Oak & Oak Forests in Caucasia
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OAIQ Al'D OAK FORESTS I CAL CASL\ Peter A. chmidt Profc,.;er, Dre..Oen CnJ\eNt~ of Technology Depanment ot Fore't Sc1ence,, ln,otute of General Ecolog) and FnHronment \fanagement D-01737 Tharandt. German) Introduction to the Cauca! lb ReWoo When reganlmg the .. Caucasu~:· some authol"> reh:r onl) to the h1gh mountam~ of the Great Caucasu\. Here the term "Cauca,iu'' '' 1ll be: u'ed to reler to the rnure area located bet\\.een the Black Sea and the Ca..,p1an Sea nl the Cauca'u' Reg1on . Be..,1de~ the Great Cauca~u~. the Nonh Cau~.:<hiUn Plain belonging to the Ru\\lan Federation, ~well ,L\ the depre\siOm. and lnwlanth. muumam rangcs and the h1ghland south of the Great Cauca~u' up tn t11e 'outhem border of the ..,t,•le' Armema. AserbaiJan. Georg1a <South Cauca,ta, often called ''Transcaucasia l are abo included. The Great Cauca'u' extend~ betv.een the Black Sea and Casp1an Sea o\er a t.l1..,tan~.:c ul I 500 km and a breadth of 30-180 km. It wn'1't' of sc\ eml parallel mountain nrnges. The mam ndge i~ preceded toward' the nonh by a 'crond ndge (here the Elbrus, the highe't peak 5633 m. and the Ka,bel..:. the h1ghN peak 5C.I-W ml. a~ well a' the rock) ndge Cch3Jll of bme..tone mas 1h). The e h1gh mountam' form an 1mpo1U11t chmate and "ater di' ide bet\\ een Ea't Europe and South\\e't Asia, and thu' hetween t\\O continent'. Geological de,eJopment, a' \\ell as thc unique omgraph1c :md climatic 'iruation, gi\es ri-.c to a unique naturo~l semng and b1olog1cal dJ\eNt)' of Cauca,ia. The cultural dt\ei">Jt) charaden,uc ol moun tamous reg1on' 1s not le<>s unique. Likewbe. the cultural hJstol') and land u'e dat.ing back for millennia have influenced the land,cape' and habnut-.. Characteri,tlc orobiomes rich in relics and endemicc, (from ne<~r- ,hore montane tore't... up t<' the alpme gra~~lands) meet withm J 'mall geographic area with submednenanean and "·arm-humid b10mes. as v\cll a~ cnnuncntal sernwrid biomes <ct. BIN 2CX>O. Zazanashvili et a1 2000. Krever et al. 2CX)J St·hmidt 2C>O:!J. In the nonh Caucas1an Plain steppe and woodland 'teppc d!cas. and m the ea.st, Caspian 'emide,ens clo-.cl) approach the C:tuca,us. The 'tcep 'outhern ~lopmg of the Great Cauc~u' contatns very different ccntope' and dJtlerent ph} togcographie pro' mces of ~outhem Cauca~•a : In th ''e't wuh Its hum1d climate. it tran,forms intll the coastal area ul the Blad.. Sea and the Lo" land of Colkheti. Ea't of the Sur..m ~tountam-.. 11 forrru. a bndge to \1mor Cauca'u' hl\\arU' the Kura um l..nd. extendmg. m the ea~t 10 the and Kura-Ara:o. l.o\\ land at the Casptan Sea To the south ascend thc mountainous ')'tern <Jf !