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Zelkova carpinifolia (Pall.) C. Koch (Ulmaceae) in Turkey (Relict Tree): Floristics, Ecology, Distrubition and Threats

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Zelkova spach. (an ancient tree) is a member of the Ulmaceae family. Zelkova of about six species (Zelkova sicula Di Pasq., Garfì & Quézel, Zelkova abelicea (Lam.) Boiss., Zelkova carpinifolia (Pall.) C. Koch, Zelkova serrata (Thunb) Makino, Zelkova schneideriana Hand.-Mazz. and Zelkova sinica C.K. Schneid.) of deciduous, semi-evergreen trees occurring in written in woodland, thickets and in Italy, Greece, Turkey, Iran and Asia. Zelkova have alternate, ovate to elliptic toothed leaves. Most species display good fall colour of yellows and orange-browns. Zelkova carpinifolia, known as Caucasian elm, Caucasian zelkova or just zelkova is a species of Zelkova, native to the Caucasus, Kaçkar, and Alborz mountains in the extreme southeast of Europe and southwest Asia (Turkey, Azerbaijan, Iran, Georgia, Armenia). Zelkova carpinifolia is highly valued as an ornamental owing to its unusual outline, having a relatively short, wide trunk that divides into many ascending branches. It is rare in many regions and listed as Near Threatened according to IUCN Red List criteria. In Turkey, for example, only a few highly isolated populations are known (Kars, Hakkâri, Siirt, Muş). It seems very probable that other yet undiscovered sites with Z. carpinifolia still exist on Turkey, therefore additional field surveys should be undertaken. Currently there is no legal protection for this species. In this study, the genus of Zelkova, floristic and ecological characteristics were studied. Distribution areas in the world and Turkey is shown on the map and threat categoriesis based on the IUCN Red List criteria. In addition, pictures and images of the Zelkova carpinifolia species is given. Photos are taken in Turkey's Hakkari province. Türkiye'de Zelkova carpinifolia (Pall.) C. Koch (Ulmaceae) (Relikt Ağacı): floristik, ekoloji, dağılım ve tehditler Özet Zelkova spach. (relikt bir ağaç), Ulmaceae familyasının bir üyesidir. Ormanlık ve çalılıklarda yaprak döken ve dökmeyen Zelkova'nın İtalya, Yunanistan, Türkiye, İran ve Asya'da tanımlanmış yaklaşık altı türü vardır (Zelkova sicula Di Pasq., Garfì & Quézel, Zelkova abelicea (Lam.) Boiss., Zelkova carpinifolia (Pall.) C. Koch, Zelkova serrata (Thunb) Makino, Zelkova schneideriana Hand.-Mazz. and Zelkova sinica C.K. Schneid.). Zelkova alternat, ovat-eliptik ve dişli yapraklıdır. Çoğu türü sarı ve turuncu-kahverengi, sonbahar renklerindedirler. Kafkas karaağacı, Kafkas zelkovası ya da sadece zelkova olarak bilinen Zelkova carpinifolia, Kafkasya, Kaçkar, Avrupa ve güneybatı Asya (Türkiye, Azerbaycan, İran, Gürcistan, Ermenistan) güneydoğunun en uçlarında Elburz dağlarının doğal bir Zelkova türüdür. Zelkova carpinifolia çok sayıda yükselen dallara ayrılan nispeten kısa, geniş gövdeli; sıra dışı görüntüsü sayesinde çok iyi bir süs bitkisi olarak değerlendirilir. Pek çok bölgede nadir bulunur ve IUCN Kırmızı Liste kriterlerine göre NT olarak listelenir. Türkiye'de izole halde birkaç populasyonu bilinmektedir (Kars, Hakkâri, Siirt, Muş). Türkiye'de Z. carpinifolia'nın mevcut olduğu henüz keşfedilmemiş yerler hala çok muhtemel görünmektedir, bu nedenle arazi araştırmaları yapılmalıdır. Şu anda bu tür için yasal koruma bulunmaktadır. Bu çalışmada Zelkova cinsinin floristik ve ekolojik karakterleri araştırılmıştır. Türkiye ve Dünya'da yayılışı, harita üzerinde ve tehlike kategorilerisi IUCN Kırmızı Liste kriterlerine göre gösterilmiştir. Buna ek olarak Zelkova carpinifolia türüne ait şekiller ve çizimler verilmiştir. Fotoğraflar Türkiye'nin Hakkari ilinde çekilmiştir.
