Plant Diversity in the Central Great Caucasus: A Quantitative Assessment

175 Pages·2017·4.73 MB·English

by  George Nakhutsrishvili

Geobotany Studies
Basics, Methods and Case Studies
George Nakhutsrishvili
Otar Abdaladze
Ketevan Batsatsashvili
Eva Spehn
Christian Körner E ditors
Plant Diversity
in the Central
Great Caucasus:
A Quantitative
Assessment
Geobotany Studies
Basics, Methods and Case Studies
Editor
FrancoPedrotti
UniversityofCamerino
ViaPontoni5
62032Camerino
Italy
EditorialBoard:
S.Bartha,Va´cra´to´t,Hungary
F.Bioret,UniversityofBrest,France
E.O.Box,UniversityofGeorgia,Athens,Georgia,USA
A.Cˇarni,SlovenianAcademyofSciences,Ljubljana,Slovenia
K.Fujiwara,YokohamaCityUniversity,Japan
D.Gafta,“Babes-Bolyai”UniversityCluj-Napoca,Romania
J.Loidi,UniversityofBilbao,Spain
L.Mucina,TheUniversityofWesternAustralia,Perth,Australia
S.Pignatti,Universita` degliStudidiRoma“LaSapienza”,Italy
R.Pott,UniversityofHannover,Germany
A.Vela´zquez,CentrodeInvestigacio´nenScie´nciasAmbientales,
Morelia,Mexico
R.Venanzoni,UniversityofPerugia,Italy
About the Series
The series includes outstanding monographs and collections of papers on a given
topic in the following fields: Phytogeography, Phytosociology, Plant Community
Ecology,Biocoenology,VegetationScience,Eco-informatics,LandscapeEcology,
Vegetation Mapping, Plant Conservation Biology and Plant Diversity.
Contributions are expected to reflect the latest theoretical and methodological
developments or to present new applications at broad spatial or temporal scales
that could reinforce our understanding of ecological processes acting at the
phytocoenosis and landscape level. Case studies based on large data sets are also
considered,providedthattheysupportrefinementofhabitatclassification,conser-
vation of plant diversity, or predictionof vegetation change. “Geobotany Studies:
Basics, Methods and Case Studies” is the successor to the journal Braun-
Blanquetia, which was published by the University of Camerino between 1984
and 2011 with the cooperation of the Station Internationale de Phytosociologie
(Bailleul-France) and the Dipartimento di Botanica ed Ecologia (Universita` di
Camerino, Italy) and under the aegis of the Socie´te´ Amicale Francophone de
Phytosociologie, the Socie´te´ Franc¸aise de Phytosociologie, the Rheinold Tüxen
Gesellschaft, andthe Eastern AlpineandDinaricSociety forVegetation Ecology.
Thisseriesaimstopromotetheexpansion,evolutionandapplicationoftheinvalu-
ablescientificlegacyoftheBraun-Blanquetschool.
Moreinformationaboutthisseriesathttp://www.springer.com/series/10526
George Nakhutsrishvili (cid:129) Otar Abdaladze (cid:129)
Ketevan Batsatsashvili (cid:129) Eva Spehn (cid:129)
€
Christian Korner
Editors
Plant Diversity in the
Central Great Caucasus:
A Quantitative Assessment
Editors
GeorgeNakhutsrishvili OtarAbdaladze
InstituteofBotany InstituteofEcology
IliaStateUniversity IliaStateUniversity
Tbilisi,Georgia Tbilisi,Georgia
KetevanBatsatsashvili EvaSpehn
InstituteofEcology InstituteofPlantSciences
IliaStateUniversity UniversityofBern
Tbilisi,Georgia Bern,Switzerland
ChristianK€orner
InstituteofBotany
UniversityofBasel
Basel,Switzerland
ISSN2198-2562 ISSN2198-2570 (electronic)
GeobotanyStudies
ISBN978-3-319-55776-2 ISBN978-3-319-55777-9 (eBook)
DOI10.1007/978-3-319-55777-9
LibraryofCongressControlNumber:2017938577
#SpringerInternationalPublishingAG2017
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Foreword
TheGreatCaucasusiswellknownforitsspectacularflora.Arichtopographyand
geology, steep climatic gradients, and the position between the flora regions of
Asia,Europe,andtheMediterraneanhavecontributedtoevolutionofthisbiodiver-
sityhotspot.Intotal,6350vascularplantspeciesoccurintheCaucasus,including
more than 2900 endemic species. The Great Caucasus represents all elevational
vegetationbeltsfromsemidesertsoftheCaspianSeadepression((cid:1)28ma.s.l.)to
thenivalbeltoftheice-cappedMt.Elbrus(5642ma.s.l.).