\1mor Cauc:a.,us characten ed b) us dJ\e!"e, but human-mfluenced, montane \egetauon The ~mor Cauca,us lonm the margin of the Armeman llig.hlund It ., poor 111 lore'''· bemg. C(J\ered o\er large trat:t' "ith xeroph)tJc \egetaunn. mcludmg l•pcn woodland up to mnnt.rne 'teppes and Oriental thom-cush1on fom1auons. T\\O land ...capcs st.md out here. dl\linguished by 'ummer·\\:trm .md \\Jnh:r-mlld climate, abundant rainfall, and high air humidll)' They rcprt•,cnt refuge' of TertJ:tr) dec•duou., broad-leaved trees. \\hich othCT\1.1\C 111 \\'c,tem Euras1a are 9 cxtmct. as Yo ell us habttats oft tden K oak 'pe"tes (rehch) Colkheu at the Bla k Sc.t .md l.cnknr.tn .11 the C.t,plan ()ea The Colkhtc .uu.l H) r~,;antan ue~;tJUilU' hlllad-lea\cd lore~t are n~;h 10 e en!reen Y.Olld) 'peetcs 10 the unJergro" th. \\ llh plenty ol hana' and eptphytl!s. and are characteri ... u.: of a \\arm-hurmu chm:uc of suhmendtonallalltuJe, (not 'ubtropil:al \ egctauon•• 1, ollen referrcJ to m Caucaswn and Ru,,tan lneratun:). ~ pecil"i con c~pl'- -.eu:nteen \ en.us eight oak pecies Dcpendmg on the ta\Onomic conception ot the authors, the d.ua about the numllcr ol Q11t n 111 s~ctc' occurring in the Cauca'u' Reginn "Uf) con,iucrahl} t !i to rnurc th.m 17 'pcctc,). The "~pliuer·· concept ul ~pccu:' accordmg to the KOMi\ROV s(houl (t'.g. Ctcrepano\ 1995). and accepted hy most uf the author... ol the Cuw.:a'u' state' (e.g. Flora Korlu1:.al Gro~~hcim 1945. Flora A:allcl]c/?mwl Bandm 1952. Drndmj1ora Kavlu1:al Matika'!~ ih 1961. Flora Gru111 I 975. ArtJUntan & ArtJunjan 1985. and other floras and dendmlogtcal ham.lhouk,, or Krc\ er et al. 200 I). contra,ts \\ ith the "Jumper"' concept (e.g. Go\aen' < 1-ruJm 11)98) \\hich is al\o referred to b) the monograph of the East European. Cauca tan and A'ian oaks. b) Menitsk) ( 1968-::!IXJ2). II. ho\\c\er. hi' 'uh)~ct• are t.1kcn mto consiJerauon. the number of the taxa rcgard~J a' c'~cnual de\ i.ue lmm c<K·h other to a le~~er e.\tem hec Table I ). Up to no\1. most ot the Cauca,ian author... dtlterenuate bct\\ecn 17 oak 're~•c,, \1.1th Jc\tallnn' of ,mgle author' bcmg po"tble, panl) due to recogmL· mg adJullln:~l pcctc' (e.g Q h)poc·lrry~al or le"er 'pectc' (e.g. ol the 'pcctc' complc:" Q urft'cU•na). Semag:ina ( 1999). for in,tan~·e. refer; to 7 '~etc' ;~lone from the hmn~d an:a m the \\C,tem Great Cauca'u' !Caucll)U~ State Na1ure Resen e 1. Kol.tkc." ,kj ( 198::! 