
‫و‬
‫ﻦ ﺮات ﻨﺎی ﻮ ای ا ﻦ ﯿﺎس و ا ﻦ ﻻ رخ داد‪ .‬ﮫﺎر ﺳﺎ ﺘﺎر ﺺ ﻮ ای‬
‫ﺖ و ﺪرت ز ی ﻮا ﯽ ز ﯽ ﺑﺎﯾﺪ ﻮر ﺪا ﮔﺎ ای ﻮرد ﻄﺎ ﻪ ار ﺮد‪.‬‬
‫ا ﯿﺎ ﯽ روی ﻨﺎی ﻮ ای و ﺳﺎ ﺘﺎر ﻮ ای ﻮر‬
‫اﻣﺎ‬
‫را ﻮل ﻻ ی ﻮﭼﮏ رگ‬
‫ﯽ وا ﺪ ی ز ﺖ‪ -‬ا ﯿﺎ ﯽ ﻮر‬
‫ﻪ ی ﻮا ﯽ ز ﯽ ‪WWF‬‬
‫ﻨﺎ ﺐ ای ﻄﺎ ﻌﺎت ﻔﺎ ﯽ ﯽ ﺑﺎ ﺪ د ﻞ آﻧ ﻪ‬
‫‪ WWF‬ﻮد‪ .‬ا ﻦ ﯾﺎ ﻪ ﺸﺎن ﯽ د ﺪ‬
‫) ﻞ ‪ ۴‬و ‪.(۵‬‬
‫ﺤ‬
‫و ﻞ داده ی ﻨﺪﻻ ای ا ا ﺶ ﻨﺎی ﻮ ای ﻮر‬
‫ﯽ را ﯽ ﻮد‪.‬‬
‫ھ‬
‫ﺸﺎن داد‪ .‬ﺷﺎ ﺺ ﻮاری ﻮل ا ﻦ ﺳﺎ ﺘﺎر‬
‫ﻞ ا ﺑﺎ ء ﻨﺪی‬
‫ی ا ان ﻮرد ر ﯽ و‬
‫ﺤ‬
‫و ﻞ ار‬
‫ﻮل ا ﯿﺎ ﯽ ﺘﺎ ﺞ ﺸﺎن داد ﻨﺎی ﻮ ای دا ﻞ‬
‫ﺑﺎ ر ﯽ ﺳﺎ ﺘﺎر ا ﻤﺎ ﯽ ﻮر‬
‫ﺤ‬
‫ای ﺳﺎ ﺘﺎر ا ﻤﺎ ﯽ ﻮر‬
‫ﻮن ﻮرت ﺰ‬
‫ﻨﺪ‪ .‬ﺣﺎ ﻞ ﻘﺎت‬
‫‪ ‬‬
‫‪ ‬‬
‫‪Summary‬‬
‫ﻮ ﯿﮏ و ﻮ ﻮ ﯿﮏ روی ﻮر ی ا ان ارش ﯾﮏ ز ﺧﺎ ﻮاده‪ ،‬ﺖ ﺲ و ﯽ و ﻮ ﺪﯾﺪ ای ا ان و دو ﻮ ﺪﯾﺪ ای د ﯿﺎ ﻮد‪ .‬ﻋﻼوه‬
‫ا ﻦ ای او ﻦ ﺑﺎر ﺖ ﻮ ی ﻮر ی ا ان ﺰ ﺮ دﯾﺪ‪.‬‬
‫ﺧﺎ ﻪ‪ ،‬ﻮا ﯽ ﺸﮏ و ﻪ ﺸﮏ ا ان دارای ﻨﺎی ﻮ ای ﺘﺎ ﺑﺎﻻ ﯽ ﯽ ﺑﺎ ﻨﺪ‪.‬‬
‫ﺳﺎل ﻊ آوری دﯾﺪ‪ .‬ا‬
‫ﻨﺎی ﻮ ای دا ﻞ‬
‫ﻪ ای ﺘﺎ ﭘﺎ‬
‫از ﻮد ﻮ ﻮر‬
‫دا ﻞ ﮫﺎرده‬
‫ﺤ‬
‫و ﻞ ی ا ﺠﺎم‬
‫ﻮد و از ﺪه ﺠﺎوز ﻮد‪،‬‬
‫ﺮات ﻮ ای ﻤﺎ ﯽ ﯿﺎس ی ا ﯿﺎ ﯽ زﯾﺎد و ﯽ دار ﯽ ﺑﺎ ﺪ ﺸﺎن ﯽ د ﺪ از ﻮ ی ﯿﺎری از ﻮر‬
‫دارای ﺮﭘا ﻨﺪ ﯽ ﺪود ﯽ ﺑﺎ ﻨﺪ‪ .‬ا ﻦ اﻟ ﻮی ﺮﭘا ﻨﺪ ﯽ ﻮر‬
‫ﺮﭘا ﻨﺪ ﯽ ﺪود ﺪه ﯿﺎری از ﻮ ی ﻮر آ ﮫﺎ را ا ا اض ﻪ ای و‬
‫ﻮا ﯽ ﺸﮏ و ﻪ ﺸﮏ ا ان‬
‫ﻪ ای آ‬
‫ﻮا ﺪ دا ﺖ‪.‬‬
‫ﻮع ﻮ ای ﻮر ﭘﺪﯾﺪ ﯽ آورد‪.‬‬
‫ﭘﺬ ﯽ ﻤﺎﯾﺪ ‪ .‬ﻋﻼوه ا ﻦ از آ ﺠﺎ ﻨﺎی ﻮ ای و ﺳﺎ ﺘﺎر‬
‫ﺰان زﯾﺎدی ﺮا ﻂ آب و ﻮا ﯽ ﺘ ﯽ دارد‪ ،‬ﺮات آب و ﻮا ﯽ ﺶﯿﭘ‬