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
Zelkova carpinifolia (Pall.) C. Koch (Ulmaceae) in Turkey (Relict
Tree): Floristics, Ecology, Distrubition and Threats
*Fatmagül GEVEN1, Nezaket ADIGÜZEL2
1Ankara University, Faculty of Science, Ankara / TURKEY
2Gazi University, Faculty of Science, Ankara / TURKEY
*Corresponding Author: geven@science.ankara.edu.tr
Abstract
Zelkova spach. (an ancient tree) is a member of the Ulmaceae family. Zelkova of about six species
(Zelkova sicula Di Pasq., Garfì & Quézel, Zelkova abelicea (Lam.) Boiss., Zelkova carpinifolia (Pall.) C.
Koch, Zelkova serrata (Thunb) Makino, Zelkova schneideriana Hand.-Mazz. and Zelkova sinica C.K.
Schneid.) of deciduous, semi-evergreen trees occurring in written in woodland, thickets and in Italy,
Greece, Turkey, Iran and Asia. Zelkova have alternate, ovate to elliptic toothed leaves. Most species
display good fall colour of yellows and orange-browns. Zelkova carpinifolia, known as Caucasian
elm, Caucasian zelkova or just zelkova is a species of Zelkova, native to the Caucasus, Kaçkar,
and Alborz mountains in the extreme southeast of Europe and southwest Asia (Turkey, Azerbaijan, Iran,
Georgia, Armenia). Zelkova carpinifolia is highly valued as an ornamental owing to its unusual outline,
having a relatively short, wide trunk that divides into many ascending branches. It is rare in many regions
and listed as Near Threatened according to IUCN Red List criteria. In Turkey, for example, only a few
highly isolated populations are known (Kars, Hakkâri, Siirt, Muş). It seems very probable that other yet
undiscovered sites with Z. carpinifolia still exist on Turkey, therefore additional field surveys should be
undertaken. Currently there is no legal protection for this species.
In this study, the genus of Zelkova, floristic and ecological characteristics were studied. Distribution
areas in the world and Turkey is shown on the map and threat categoriesis based on the IUCN Red List
criteria. In addition, pictures and images of the Zelkova carpinifolia species is given. Photos are taken in
Turkey's Hakkari province.
Keywords: Zelkova carpinifolia, Caucasian elm, Ulmaceae, Turkey
Türkiye'de Zelkova carpinifolia (Pall.) C. Koch (Ulmaceae) (Relikt Ağacı): floristik,
ekoloji, dağılım ve tehditler
Özet
Zelkova spach. (relikt bir ağaç), Ulmaceae familyasının bir üyesidir. Ormanlık ve çalılıklarda yaprak
döken ve dökmeyen Zelkova’nın İtalya, Yunanistan, Türkiye, İran ve Asya'da tanımlanmış yaklaşık altı
türü vardır (Zelkova sicula Di Pasq., Garfì & Quézel, Zelkova abelicea (Lam.) Boiss., Zelkova
carpinifolia (Pall.) C. Koch, Zelkova serrata (Thunb) Makino, Zelkova schneideriana Hand.-Mazz. and
Zelkova sinica C.K. Schneid.). Zelkova alternat, ovat-eliptik ve dişli yapraklıdır. Çoğu türü sarı ve
turuncu-kahverengi, sonbahar renklerindedirler. Kafkas karaağacı, Kafkas zelkovası ya da sadece zelkova
olarak bilinen Zelkova carpinifolia, Kafkasya, Kaçkar, Avrupa ve güneybatı Asya (Türkiye, Azerbaycan,
İran, Gürcistan, Ermenistan) güneydoğunun en uçlarında Elburz dağlarının doğal bir Zelkova türüdür.
