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Pollen analysis of coprolites from a late PleistoceneeHolocene cave deposit(Wezmeh Cave, west Iran): insights into the late Pleistocene and late Holocenevegetation and flora of the central Zagros Mountains

Morteza Djamali a,*, Fereidoun Biglari b,c, Kamyar Abdi d, Valérie Andrieu-Ponel a,Jacques-Louis de Beaulieu a, Marjan Mashkour e, Philippe Ponel a

a Institut Méditerranéen d’Ecologie et de Paléoécologie (IMEP), UMR 6116 CNRS, Europôle Méditerranéen de l’Arbois, F-13545 Aix-en-Provence Cedex 04, Franceb Paleolithic Department, National Museum of Iran, 30 Tir St., Emam Khomeini Ave., P.O. Box 11365/4364, Tehran, IrancUMR 5199-PACEA, University Bordeaux 1, Avenue des Facultés, F-33405 Talence, FrancedDepartment of Archaeological Sciences, Shahid Beheshti University, Tehran, Irane Laboratoire d’Archéozoologie et Archéobotanique, UMR 7209 CNRS, Département Ecologie et Gestion de la Biodiversité, Muséum National d’Histoire Naturelle (MNHN),55 rue Buffon, F-75005 Paris, France

a r t i c l e i n f o

Article history:Received 4 March 2011Received in revised form7 July 2011Accepted 3 August 2011

Keywords:CousiniaTulipaIrano-Turanian regionMiddle EastCoprolitesLate Quaternary

a b s t r a c t

Coprolite pollen analysis is increasingly used by palynologists to reconstruct the palaeovegetationespecially in the arid to semi-arid regions where lakes and peat bogs are scarce. This study is the firstpalynological analysis of coprolites in Iran to examine their potential in palaeoenvironmental recon-structions. Four pollen-rich coprolite samples were studied from the Wezmeh Cave in western Iran witha recently discovered late Quaternary (70 ka to sub-recent) faunal assemblage dominated by carnivoreremains. Pollen analysis of coprolites shows that a mountain steppe dominated the glacial landscapes ofthe area. Only one sub-recent sample indicates the presence of dispersed tree stands. One sample wasrich in Tulipa pollen suggesting that tulips were relatively abundant during the glacial periods. Cousinia(Asteraceae) pollen was found in all samples with considerable values in one sample indicating itsimportance in the glacial landscapes of the Zagros Mountains. This study revealed that coprolite pollenanalysis can provide an invaluable source of information to understand the floristic composition ofpalaeolandscapes of the Irano-Turanian region.

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1. Introduction

Coprolites are increasingly used by palynologists for pastvegetation reconstructions (e.g. Scott, 1987; Vivent, 1989;Fernández Rodríguez et al., 1995; Carrión et al., 2001, 2005, 2007;Guérin et al., 2004; Gonzáles-Sampériz et al., 2003; Scott et al.,2003). They are especially a good source of palaeoenvironmentalinformation in arid and semi-arid regions, where themost commonpollen traps i.e. lakes and peat bogs, are rare or absent (Scott, 1987;Gonzáles-Sampériz et al., 2003). Hyena coprolites have beendemonstrated to reflect a relatively unbiased picture of the regionalterrestrial vegetation (Carrión et al., 2001; Gonzáles-Sampérizet al., 2003; Scott et al., 2003). However, few studies have treatedthe role of animal behaviour and taphonomic history in the

incorporation and preservation of pollen grains in the hyenacoprolites (e.g. Scott et al., 2003). Several sources of biases mayhave been underestimated in the interpretation of the coprolitepollen assemblages such as the possible incorporation of olderpollen in the soils, sediments, and cave deposits dug by the animal.Future studies on the complex taphonomy and taphonomic bias ofcoprolites are therefore required to test the validity of vegetationreconstructions based on coprolite pollen analysis.

A recent discovery of several coprolites associated with a latePleistocene faunal assemblage in the Wezmeh Cave in western Iran(Mashkour et al., 2008), offers an opportunity to examine thepotential of coprolite pollen analysis to reconstruct the past vege-tation and flora of the central Zagros Mountains. TheWezmeh Cavefaunal assemblage was recovered in 1999 and 2001 from the lootedand reworked deposits of the cave and was studied by Mashkouret al. (2008). The site is the only known cave with a long and richsequence of Upper Pleistocene deposits in the Islamabad Plainwhich based on U-series dating have been used mainly by local

* Corresponding author.E-mail address: morteza.djamali@yahoo.com (M. Djamali).