‫ﺮده د ر ﺰان ﺑﺎرش ﺑﺎران ا ﻤﺎل زﯾﺎد ﺮ زﯾﺎدی روی ﻮر‬
‫ﻮا ﺪ ﻮد و ﻮر‬
‫ﻪ ﺸﺎن داد‬
‫ﺰان‬
‫ﺸﺎن ﯽ د ﺪ ﻨﺎ ﻖ ﺸﮏ ﮫﺎ ﻨﺎ ﯽ و ﻊ و ﯾ ﻮا ﺖ ﯽ ﺑﺎ ﻨﺪ‪ .‬ﺧﻼف آن ا ﻦ ﻨﺎ ﻖ‬
‫دارای ﺳﺎ ﺘﺎری ﺠﺎ ﺲ ﻮده ﺮﭘا ﻨﺪ ﯽ ﯿﺎری از ﻮ را ز ‪ A‬ﮕﺎه ی ﻮﭼﮏ ﺪود ﯽ ﻤﺎﯾﺪ و ﺳﺎ ﺘﺎری ﺗ ﻪ ﺗ ﻪ‬
‫ﻮ ای ﻮر‬
‫ﻪ ﻔﺎ ﺖ ﺪه ﻮل دو‬
‫ﻪ ﻣﺎ ﻨﺪ ا ا ﺶ ﺰان ﯽ و دﻣﺎ و ﺮات‬
‫ﺪه‬
‫ا ﻦ ﺮان ا ﻤﺎل اوان ‪ ،‬ﺮ ﺮﭘا ﻨﺪ ﯽ‬
‫ﯽ ﻮا ﯽ ﺳﺎزﮔﺎری ﺑﺎ ﺮا ﻂ ﺪﯾﺪ را ﺪا ﻪ ﺑﺎ ﻨﺪ و ﻮا ﻨﺪ ﭘﻨﺎ ﮕﺎه ﻨﺎ ﯽ ﯿﺎ ﻨﺪ‬
‫ﯿﺎس‬
‫ﻪ ای و‬
‫ﻪ ای‬
‫ﯽ از ﻮر‬
‫ض ﻮا ﻨﺪ ﺪ‪.‬‬
‫‪146 ‬‬
‫‪ ‬‬
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.
148
General Appendix
Figure 3: Pictures of the second field work in 2008. From tope left to right down: 1Plastic quadrate used for measuring habitat heterogeneity in sampling sites, 2Measuring 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.
149 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
151 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.
152 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
University.
Research scientist and instructor, Institute of Biology, Gorgan
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
153
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: 151159
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.
155
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
156
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
157 158