1 con,tders Cauca.,ta a' one of the centre~ ul 'pectallon 10 the genu~ Q11nnt1 and of con sen atwn of relic~ m EuraSia The 'pccie\ concept lollo\\ed here largcl) corresponds Ill that of Menitsk> 1 llJ6!i-:!002) Sun liar arc also the 'pecies interpretation m the 1/nra oj T11rkev ( ll edg~ & Yaltink 191!2) anJ in the World Check/ill b) Govaem & Frodm ( 1998). C\en though the numher of l.ubspectes accepted by these authors i~ lower and the tklirnitallnns ul' '"htch (and thus data of distribution) ma) differ. Hence. in Cauc.:.tsia \\ ith regard tu broader interpretation of Q. rohur. Q. pt•traea anJ Q. in(i•cwna. R ~pccu:s anJ II· I~ taxa. respecti\ely. lli.:(Uf, being acccptt'J a' 'pc~·ie . or at lc.tst 111 the r.ml.: of ... ub,pectc~. In '>(lllth\\est A ia h)bridtsauon seems to e,senuall) ~,;ontnbute to the fom1ation of ta'o.a anJ t1l the blurring of ,pecje, limn,, respt.'CII\e1y let. Menn'k) 196lS-19 4, Ka,aphgtl 1981) A,cording 10 Kolako,,k.i (IIJ, :!) c g m the o\crlapping area of the re,pecthe close!) related taxa of Q. mbur sen'u Jato and Q pelrat c1 'en u Jato. a great number of h) brid form' occur 111 Cauca,ia. but he accepts the Caucastan taxa each as mdependent enc.lcmt, 'P.:Cit' (Q. mrernma. Q. ibenca). Concerning the European "pecie,. h) hridt~aunn ha been \ erilicd ba::.cc.l on cros~mg cxpenment~ and geneuc analyse,. 1'\umeruu-. comhmauon~ tdenulied 11' h) hrid • or those a'~umeJ ;c. ;,uch, Yo ere gt\ en bmary name,. "htch aJ,o apph~ to the three occumng bct\\een the three far-,pread European· We!.t A'tan <.pectc.' hkewtsc tnckmg m Cauca~ia. Q f'l!lrat•a \ Q. pllht• Ctllf IQ. \ Wt'/11111 H~ufl or Q. x calrt•1cem Vuk.). Q perraw" Q. ml'llr CQ. "< mwn•a Becht.). Q pu~l<l'llf x Q. mhur (Q. x. ken1trr Simnnk.l. If the three 'pccie' are -..ccn 111 a narro" ta.xonnmtc context and 'ever.tl ... pccte' de,tgnateJ each, the 10 Table: I. Sur.c> of the oak' ot Cau~a.,ta accordmg co \'lli10U' authoN c-/.I:REPA:-:OV GOVAERTS & 1 1995 ), tlor.h ol HWDI:'\ 1 19Y8) the Caucasu' Region Q Cl11tcmt·i}olia Q. ca\faneifolia Q ccHttlltt•ifolw Q han1untallt1 Q. hanwissianD Q. hamurrwnu Q macramhera Q. macraOmhera Q. llwcranthem Q. ponll~a Q. pontica Q. ponuca Q. puhe1n·m Q. pubescens Q. puhe.1c t'/11, hue nol mentioned lor Caucas1.1 Q. infectoria ~en~u lalo Q tii'CI.\111(1 Q. mjcctorw ''P· boi.1 \tl'tt Q mfcctoria ssp. 1 urt'rH Q botHit•n Q. mji·t Uma ''P· lmis.\lcrt Q mjl'ctoria ''P· 1 c•nc•rr 1 Q mfi c tona Q mj(·c tona "~P mfectnt w Q mjc 0111 ia ''P· not m Cau.:a~ta mjec torra not m Cauca ta Q. petraea ~n.<.u lato Q dulec hampit Q. petraea s~p. mt•d11edtehll s p. /11( c/11 ecftt'IHI = "P thc ru a Q daleclwmpii nnt m Cauca~ia Q. d1hnroc hen s11 Q. pttrm·a ~'P· d>lwmc ht'll\1 Q pc•traea "P· tho ica Q ~:agnuna Q petrum "P· iberica Q. thenca Q petmra "P tl>< ricu Q petraea ssp. iberica Q pctrtl('(/ Q. pc·traea '"P· petrtlt'll Q. pc trat•a ''P· petraea Q pimwtiloha Q. pt'lraca ''P· pinnatiloha Q. petram "P· pmnauloba (not mentioned lor Cauc,t'IJ) Q.