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carnivores since ca. 70 ka BP. The site has also yielded the onlyknown Upper Paleolithic human remain in western Zagros datingback to about 20e25 ka BP based on non-invasive spectrometrygamma dating (Trinkaus et al., 2008). However, the environmentalcontext of the Wezmeh Cave is almost completely unknown. Thisstudy will try to provide some palaeoenvironmental informationbased on coprolite pollen analysis of the cave.

Several coprolite samples were discovered and collected by oneof the authors (F.B.) in a recent visit to the site in October 2008.During that visit sizeable samples of faunal remains and somecoprolites were collected on the reworked deposits left by looterson the slope in front of the cave entrance. Unfortunately, thenumber of these coprolites was limited and only seven samplescould be used for pollen analysis. They were allocated to hyena

(Crocuta crocuta) according to their morphology and size compa-rable to the modern feces found in the wild in Africa (PhilippeFosse, pers. comm.).

A few pollen diagrams available from the Zagros Mountains(Lakes Mirabad, Zeribar, and Urmia; Fig. 1A) suggest that thevegetationwas profoundly affected by Quaternary climatic changes(van Zeist and Bottema, 1977; Bottema, 1986; Djamali et al., 2008a).However, these diagrams do not give much detailed informationabout the floristic changes of vegetation and the specific taxapresent in the upland areas of the Zagros in glacial and postglaciallandscapes. This is partly because of the biased picture of vegeta-tion reflected in the pollen diagrams due to the absence or under-representation of some autogamous or entomophilous plants (e.g.Fabaceae, Plumbaginaceae, Rosaceae) and the over-representation

Fig. 1. A. Geographical location of the Wezmeh Cave in the northwestern Zagros Mountains. B. General view of the Wezmeh Cave area. The cave entrance measures 2 m wide and1.2 m high.

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of other anemophilous groups (e.g. Artemisia, Chenopodiaceae) inthe pollen rain (Moore and Stevenson,1982; Djamali et al., 2009). Inaddition, the abundant pollen produced by local aquatic andhalophytic vegetation in marginal zones of the aforementionedlakes strongly masks the pollen rain produced by the uplandvegetation (e.g. Wasylikowa, 2005; Djamali et al., 2008b). Pollenanalysis of the Wezmeh Cave could, therefore, potentially help toinfer the past vegetation composition of the well-drained slopes ofthe central Zagros.

This short contribution aims at (i) improving the chronologicalframework of the fossil fauna of the Wezmeh Cave by a fewradiocarbon datings on the organic content of the coprolites and(ii) deducing the local and regional vegetation and its floristiccomposition of the landscapes around the cave during the lastglacial time, as reflected in the pollen assemblages of the coprolites.Here, we discuss the potential of pollen analysis of coprolites as analternative palaeoecological technique in the semi-arid to aridareas of the Irano-Turanian region.

2. Wezmeh Cave and its research background

The Wezmeh Cave is located at about 12 km to the southeast ofthe town of Islamabad-e-Gharb (34�0300400 N, 46�3804200 E), at anelevation of 1430 m a.s.l., in a small valley side tucked into thewestern foothills of the Qazivand Mountain. The mouth of the cavefaces north and is 2 mwide and 1.2 m high. The cave is about 27 mlong and has about 45 m2 of floor area.

The site was located and preliminarily sampled during thesecond season of archaeological research in the Islamabad Plain in1999 directed by K. Abdi (Abdi et al., 2002). About 4 years before thediscovery of the site, the main body of the Upper Pleistocenedeposits at the rear of the cave was dug by looters to the depth ofabout 4 m and was dumped on the slope in front of the caveentrance. The closest source of water to the Wezmeh Cave isa spring at about 200 m to the east which dried up long time ago.However, a stand of a few elm trees (“Wezm” in local Kurdishdialect thereby the name Wezmeh) is found around the springsuggesting some near surface groundwater.

The first survey and later excavations in the cave yielded Earlyand Middle Chalcolithic archeological remains, but the surfacecollecting on the reworked deposits on exterior slope in 1999, 2001and 2008 yielded an abundant late Pleistocene faunal assemblage,particularly rich in carnivore remains including a human premolardating to 20e25 ka BP (Trinkaus et al., 2008).