This book is based on extensive botanical-ecological research in the Central
Great Caucasus, the Kazbegi region in particular. Community ecological field
surveys during the last 50–60 years have created knowledge and an extensive
vegetation database that is synthesized in this volume. Botanists have visited and
surveyedthefloraandvegetationofthatregionovermorethan200years.Mostof
theseearlierworkswerefacilitatedbytheancientNorth-Southroadwhichcrosses
the Central Great Caucasus at its eastern border. Among the first who traveled in
these mountains in 1770 was the naturalist J. A. Gueldenstaedt. Later, this region
was visited by numerous outstanding botanists, like M. F. Adams, A. A. Mussin-
Pushkin, J. J. F. W. Parrot, K. H. E. Koch, F. J. Ruprecht, G. F. R. Radde, N. A.
Desulavi,A.Rehman,andB.F.Busch,whomadevaluableherbariumcollections.
The first botanical survey that took a community approach was that by Seifriz
(1931)intheKazbegiregionintheearlytwentiethcentury.Detailedandsystematic
studies have begun only since 1928 when the prominent Georgian botanist
A. Kharadze pioneered this field. She collected rich herbarium material and
described many new species. She was assisted by the local resident botanist
E. Khutsishvili. These rich collections laid the foundation for a first synopsis of
thefloraofvascularplantsoftheKazbegiregion(SakhokiaandKhutsishvili1975).
AnumberofregionalassessmentsoffloraandvegetationofthispartoftheCentral
Great Caucasus were published by Kharadze (1944, 1948a, b, 1965), Kimeridze
(1965a, b), Sakhokia (1983), Nakhutsrishvili (1971, 1974, 1999, 2003, 2013),
Gamtsemlidze (1979), Nakhutsrishvili and Gamtsemlidze (1984), Bedoshvili
(1985), Nakhutsrishvili et al. (1990, 2005, 2006), Zazanashvili (1990), Kikvidze
and Nakhutsrishvili (1998), Zazanashvili et al. (2000), Shetekauri (1999),
Shetekaurietal.(2012),Tephnadzeetal.(2014),andAbdaladzeetal.(2015).
The Foundation of the “Kazbegi High-Mountain Research Station” in 1969
undertheleadershipofthewell-knownGeorgianBotanistNikoKetskoveliplayed
v
vi Foreword
a key role for the research activities in this area. Located at 1800 m a.s.l. at the
oustskirtsofthetownofKazbegi(nownamedStepantsminda),thisstationbecame
a focal point for decades of plant eclogical research. Soon, the Stepantsminda
station attracted researchers from all over the world and became an international
center for high-mountain research. In the 1970s, Prof. W. Larcher (University of
Innsbruck,Austria)introducedexperimentalresearchinhigh-mountainplantecol-
ogy at this station which led to works by the Austrian scholars Prof. A. Cernusca
and Prof. C. K€orner and the establishment of a young generation of Georgian
botanists under the leadership of Prof. Nakhutsrishvili (Gamkrelidze 1986;
Sanadiradze1986;Abdaladze1987;KikvidzeandAbdaladze1988).Morerecently,
Prof. C. K€orner and Dr. E. Spehn (University of Basel, Switzerland) facilitated
research projects on the functional significance of biodiversity in the Kazbegi
region.OneoutcomeoftheseworkswasaprojectsupportedbytheSwissNational
Science Foundation in cooperation with the Swiss Agency for Development and
Cooperation(SCOPESprogramme),theaimofwhichwasadigitalassessmentofa
40-yearseriesofphytosociologicalstudiesandcollectionofvegetationdatainthe
Kazbegiregion(inessencedigitizingG.Nakhutzsrishvilis’sfieldbooks),whichis
partofthisbook.ThedigitaldatacoverthecentralpartoftheGreatCaucasusrange,
specifically the geographical region of Kazbegi (1400–3700 m a.s.l.), Mamisoni
Pass(2750–3650ma.s.l.),andthesubnivalbelt(3000–3750ma.s.l.)ofMt.Elbrus.