Zelkova carpinifolia çok sayıda yükselen dallara ayrılan nispeten kısa, geniş gövdeli; sıra dışı görüntü
sayesinde çok iyi bir süs bitkisi olarak değerlendirilir. Pek çok bölgede nadir bulunur ve IUCN Kırmızı
Liste kriterlerine göre NT olarak listelenir. Türkiye’de izole halde birkaç populasyonu bilinmektedir
(Kars, Hakkâri, Siirt, Muş). Türkiye'de Z. carpinifolia’nın mevcut olduğu henüz keşfedilmemiş yerler
hala çok muhtemel görünmektedir, bu nedenle arazi araştırmaları yapılmalıdır. Şu anda bu tür için yasal
koruma bulunmaktadır.
Bu çalışmada Zelkova cinsinin floristik ve ekolojik karakterleri araştırılmıştır. Türkiye ve Dünya’da
yayılışı, harita üzerinde ve tehlike kategorilerisi IUCN Kırmızı Liste kriterlerine göre gösterilmiştir. Buna
ek olarak Zelkova carpinifolia türüne ait şekiller ve çizimler verilmiştir. Fotoğraflar Türkiye’nin Hakkari
ilinde çekilmiştir.
Anahtar Kelimeler: Zelkova carpinifolia, Kafkas karaağacı, Ulmaceae, Türkiye
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
Introduction
Zelkova (Ulmaceae s.s.) is a small
genus that comprises six monoecious tree
species in the northern hemisphere. At
present, it shows a disjunct distribution in
east Asia (three species), western Asia
(one species) and the Mediterranean (two
species), whereas it is absent from North
America (Denk and Grimm 2005).
During past geological epochs, Zelkova
trees were widespread in Europe, Eurasia
and North Africa They were present in
many parts of Europe from the Oligocene
until the terminal Pliocene, in Italy until
approximately 32000 14C yr BP, and
have modern populations in southeastern
Sicily (Zelkova sicula), Crete (Zelkova
abelicea), Iran and Caucasus (Zelkova
carpinifolia), and Eastern Asia (Zelkova
schniederiana, Zelkova serrata, and
Zelkova sinica (Kvavadze and Connor
2005). Zelkova carpinifolia a Tertiary
relict tree endemic of Turkey. The genus
has a particularly disjunct distribution
(Güner et al. 2000). Zelkova carpinifolia
is one of the most iconic relict trees in the
Transcaucasus, between the Black and
Caspian Sea. The generally warm and
humid environmental conditions in this
region since the late Tertiary provided a
shelter for relict species during the ice
ages. As a result, the Transcaucasian
forests, especially those in the Colchis
and Hyrcanian regions, are among the
oldest, most diverse and richest habitats
housing endemic, woody species in west
Eurasia (URL1 2016).
The purpose of this paper is to review
and evaluate the ecological features and
distribution area of the Zelkova
carpinifolia.
Material and Methods
We collected distribution, threat and
population structure data by reviewing
literature and through field surveys at
two study plots throughout the range of
the species. It has also benefited from
various herbarium (ANK., GAZI.,
VAN.). The geographical distributions of
Zelkova carpinifolia was analyzed using
Grid system (Davis 1965- 1988), (Figure
1). In addition, photos were taken (Figure
2, Figure 4).
Research in the area of Şemdinli and
Yüksekova (Altısu village, 1500m.).
There is a 15. km long old road.