3. Climate and vegetation

In the Global Bioclimatic Classification System (Rivas-Martínezet al., 1999), the climate of this area is Mediterranean pluviseasonalecontinental with two peaks of precipitation during the winter andspring months (P ¼ 445 mm/yr, T ¼ 15 �C at Kermanshah). However,the continentality index (Ic¼ Tmax (July temp.)� Tmin (January temp.))is higher compared to the Mediterranean region (Ic ¼ 26.9 �C atKermanshah). Today, the regional vegetation is disturbed by humanactivities and the area is also under extensive dry farming of cereals(see vegetation map of Kermanshah, 1998 by Iranian Ministry ofAgricultural Reform). The potential vegetation is most probablya xerophytic deciduous oak woodland according to the vegetationmap of Zohary (1973).

4. Materials and methods

Seven coprolite samples were selected for pollen analysis(Table 1; Fig. 2). Surface of coprolites was cleaned to avoid anycontamination with modern pollen. Pollen grains were extracted

following Moore et al. (1991) and were then boiled in acetolysissolution for 5e6 min. Over 330 pollen grains were counted for eachsample under the light microscope at 500� magnification. Pollengrains were determined using different pollen atlases (Reille, 1992,1995, 1998; van Zeist and Bottema, 1977) and pollen referencecollection of IMEP (Institut Méditerranéen d’Ecologie et dePaléoécologie). The pollen percentage values were calculated inTilia and then illustrated in TGView (Grimm, 2004/2005). In orderto estimate the pollen concentrations in the samples (Stockmarr,1971), one Lycopodium tablet was added to each sample beforestarting the pollen extraction. For 14C dating, organic matter wasextracted from three coprolites by successive treatments in HCl andHF avoiding the utilization of any chemical composition containingcarbon atoms in its structure which might be a source of error inradiocarbon dating.

5. Results

5.1. Chronology

Three of the coprolites were successfully dated and the cali-brated 14C ages are reported in Table 2. Calibrations were per-formed using the last version of Calib software (Calib6) in whicha new radiocarbon calibration dataset (IntCal09) is included(Reimer et al., 2009). Here, we use the mean ages calculated as themean of the 2s-ranges with highest probability. However, thereader is referred to Table 2 for all possible age-ranges for eachcoprolite. Of two polliniferous samples (see Table 1; Fig. 2), WC-1 isof the last glacial age (19228 � 352 cal BP) andWC-2 is of the latestHolocene (405� 59 cal BP). The sampleWC-3 was devoid of pollen,however, its age (12744 � 370 cal yr BP) adds to the chronologicaldata of the Wezmeh Cave faunal assemblages assigned to latePleistoceneeHolocene times (Mashkour et al., 2008). Based on the

Table 1Weights, pollen sum, and pollen concentrations of the examined coprolites.14C-dated samples are marked with asterisks (see Table 2).

Sample number Weight (g) Pollen sum Pollen concentration(no. per g)

WC-1* 8.4 350 3239WC-2* 4.3 314 528WC-3* 3.0 Barren BarrenWC-4 3.9 330 3797WC-5 2.4 Barren BarrenWC-6 3.5 Barren BarrenWC-7 2.2 331 2114

Fig. 2. Photos showing two studied coprolites WC-1 and WC-2.

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similarities of pollen spectra, the WC-4 and WC-7 coprolites arealso assigned to the last glacial period (see below).

5.2. Pollen content

Pollen percentage histograms are illustrated in Fig. 3. Only foursamples (WC-1, WC-2, WC-4, and WC-7) contained pollen grainswith a wide range of concentration values (Table 1). Three othersamples were entirely barren (WC-3, WC-5, and WC-6). Overall thefour samples are dominated by pollen of herbaceous plants(88.2e99.7%) in which the percentages of terrestrial pollen highlyexceed those of aquatic plants (compare Sparganium-type andCyperaceae pollen percentages, with those of total herbaceousplants).

The pollen composition of sample WC-2 is quite different fromthree other samples. It is the only sample displaying significantvalues of tree pollen (11.8%) mainly composed of Quercus,Juniperus and cf Daphne (Fig. 3) and shows the lowest values ofChenopodiaceae and Cichorioideae. This sample also representsa very high content of micro-charcoal grains. However, its mostinteresting aspect is the high percentage of the Cerealia-typepollen (21%) of which 3.2% is almost certainly produced byTriticum species (Fig. 4).

