Eike J. Jablonski LTA, Dept. Horticole, Arboretum 72, Avenue Salentiny L-9001 Ettelbruck, Luxembourg Phone: +352 691 864079 [email protected]
ABSTRACT
The GenGroʐora oI the iVlanG oI &ypruV VhoZV Vome remarkable IeatureV. The oakV, Zith only three inGigenouV VpecieV, are a prominent part oI the GenGrological inventory, incluGing the endemic Quercus alnifolia Poech. The three indigenouV oak taxa, together Zith a recently deVcribed Iorm oI Q. alnifolia and a recently deVcribed hybrid betZeen Q. alnifolia and Q. coccifera L. VubVp. calliprinos :ebb. Holmboe are deVcribed, and their taxonomic VtatuV diVcuVVed.
Keywords: Cyprus, oaks, Mediterranean oaks, Quercus alnifolia, Quercus coccifera subsp. calliprinos, Quercus infectoria subsp. veneris, Quercus ×campitica, Quercus alnifolia var. argentea
27 Introduction
&ypruV iV the third largeVt iVland oI the 0editerranean, Vituated in the eaVtern 0editerranean Sea, about 60 km 40 mileV Vouth oI the TurkiVh coaVt, and 100 km 65 mileV ZeVt oI Syria, Zith an area oI approximately 9,251 km2 (3,572 mi2 . )rom Vea level at the coaVtal belt the land riVeV to Vteep, rocky, diabaVe and Verpentine peakV at the TroodoV range Zith the higheVt peak at 2lympoV (&hioniVtra at 1,952 m (6,401 It. . The climate iV typical arid 0editerranean, Zith a Vhort, cool, Zet Zinter and a long, dry, hot Vummer. :inter temperatureV in the TroodoV range can drop doZn to -10 p&14 p), Zith an average oI 10 ZeekV oI VnoZ above 1,400 m (4,000 It. 5ainIall varieV betZeen only 300 mmyear (12 in on the central plain and more than 1,100 mm (43 in on the upper VlopeV oI 2lympoV peak. ,n ancient timeV &ypruV ZaV largely covered Zith IoreVtV but today moVt oI thiV Zoodland iV long gone. 5emnantV oI it Vurvive on the TroodoV and .yrenia rangeV, Zith 18 oI the iVlandȊV vegetation being claVViʏed aV Zoodland (31 oI the land above 1,000 m3,280 It , that in Vome placeV iV replaced by tall, Vhrubby maquis (4-6 m13-20 It high: Arbutus andrachne L., Pistacia terebinthus L., Olea europaea L., 6t\ra[ ofʏcinalis L. and Quercus coccifera L. , or moVtly by Vub-Vhrubby garigue (leVV than 3 m9.9 It high: Cistus Vpp., Lithodora hispidula (Sibth. Sm. *riVeb. Genista sphacelata Decne. etc. The ʐora oI &ypruV iV remarkably diverVe, due to the varied edaphic and topographic character oI the iVland. There are 128 plant VpecieV endemic to &ypruV, including the Zell-knoZn &ypruV cedar (Cedrus brevifolia (Hook. I. A. Henry and the golden oak oI &ypruV (Q. alnifolia . The plant endemiVm rate oI the eco-region iV about 7 oI the total ʐora oI 1,750 VpecieV (::), 2007 . Human impact haV been diVaVtrouV Ior the ZoodlandV over the laVt 1,000 yearV, and today only Vmall IoreVt habitatV are leIt intact. 2ld-groZth black pine IoreVt (Pinus brutia Tenore iV Iound only in the high mountain rocky VummitV. The endemic Cedrus brevifolia IoreVt iV repreVented by only a IeZ hundred hectareV. TheVe IoreVtV oZe their exiVtence to none other than :inVton &hurchill. ,n 1907, aV 8nder Secretary oI State (and -unior 0iniVter reVponVible Ior &ypruV, he initiated a reIoreVtation program to replace the IamouV IoreVtV deVtroyed by centurieV oI Ielling that provided Iuel Ior Vmelting but alVo cauVed extenVive VyVtematic eroVion (%arton, 2002 .