Figure 1. Distribution map of Zelkova
carpinifolia (Pall.) K. Koch
Results and Discussion
Taxonomy
Kingdom: Plantae
Phylum: Tracheophyta
Class: Magnoliopsida
Order: Urticales
Family: Ulmaceae; Ulmaceae,
the elm family of the nettle order
(Urticales), with 15 genera of trees and
shrubs, distributed primarily throughout
temperate regions. Members of the
family have watery sap, and its leaves
alternate along the stem. The leaves
usually have toothed edges and often are
lopsided at the base. The flowers lack
petals. Male and female flowers are
borne together or apart on the same plant.
The fruit, a samara, may be winged,
fleshy, or nutlike (Zieliński 1979, UR L2
2016).
Genus: Zelkova Spach; Zelkova is a
genus of six species of deciduous trees in
the elm family Ulmaceae, native to
southern Europe, and southwest and
eastern Asia. They vary in size from
shrubs (Z. sicula) to large trees up to 35
m tall (Z. carpinifolia). The leaves are
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
alternate, with serrated margins, and
(unlike the related elms) a symmetrical
base to the leaf blade. The fruit is a dry,
nut-like drupe, produced singly in the
leaf axils (UR L3 2016).
Species: Zelkova carpinifolia (Pall.) K.
Koch; Western Asia (Figure 2, Figure 3).
Figure 2. Zelkova carpinifolia (Pall.) K. Koch
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
Figure 3. Zelkova carpinifolia (Pall.) K.
Koch
Synonym for Zelkova carpinifolia (Pall.)
K. Koch;
1. Abelicea crenata (Desf.) Baill. species,
2. Abelicea sibirica Rafin. species,
3. Abelicea ulmoides Kuntze species,
4. Planera carpinifolia P.Watson species,
5. Planera crenata Desf. species,
6. Planera repens Hort. ex
Lavallee species,
7. Rhamnus carpinifolius Pall. species,
8. Ulmus nemoralis Ait. species,
9. Ulmus polygama Juss ex Poir. species,
10. Zelkova carpinifolia subsp.
yomraensis Ansin & Gerçek infraspecific
name,
11. Zelkova crenata (Desf.) Sp. species,
12. Zelkova crenata var. repens
Lavallee infraspecific name (URL3 2016)
and Zelkova ulmoides Schneid. (Denk
and Grimm 2005). Synonym of other
species and distribution areas (names in
parentheses are synonyms);
-Zelkova schneideriana Hand.-Mazz.,
China
-Zelkova sinica C. K. Schneider, China
-Zelkova serrata (Thunb.) Makino,
Japan, (Zelkova formosana Hayata),
(Zelkova keaki Maxim.), (Zelkova.
acuminata Planch.)
-Zelkova abelicea (Lam.) Boiss., Crete,
(Zelkova cretica Spach.)
-Zelkova sicula Di Pasquale, Garfi &
Quezel Sicily, (Denk and Grimm 2005).
Distribution: Turkey (E-Anatolia, SE-
Anatolia), Iran (EC-Iran, E-Iran, NE-
Iran: Mts., N-Iran: Mts., NW-Iran:
Iranian Aserbaijan), Caucasus,
Azerbaijan, Armenia, Gruziya, Crimea
(introduced) (Güner and Zielinski 1998,
URL3 2016). The geographical
distributions of Zelkova carpinifolia
was analyzed using Grid system (Kars
A9, Hakkari B8, Siirt C9, Muş C10),
(Figure 1), (Davis 1965-1988, Davis and
Tan 1988, Güner et al. 2000).
Habit: Z. carpinifolia is a large tree,
attaining an average height of 20-35 m,
and measuring up to 2 m in diameter. The
vase-shaped crown and short, broad trunk
dividing into numerous nearly erect,
strong branches, give the species its
characteristic and distinctive form. The
leaves, especially on young shoots, are
generally much larger than those of the
Mediterranean Zelkova species: they can
measure more than 10 cm in length and 6
cm in width, with a dentate leaf base, and
7-12 secondary veins (Kozlowski and
Gratzfeld 2013). Z. carpinifolia is
observed in the region of 2-3 meters tall
and shrubs form (Figure 4). Whatever the
causes of this fragmentation (a complex
colonization history, climate change,
browsing by large Pleistocene herbivores
or more recent agricultural practices),
gene flow between remote populations is
improbable because of the limited
dispersal capacity of seeds, as is the case
for all Zelkova species (Kozlowski et al.