Three other coprolite samples are similar in general character-istics (Fig. 3). However, they show some slight but significantdifferences in the frequencies of some pollen types. The mostremarkable features of these samples are as follows:

(i) predominance of Asteraceae sub-family Cichorioideae pollen(43.3e57.6%),

(ii) considerable values of Chenopodiaceae (Atriplex-type) pollen(5.4e18%) but total absence of Artemisia pollen,

(iii) significant values of Tulipa pollen (Tulipa agenensis-type, seeReille, 1998) in samples WC-4 (1.8%) and WC-7 (1.5%) andrelatively high values of Sisyrinchium-type pollen in WC-4(17.6%),

(iv) presence of Cousinia-type pollen in all the samples but withsignificantly high percentages in sample WC-7 (10.9%),

(v) relatively high values of undetermined pollen (3.3e7.7%).

6. Discussion

6.1. Vegetation

Overall, the pollen assemblages of the four studied coprolitesshow that steppe vegetation dominated the landscape at the timeof the deposition of the Wezmeh Cave faunal assemblage. Onlysample WC-2 suggests the presence of significant stands of treesand shrubs including Quercus, Juniperus, and Thymelaeaceae(most likely Daphne or Dendrostellera). The radiocarbon age is405 � 59 cal BP (Table 2). The contemporaneous vegetation wascomposed of a mosaic of grasslands and/or arable lands with cereal

cultivation (see the high percentages of Poaceae pollen in Fig. 3),with dispersed tree stands. To understand the degree of theopenness of the landscape contemporaneous with sample WC-2,a comparison of the pollen percentages of oak in this sample(8.6%) with surface samples No.12 (15%) and No.13 (30%) of Wrightet al. (1967) is informative. These latter surface pollen samples havebeen taken very close to the Wezmeh Cave and show significantlyhigher values of oak pollen indicating that the tree cover at405 � 59 cal BP was probably lower than today. As the WezmehCave area is located inside the natural distribution range ofthe Zagros oak woodlands, these low pollen values most probablyreflect the human-induced firing of the landscape that favouredthe cereal cultivation and prevented the establishment of densewoodlands four centuries ago. High micro-charcoal content ofWC-2 (Fig. 3) indicates the abundant firing events most likely ofanthropogenic origin. It should, however, be noted that seasonalchanges in oak pollen content in the air may also explain the aboveobservation.

6.2. Notes on floristic composition of glacial steppes of centralZagros

Poaceae pollen percentages of sample WC-2 (38.5%) are mainlycomposed of Cerealia-type pollen (21%) displaying at least 3.2% ofTriticum-type pollen which is almost certainly produced by genusTriticum. It is well-known that the Cerealia-type pollen is notexclusively produced by cultivated cereals (species of Triticum,Hordeum, and Secale). Some wild Poaceae species can also producethis pollen type especially in the Middle East e.g. Aegilops spp. andthe wild ancestors of wheat and barley (van Zeist et al., 1975;Bottema, 1992). Indeed, the Wezmeh Cave area is located withinthe natural distribution range of the wild einkorn wheat (Triticummonococcum subsp. boeoticum), the ancestor of the cultivatedeinkorn (T. monococcum) and the wild barley (Hordeum vulgaresubsp. spontaneum), the ancestor of the cultivated barley i.e.H. vulgare (see maps 1 and 5 in Zohary and Hopf, 1994). TheCerealia-type pollen recorded in the WC-2 sample could have beenproduced by the cultivated cereals as well as the wild grasses fourcenturies ago. The pollen assemblage composition of this samplemost probably indicates an intensive cereal cultivation in the areaat this time.

The long pollen records of Lake Urmia (Djamali et al., 2008a)suggest that the wild cereals were present in very significantamounts in the glacial landscapes of NW Iran and E Anatolia. Incontrast to the modern cultivated wheat and barley which areextremely under-represented in the pollen rain, the wild cerealshave a better representation because the percentages of cross-fertilization and wind-pollination in wild cereals are higher thanin cultivated cereals (Bottema, 1992; Zohary and Hopf, 1994). This issuggested by Bottema (1992) as the main reason why the wildcereals are frequently found in the fossil records. The presence ofwild Cerealia-type pollen in WC-7 is significant and most likely

Table 2Radiocarbon ages of three organic-rich coprolites fromWezmeh Cave. Calibrated dates are reported as means of the 2s-ranges with highest probability. However, all possibleage-ranges with different probabilities (in parentheses) for all dates are reported as a separate column (2s-ranges).