Figure 1/ Forest cover in Cyprus. 28 Oaks of Cyprus
Native oak species of Cyprus
Three VpecieV oI oak are indigenouV to &ypruV: the deciduouV Q. infectoria 2liv. VubVp. veneris (A. .ern 0eikle (Section Quercus and tZo evergreen VpecieV, Q. coccifera L. VubVp. calliprinos (Webb) Holmboe and Q. alnifolia Poech. (Section Cerris). Additionally, one poVVibly diVtinct variety oI Q. alnifolia and one rare hybrid have been obVerved. 1/ Ripening acorns of Q. infectoria subsp. veneris. Quercus infectoria 2liv. VubVp. veneris (A. .ern) 0eikle (Section Quercus) &ypruV oak, Aleppo oak: DeciduouV to Vemi-deciduouV tree, up to 18 25 m59 x 82 It (old VpecimenV oIten Zider than high Zith a Vpreading croZn and a trunk that can eaVily reach a girth oI more than 7 m23 It), bark dark grey, vertically ʏVVured on the trunk tZigV hairy Zhen young, leaveV variouVly Vhaped, up to 15 7 cm (6 2.7 in), glabrouV, Vhiny green abaxially, Vlightly tomentoVe beneath, marginV Vinuate to Vinuate-Verrate, petiole 5-30 mm (0.2-1.2 in) acorn Vhortly peduncled or VeVVile, 3-5 cm (1.2-1.6 in) long, cupule Zith adpreVVed VcaleV, encloVing one-third oI the acorn, ripening in the ʏrVt year. 1omenclatural note: Like other VpecieV oI the genuV, the taxonomy oI Q. infectoria VubVp. veneris iV Vtill controverVial and Veveral authorV do not accept the VubVpecieV epithet. %aVionym: Q. veneris A. .ern. (1904). 2Iten incorrectly reIerred to aV Q. infectoria VubVp. boissieri (5eut.) 2. SchZar] (1934), but Zith the incluVion oI Q. veneris A. .ern. (1904) at VubVpeciʏc rank Zithin thiV taxon, the name haV to be Q. infectoria VubVp. veneris due to itV priority at that rank being eVtabliVhed by Q. lusitanica VubVp. veneris (A. .ern.) Holmboe (1914) Vee ,&%1 Art 11.4 and itV Ex. 15. Some authorV reIer to Q. infectoria VubVp. boissieri (5eut.) *¼rke, but *¼rke in .. 5ichter, Plantae Europaea 2: 69 (1897) ZaV unclear aV to Zhich rank he applied the epithet, Vo hiV combination cannot be taken into account Ior purpoVeV oI priority Vee ,&%1 Art. 35.3. (Trehane, 2010). Some authorV place thiV taxon at VpecieV level becauVe it iV tree-like Zith larger, diVtinctly petiolate leaveV, and large acornV, ZhereaV Q. infectoria VubVp. infectoria iV Vhrub-like Zith Vmall leaveV, Vhort petioleV and VmalliVh acornV. ThiV VubVpecieV groZV in &ypruV, Turkey and eaVtZardV to ,ran. ,n &ypruV, it iV Iound in all diVtrictV in open Zoodland and cultivated ʏeldV Irom Vea level up to 1,700 m (5,576 It) altitude, but particularly in the ZeVtern partV oI &ypruV, in deep VoilV. A Vmall IoreVt oI Q. infectoria VubVp. veneris iV Iound near the village oI AgroV, on the VouthZeVtern VlopeV oI the TroodoV 5ange, at 1,100 m (3,608 It). Although the durable hard Zood Zould make excellent ʏre Zood or could be uVed in making toolV, the tree iV rarely cut doZn, poVVibly becauVe the Vhade caVt by their enormouV croZn ZaV important Ior the people and huVbandry (&hapman, 1945) or becauVe oI Vacred connotationV. ,n &ypruV many old, giant treeV oI thiV VpecieV are Iound, Vcattered on the iVland and oIten aV Vingle treeV or in Vmall VtandV, many oI them noZ protected aV natural monumentV. The acornV have been an important Iodder Vource Ior Vemi-Zild pigV.