2013).
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
Figure 4. Habit of Zelkova carpinifolia (Pall.) K. Koch
Ecology: Generally favouring moist,
humus-rich soils but not tolerating
waterlogged or swampy conditions,
Zelkova carpinifolia is a light-demanding
canopy tree. Observation area; generally
mountainous area (1300-1800 m), 45%
of forestland, unfit for agriculture 41%,
10% meadows and pastures can be used
four or five months and 4% are arable
lands. Genus of the tree in the forest is
usually oak. Quercus brantii Lindl. and
Quercus cerris L. are the dominant plant
species of oak forest (Figure 5).
Figure 5. Habitat of Zelkova carpinifolia (Pall.) K. Koch
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
Climate: Hakkâri is seen in cold and
temperate climates. much more rainfall in
the summer months during the winter.
According to the Köppen-Geiger climate
classification it can be called Ds. The
average annual temperature is 10.3
province of Hakkari. The average annual
rainfall: 789 mm (Figure 6, Figure 7,
Table 1), (UR L4 2016).
Figure 6. Graphic of precipitation (mm)
and temperature (°C, °F)
24.2 temperature is the hottest month of
the year in July. The average temperature
in January is the lowest average of the
year is -3.8.
Figure 7. Graphic of temperature (°C, °F)
Table 1. Table of precipitation (mm) and temperature (°C, °F)
month
1 2 3 4 5 6 7 8 9 10 11 12
mm
37 36 44 59 53 21 5 4 12 50 51 37
°C
-3.7 -3.0 1.5 7.5 12.4 17.1 21.2 20.9 16.9 10.9 4.9 -0.3
°C (min)
-8.2 -7.7 -3.0 2.6 6.8 10.5 14.4 13.9 10.0 4.9 0.0 -4.4
°C (max)
0.8 1.8 6.0 12.4 18.0 23.7 28.1 28.0 23.9 16.9 9.8 3.9
Driest and wettest month of the year
rainfall between 128 mm average
temperature throughout the year varies
around 28.0. (URL4, 2016).
Red List Category & Criteria:
Lower Risk/near threatened (Figure 8).
Occurring in lowland as well as mountain
forests across the Caucasus, including
two main population centres in the east
(Hyrcania) and the west (Colchis), Z.
carpinifolia has the widest distributional
range (some 100 populations) of the three
Eurasian species. Molecular studies
revealed a significant genetic
differentiation between the two main
groups which is indicative of the many
million years gone by since these
populations fragmented into their current
distribution. By and large, in all regions
where the species occurs, overgrazing,
expanding mass tourism and related
infrastructure development in areas of
scenic beauty, are major drivers of
change. Although numerous natural
reserves and parks have been created in
Georgia, Azerbaijan and Iran, containing
important forest stands with Z.
carpinifolia, the pressure on these
habitats is steadily rising (Güner and
Zielinski 1998, Kozlowski and Gratzfeld
2013, URL3 2016).
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International Forestry Symposium (IFS 2016) Proceedings
07-10 December 2016, Kastamonu/TURKEY
Figure 8. Conservation status of Zelkova carpinifolia (Pall.) K. Koch.
Diseas: Zelkova carpinifolia (Pall.) K.
Koch is native to the Caucasus and
Alborz mountains in northern parts of
Iran. The decline began with browning of
the leaves, viscous liquid exudatation on
the branches and trunks resulting in a
brown-black discoloration of bark and
woody tissues. In the winter of the next
year fungal growth induces a typical
charcoal-black surface on diseased
branches and trunks (Mirabolfathy 2013).