Sample number Lab code 14C age (yr BP) Calibrated age (cal yr BP) Age-ranges (cal yr BP) Material dated

WC-1 Poz-39131 16,180 � 150 19,228 � 352 18,876e19,580 (0.99) Coprolite19,756e19,761 (0.001)

WC-2 Poz-39132 315 � 30 405 � 59 303e343 (0.24) Coprolite346e464 (0.76)

WC-3 Poz-39133 10,790 � 180 12,744 � 370 12,164e12,182 (0.004) Coprolite12,217e12,288 (0.019)12,314e12,352 (0.009)12,374e13,115 (0.96)

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shows the importance of wild cereals in the glacial landscapesaround the cave.

A remarkable feature of the sample WC-4 which is certainlyof last glacial age, is the significant percentages of Sisyrinchium-type and T. agenensis-type pollen. Sisyrinchium-type pollen wasfirst reported by van Zeist and Bottema (1977) from the glacialsediments of Lake Zeribar in the Iranian Kurdistan, NW Iran. Itshows a sporadic occurrence in the BH2 and BH3 pollen diagramsof Lake Urmia during both glacial and interglacial periods(see Djamali et al., 2008a and the website of European PollenDatabase at http://www.europeanpollendatabase.net/). In Europe,this pollen type is produced by Sisyrinchium bermudianum L.

(Iridaceae) (see Reille, 1992, 1998). Its origin in the Zagros regionis still unknown (van Zeist and Bottema, 1977) but the most likelyproducer seems to be Gynandriris sisyrinchium, a species ofIridaceae with a large distribution range in the western part of theIrano-Turanian region including Iraq, Iran, and Turkey (Mill, 1984;Mathew, 1985). In Iraq, G. sisyrinchium is mostly found inmountain slopes and grassy steppic hills and plains mostcommonly in poorly-drained soils with good moisture availability(Mathew, 1985). It is more frequent in hilly and arid regions. InTurkey, it is more frequent in the European part of the country,Outer Anatolia, and Islands (Mill, 1984). In Iran, it has beenreported from the Khuzestan plain (SW), Gorgan (NE), the Zagros

Fig. 3. Histograms showing the pollen percentages of four examined coprolite samples.

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Mountains including the vicinity of Kermanshah which is not toofar from the Wezmeh Cave. It is more frequently found in semi-arid areas but in habitats with high soil moisture. Floweringtime is in the early spring.

van Zeist and Bottema reported a pollen morphological typedescribed as “unidentified type” from Lake Zeribar sediment cores(1977, Fig. 17, page 51). These pollen grains perfectly correspond toTulipa pollen (T. agenensis-type). Tulipa species show a range ofmorphological variations (e.g. Reille, 1998; Kosenko, 1999) and it ishard to relate the studied tulippollen to any species in Iranbecauseofthe lack of the reference materials in IMEP and because the authorsfound no paper treating the pollen morphology of the genus in Iran.Tulipa is most commonly associated with Irano-Turanian mountainsteppes with grasses, thorn-cushions, and shrubs as well as theArtemisia steppes (e.g. Akhani, 1998). The flowering season of thetulip species is from early to middle spring. In Lake Urmia longrecords, the Tulipa pollen also shows a sporadic presence but seemsto bemore frequent during theglacial periods. Regarding the ecologyand flowering time of both Tulipa and Gynandriris, their significantpollen percentages in the sample WC-4 as well as the Urmia pollenrecords suggest that the coprolites may provide some informationabout the seasonal occurrences of specific plant species. The prob-able effects of seasonality on the pollen content and composition ofcoprolites have also been suggested by some authors (e.g. Scott et al.,2003; Carrión et al., 2006). The WC-4 sample was most probablydeposited during the early to middle spring time when soils onmountain slopes were recharged with water from the snow meltand/or the early spring precipitation.