Prof. E. Hübl, Prof. F. Ehrendorfer, Prof. M. Fischer, Prof. O. Hegg, and Prof.
A.OtteparticipatedinvegetationsurveysintheKazbegiregion.
Withthisvolume,wehopetoestablishareferenceforfuturebotanicalworksin
theGreatCaucasus.ThedatacollectedherealsorepresentaGeorgiancontribution
to regional assessments of Biodiversity inthe context of IPBES (the International
Program on Biodiversity and Ecosystem Services). We also aimed at drawing a
wider picture by comparisons with the European Alps. Hence, we also hope that
larger scale surveys and comparative works in evolutionary biology and ecology
willprofitfromthisassessment.Lastbutnotleast,thisvolumecompilesevidence
thatmayassistinconservationpolicyforthisfragilemountainworld.
Tbilisi GeorgeNakhutsrishvili
October2016 ChristianK€orner
Contents
1 AGeostatisticalandBioclimatologicalComparison
oftheCentralGreatCaucasusandtheCentralAlps. . . . . . . . . . . . 1
ChristianK€ornerandJensPaulsen
2 VegetationoftheCentralGreatCaucasusAlongW-EandN-S
Transects. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
GeorgeNakhutsrishviliandOtarAbdaladze
3 PlantDiversityoftheCentralGreatCaucasus. . . . . . . . . . . . . . . . . 17
GeorgeNakhutsrishviliandOtarAbdaladze
4 QuantitativeAnalysisofthePhytosociologicalReleve´s
fromtheCentralGreaterCaucasus. . . . . . . . . . . . . . . . . . . . . . . . . 113
GeorgeNakhutsrishviliandKetevanBatsatsashvili
5 AComparisonofClimaticNichesoftheSameAlpine
PlantSpeciesintheCentralCaucasusandtheAlps. . . . . . . . . . . . . 133
ChristopheF.Randin,Lo¨ıcPellissier,AntoineGuisan,
andGeorgeNakhutsrishvili
6 NewIndicatorValuesforCentralCaucasusFlora. . . . . . . . . . . . . . 145
GeorgeNakhutsrishvili,KetevanBatsatsashvili,KatrinRudmann-Maurer,
ChristianK€orner,andEvaSpehn
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
vii
1
A Geostatistical and Bioclimatological
Comparison of the Central Great Caucasus
and the Central Alps
€
Christian Korner and Jens Paulsen
1.1 Introduction
Althoughalmost3000kmapart,theGreatCaucasusattheeasternedgeofEurope
andtheAlpsincentralEuropeshareacommonyounggeologicalage,anapproxi-
mative W-E orientation, they both belong to the Eurasian mountain chain that
formed and is still forming as a result of southern continents pushing northwards.
Usinga1000melevationminimum,theGreatCaucasusisstretchingfrom41(cid:1) 150
N to 43(cid:1) 450 N (central part at 43(cid:1)N) and the Alps from 44(cid:1) 100 N to 47(cid:1) 400 N
(central part at 46(cid:1) 300N), with both chains belonging to the temperate zone
(Fig. 1.1). The Alps experience a stronger maritime and the Great Caucasus a
more continental influence. Both ranges divide the weather systems into northern
and southern climates, and both show strong precipitation gradients. In the Great
Caucasus this is a NW-SE gradient, in the Alps (with some exceptions) a N-S
gradient, ranging from around 2000 mm per year to less than 500 mm at places.