Zelkova carpinifolia (Pall.) K. Koch is
described as a new observations for the
Hakkari region (B8) of Turkey. A
detailed morphological description,
photographs, distribution map and
illusturation of this new observations are
given. Several population was observed
in case of local communities in the
region. Overgrazing, fire, erosion,
agriculture, road construction, as a result
of activities have been identified as
fragmented habitats (Figure 9).
Figure 9. Fragmented habitats of the
Zelkova carpinifolia
Conclusion
Fires, overgrazing, tourism,
industrialization, agricultural struggle,
expansion of agriculture, urbanization
and plantation crops such factors are
threatening rare plants. In addition, by
the local people medicine, fuel,
unconsciously gathering for various
purposes such as food and ornamental
plants are also threatening a factor. These
threats are not the generation of habitat
fragmentation and species. It should
therefore take the necessary protection
measures. We can also learn a lot from
relict plant; Tertiary relict trees are,
therefore, ancient organisms which were
able to cope with changing
environmental conditions for millions of
years. They provide, thus, a unique
opportunity to understand past and recent
biogeographical and evolutionary
processes (Taberlet et al. 1998).
References
Davis, P.H., 1965-1985. Flora of Turkey and
the East Aegean Islands, Vol I-IX, Edinburgh
University Press, Edinburgh.
Davis, P.H., Mill, R.R., Tan, K., 1988. Flora
of Turkey and the East Aegean Islands
(Supplement), Vol 10, Edinburgh University
Press, Edinburgh.
Denk T, Grimm GW. 2005. Phylogeny and
biogeography of Zelkova (Ulmaceae sensu
stricto) as inferred from leaf morphology, ITS
sequence data and the fossil record. Botanical
Journal of the Linnean Society 147: 129-157.
Güner, A., Özhatay, N., Ekim, T., Başer,
K.H.C., 2000. Flora of Turkey and the East
Aegean Islands (Supplement 2), Vol 11,
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Güner, A. & Zielinski, J. 1998. Zelkova
carpinifolia. The IUCN Red List of Threatened
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Kozlowski G., Gibbs D., Huan F., Frey D.,
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Article
Zelkova carpinifolia (Pall.) C. Koch is an ancient tree of the Ulmaceae family, which is considered an essential species in the Hyrcanian forests of Iran. Therefore, we examined infraspecific genetic variation and morphological polymorphism in ten Iranian populations of this species. For the molecular study, nuclear genomes of 50 individuals of 10 populations were extracted by C-TAB modified protocol and amplified using 10 SCoT molecular primers. Additionally, 10 individuals were selected per population for morphometric analyses. We used SPSS, GenAlex, PopGene, GenoDive, and Structure software for statistical analyses. All quantitative morphological characteristics differed among and within the populations. Meanwhile, the ANOVA test showed a significant difference (P ≤ 0.05) for a few traits. Significant correlations were determined between some morphological traits with geographic parameters of habitats. No significant correlation was detected between morphological variation and geographic distance by the Mantel test. However, some qualitative features such as leaves margin, basal forms, and blade shape differed among and within the populations. The UPGMA dendrogram revealed a high level of morphological polymorphism within populations, followed by a low level of among populations morphological differentiation. However, we detected four morphotype groups among the evaluated populations. Results of the AMOVA test indicated a significant proportion of genetic diversity among the populations. The large amounts of HT and GST showed a strong genetic differentiation created by a flat rate of gene flow. Moreover, isolation by distance occurred among the populations. TCS networking indicated a low rate of mutation occurred among the individuals of most populations. According to UPGMA tree and structure analyses, two main groups of genotype were detected that were in accordance with geographical distribution of the populations. In addition, a large amount genetic dissimilarity was detected between some populations. The genetic structure of both genotypes was nearly uniform; however, some intermediated were observed. We observed some significant correlations between various genetic diversity parameters with some leaf morphological features, which indicates genetic base of these variations. For some populations, the clustering patterns were similar for both, morphology and genetic diversity, studies. The results of morphological study did not support the existence of subspecies in Z. carpinifolia, while results of molecular study were entirely different. Our findings revealed that genetic isolation by distance, a flat rate of gene flow, a low mutation level, genetic drift, and vegetative reproduction lead to a substantial population genetic differentiation, followed by a small amount of within-population genetic diversity. However, the relative uniformity in environmental condition resulted in similar phenotypic plasticity between some populations, which was supported by correlation analysis.