Perhaps the most interesting feature of the sample WC-7 is thesignificant values of the Cousinia pollen. It is also present in low butsignificant values in three other examined coprolites (Fig. 3).Cousinia is an asteraceous giant genus with characteristic spinelesspollen morphology with more than 650 species whose distribution

area is almost completely limited to the Irano-Turanian phytogeo-graphical region (Knapp, 1987). It is the only Irano-Turanian biggenuswhich is relativelywell recorded in the pollen diagrams of thecontinental inland of the Middle East (e.g. van Zeist and Bottema,1977; Djamali et al., 2008a). The other genera, particularlyAstragalus (Fabaceae) and, Onobrychis (Fabaceae), Acantholimon(Plumbaginaceae), and Acanthophyllum (Caryophyllaceae), areextremely under-represented in the modern pollen rain (Djamali,2008). These facts highlight the importance of Cousinia pollenas one of the rare sources of information in understanding thechanges in the composition of the Irano-Turanian flora duringglacialeinterglacial cycles.

In fact, Cousinia is also an under-represented plant in themodern pollen rain from surface samples of the Middle East (e.g.van Zeist et al., 1970; Bottema and Barkoudah, 1979; Moore andStevenson, 1982). The high values of Cousinia pollen found in theWezmeh Cave coprolites especially in WC-7 strongly suggest thatthis plant was an important element in glacial landscapes of theZagros Mountains. It also implies that the composition of the Irano-Turanian glacial landscape was quite different from the simplifiedpicture of Artemisia or grass steppes commonly suggested by manyauthors (van Zeist, 1967; Djamali et al., 2008a). Cousinia and otherIrano-Turanian plant taxa probably dominated themountain slopesand well-drained plains of the Irano-Turanian region below thesnowline during the glacial periods. The floristic composition ofvegetation would have been similar to the subalpine (Alti-Irano-Turanian) vegetation belt of the N Iranian mountains dominated bythorny cushion species and grasses (Klein, 2001).

7. Conclusions

As coprolites give a more detailed picture of the plant speciespresent in the landscape at a local scale compared with large lakes,

Fig. 4. Pie diagrams to show the percentages of a selection of more frequent pollen taxa counted in four studied coprolites.

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they can provide additional information about the floristiccomposition of vegetation. Thus, in the Irano-Turanian regionwhere a major part of the flora is very under-represented in thepollen rain, they could complement the data obtained fromthe traditional pollen analysis of sediment cores. This is because thecoprolites, through the diverse activities of hyenas, trap the pollenof the insect- and self-pollinated plants near their productionsources, before their concentrations decline or they are destroyedon their way to depositional lacustrine basins. Our study showedthat coprolite pollen analysis is very useful to detect the plant taxapresent in the palaeolandscapes of the Zagros Mountains inW Iran.In the case of the Wezmeh Cave, the lack of information about thestratigraphical position of the studied samples due to the looting,hampered the establishment of chronostratigraphical pollenzonation. However, our study showed that coprolite pollen analysisis a very informative tool to understand the palaeovegetation andpalaeoflora of this area. In the arid to semi-arid climate of thisvast territory, where the suitable lakes and peat bogs for pollenanalysis are scarce or where packrat “rodent” middens have notbeen reported, coprolites provide an alternative source for palaeo-botanical investigations. Recent well-organized scientific excava-tions in several sites of Iran provide new potential for future polleninvestigations of coprolites.

Our study also shows that the glacial steppes of low-latitudemountain systems of the Middle East had a rich floristic composi-tion and that the difference between the glacial and interglacialvegetation is more characterized by the arrival and establishmentof arboreal taxa in warmer and wetter interglacials. From anevolutionary point of view, this implies that the Irano-Turaniansteppes may have been the centres of plant speciation during theglacial intervals of the Quaternary period that have been longenough to allow the reproductive barriers to form in topographi-cally isolated plant populations. Less glaciated Irano-Turanianmountains compared to higher-latitude mountain systems suchas the Alps, also could have reduced the number of extinctionevents. This may explain the extremely high speciation and ende-mism of the Irano-Turanian large genera such as Astragalus,Cousinia, and Acantholimon among many others. If this hypothesisis true, then an important loss of the Irano-Turanian plant biodi-versity can be predicted in the face of the future global warmingparticularly impacting the cold-adapted high-altitude species.

Acknowledgements

We are grateful to two anonymous reviewers of this manuscriptfor their thoughtful and constructive comments that helped usenhance the quality of this paper and gave us new insights intocoprolite pollen analysis. We also wish to thank Prof. HosseinAkhani, University of Tehran, for providing us with some infor-mation on plant ecology of some taxa.

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