Both mountain systems show a mass elevation effect (‘massenerhebungseffekt’),
withahigherelevationofisothermsintheinteriorpartscomparedtofrontranges,
andadry,step-typeclimateinpartsoftheirdeepcentralvalleys.Yet,theCaucasus
formsasinglemaindividewithaseriesofsidevalleysoneitherside,whereasthe
Alpshaveseveralchainsinparallel,permittingamorepronouncedmasselevation
effect to occur in its interior valleys. The maximum elevation is similar, with the
highest peak of the Great Caucasus, Mount Elbrus 5642 m, and that of the Alps,
Mont Blanc 4809 m. Yet, some of the highest peaks in the Caucasus are former
volcanoes(MountKasbekwith5047m,isoneofthem,inthecorestudyregionof
thisvolume),whereasintheAlps,allsummitsaretectonicsummits.
C.K€orner(*)•J.Paulsen
InstituteofBotany,UniversityofBasel,Sch€onbeinstrasse6,4056Basel,Switzerland
e-mail:[email protected]
#SpringerInternationalPublishingAG2017 1
G.Nakhutsrishvilietal.(eds.),PlantDiversityintheCentralGreatCaucasus:
AQuantitativeAssessment,GeobotanyStudies,DOI10.1007/978-3-319-55777-9_1
2 C.Ko€rnerandJ.Paulsen
Fig.1.1 ThegeographicalpositionoftheGreatCaucasusandtheAlps.Thecross-sectionallines
indicatethepartthatisconsidered‘central’inbothmountainranges.Thecontentofthisvolumeis
focusingonthatcentralpartoftheGreatCaucasus
From a biogeographical point of view, these well separated mountain systems
are interesting, because both are continental centers of biodiversity, they share a
good deal (20–30%) of the same flora (the same or very closely related plant
species), but also differ in the origin of other parts of their flora, with the Alps
exhibiting a stronger influence of Arctic elements and the Caucasus showing a
stronger affiliation to central Asia (see Chap. 3). Although the mountains of the
BalkanandtheCarpatiansformabridgethatmayhavebeenstrongerduringglacial
periods,thegreatgeographicdistance,aswellasthemoderateelevationofmostof
theseintermittedmountains,andamissingmountainouscontinuum,causedahigh
degree of floristic separation. What happened to the widely separated, but partly
common flora in these two mountain systems over the presumably several recent
millions of years of isolation from each other? Which common traits have been
retained,whichgotlost?This‘experimentbynature’opensfascinatingterrainfor
evolutionarybiologists.
Here,weofferacomparative,ratherbasic,geophysicalassessmentofthenature
ofthesetwomountainsystems.Theanalysisshouldhelpbringingthegeobotanical
studies presented in this volume for the central Great Caucasus in perspective to
whatisknownfortheAlps,characterizetheelevationstructureandbiogeographic
beltsofthetwomountainsystemsfromaclimatologicalperspective,anddefinethe
geographical space this book is focussing on. We used the world topography and
the climate data base by WorldClim for driving a bioclimatic model that permits
stratifyingbiogeographicbeltsbyclimaticcriteria(K€orneretal.2011;Paulsenand
K€orner2014).
1.1.1 The Elevation Structure of the Great Caucasus and the Alps
The total area of the Caucasus and the Alps is 132,000 km2 versus 188,000 km2.
Thisareaisfortheentiremountainsystemeach,includingallinteriorlowelevation
terrain andthe hillslopes inthe periphery.This is the area aphysicalatlas would
show in brown (for higher) and yellow (for lower) mountain terrain. If a line is
drawn by best guess around these mountain territories (a mountain ‘polygon’),
thesearethenumbersforarealcoveragethatemerge.Wewilldiscusslaterhowthat
area could be separated into ‘true’ mountain terrain and other terrain. The total
mountain polygon area of these two mountains is thus, roughly 3 times and 4.5