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Maintaining living ex situ collections is one of the key conservation methods in botanic gardens worldwide. Despite of the existence of many other conservation approa-ches used nowadays, it offers for many endangered plants an important insurance policy for the future, especially for rare and threatened relict trees. The aim of this research was to investigate the global extent of living ex situ collections, to assess and discuss their viability and inform the development of conservation approaches that respond to latest global conservation challenges. We used as a model taxon the tree genus Zelkova (Ulm-aceae). The genus includes six prominent Tertiary relict trees which survived the last glaciation in disjunct and isolated refugial regions. Our comprehensive worldwide survey shows that the majority of botanic institutions with Zelkova collections are in countries with a strong horticultural tradition and not in locations of their origin. More importantly, the acutely threatened Zelkova species are not the most represented in collections, and thus safeguarded through ex situ conservation. Less than 20% of the ex situ collections surveyed
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Zelkova pollen has been found in Oligocene- to Pleistocene-aged deposits from many parts of Europe and northern Africa, but became extinct in mainland Europe prior to the last glacial maximum. This paper presents some observations on the ecology, pollen productivity and Holocene history of Zelkova carpinifolia to further understanding of Quaternary climatic trends. Georgia is one of the last refuges of this Tertiary relict tree.Based on palynological data from 20 Holocene sediment profiles in Georgia, we have established that Zelkova pollen is almost always accompanied by elevated proportions of thermophilous taxa (Castanea sativa, Quercus hartwissiana, Quercus iberica, and Pterocarya pterocarpa) in pollen spectra. These spectra are associated with phases of climatic amelioration and humidification. Zelkova carpinifolia is characterised by low pollen productivity and is underrepresented in pollen spectra by four-to-five orders of magnitude. Because of this, even single grains of Zelkova pollen may play a significant role in pollen-based climatic reconstructions.Six major climatic optima occurred in Georgia through the course of the Holocene, the longest and warmest of which occurred in the mid-Holocene and reached its maximum between 6000 and 5500 Cal. yr BP. During that period, Zelkova and Castanea forests were widespread. In Western Georgia, the upper tree line was elevated by as much as 300 m above its present-day level. In semiarid Eastern Georgia, the tree line may have been 500–600 m higher. Other climatic optima are seen in late-Holocene pollen spectra dating to 3800–2500 Cal. yr BP and 1350–800 Cal. yr BP.
Outbreak Of Charcoal Disease On Quercus spp. and Zelkova carpinifolia Trees In Forests of Zagros And Alborz Mountaıns In Iran
  • M Mirabolfathy
Mirabolfathy M. 2013. Outbreak Of Charcoal Disease On Quercus spp. and Zelkova carpinifolia Trees In Forests of Zagros And Alborz Mountaıns In Iran, Iran. J. Plant Path., Vol. 49, No. 2,: 77-79. URL1, 2016.
  • P H Davis
Davis, P.H., 1965-1985. Flora of Turkey and the East Aegean Islands, Vol I-IX, Edinburgh University Press, Edinburgh.
Phylogeny and biogeography of Zelkova (Ulmaceae sensu stricto) as inferred from leaf morphology, ITS sequence data and the fossil record
  • P H Davis
  • R R Mill
  • K Tan
Davis, P.H., Mill, R.R., Tan, K., 1988. Flora of Turkey and the East Aegean Islands (Supplement), Vol 10, Edinburgh University Press, Edinburgh. Denk T, Grimm GW. 2005. Phylogeny and biogeography of Zelkova (Ulmaceae sensu stricto) as inferred from leaf morphology, ITS sequence data and the fossil record. Botanical Journal of the Linnean Society 147: 129-157.