29 2/ One of the many impressive Quercus infectoria subsp. veneris that can be found in Cyprus. The age of this tree is estimated at 500 years. Q. infectoria VubVp. veneris iV today overall a rare tree in the 0editerranean and only Iound in &ypruV Zithin the European 8nion. The natural VtandV oI thiV VpecieV Zere added in 2003 aV an Annex , Habitat oI &ommunity ,ntereVt under the HabitatV Directive (AnonymouV, 2006). 1atural IactorV contributing to the problemV oI Q. infectoria VubVp. veneris include poor Veed diVperVal, probably due to the abVence in the loZlandV oI the EuraVian May (Garrulus glandiarius) that normally IeedV on acornV and thuV VpreadV them accidentally. Survival oI young oakV iV alVo Meopardi]ed by drought periodV that have become more IreTuent in recent yearV (AnonymouV, 2006). TreeV groZn Irom Zild-collected provenanceV have proven their hardineVV in central WeVtern Europe (Luxembourg, *ermany). TheVe mainly deciduouV treeV groZ VloZly but Zithout maMor IroVt damage in theVe climateV Zhere Zinter temperatureV drop beloZ -20 p& (-4 p)). Quercus coccifera L. VubVp. calliprinos (Webb) Holmboe (Section Cerris) .ermeV oak: Evergreen Vhrub to Vmall tree, up to 10 7 m (33 23 It), bark grey, Vmooth Zhen young but vertically ʏVVured Zith age young tZigV Vlightly pubeVcent leaveV variable in Vhape, up to 5 2.5 cm (2 0.98 in), leathery, Vhiny light dark green above, glabrouV to Vlightly pubeVcent beloZ marginV moVtly VpinoVe-dentate, oIten undulate petiole up to 5 mm (0.2 in) acorn up to 3 1.5 cm (1.2 0.6 in), cupule up to 3 cm (1.2 in) in diameter, covered Zith looVely adpreVVed or diVtinctly recurved VcaleV, encloVing one-third to three-IourthV oI the acorn ripening the Vecond year aIter ʐoZering. 1ative alVo in Turkey, Syria and ,Vrael, the .ermeV oak iV Iound all over &ypruV Irom 100-1300 m (328-4,264 It) in maTuiV and pine IoreVtV, Iorming Vmall VtandV oI VhrubV. A IeZ giant .ermeV oakV are knoZn. The Zood ZaV uVed Ior Iuel, the acornV Ior pig Iodder. Q. coccifera VubVp. calliprinos iV hoVt to the inVect reVponVible Ior carmine Vcale, Coccus ilicis Planch. (Hemiptera: Coccoidae: Dactylopididae) the larvae appear aV red
30 Oaks of Cyprus berrieV on tZigV and leaveV. A red dye ZaV obtained Irom it in the paVt, and gave the VpecieV itV name (coccus coccineus: red berry). Like in other VpecieV oI the genuV, the taxonomy oI Q. coccifera iV Vtill controverVial and Veveral authorV do not accept the propoVed VubVpecieV. 8ntil there iV a modern reviVion oI the VpecieV complex, it VeemV ZiVe to accept the Vplitting and accept VubVp. calliprinos, the eaVtern, more tree-like Iorm oI Q. coccifera. Quercus alnifolia Poech (Q. infectoria Gaud., Q. cypria -aub. Spach, Q. ilex Sibth.) (Section Cerris) golden oak oI &ypruV: Evergreen Vhrub to Vmall tree, up to 10 5 m (33 16.4 It), bark grey, Vmooth Zhen young and vertically ʏVVured Zith age young tZigV at ʏrVt denVely grey tomentoVe leaveV thick and leathery, Vlightly convex, ovate, oblong, obovate or Vuborbicular, up to 6(-10) 5(-8) cm2.3(-3.9) 2(-3.1) in, dark Vhiny green and glabrouV above, beloZ denVely golden to broZn or greeniVh tomentoVe (rarely Vilvery-grey) Zhen young, turning dark broZn or nearly black Zith age marginV dentate or lobulate tomentoVe petiole 6-12 mm (0.23-0.47 in) acorn narroZly obovate or Vubcylindical, up to 4 1.2 cm (1.6 0.47 in), cupule encloVing one-Vixth to one-Tuarter oI the acorn, covered Zith Vtrongly recurved, linear VcaleV acorn ripening the ʏrVt year. Endemic to &ypruV, very common, but occurring only on the ultra-baVic rock IormationV oI the TroodoV 0aVViI Irom 300-1800 m984-5,904 It (rarely beloZ 450 m1,476 It and Vcattered above 1,600 m5,248 It), Q. alnifolia iV the dominant VpecieV oI dry habitatV in Pinus brutia IoreVt or IormV denVe high maTuiV in meVic habitatV. 9ery IeZ giant treeV oI Q. alnifolia (10 m33 It) are knoZn, and theVe are exceptional. The VpecieV haV a high ecological value, Vince it thriveV on Vtony and rocky mountainVideV, preventing eroVion. ,t coppiceV Zell Irom the baVe and thuV reinVtateV itVelI aIter ʏreV. The acornV Iorm an important diet Ior the mountain Iauna, the hard Zood iV Vuitable Ior the conVtruction oI tool handleV and other Vmall articleV (TVintideV et al., 2002). The Zood ZaV alVo one oI the principle Iuel VpecieV uVed to make high-grade charcoal (&hapman, 1945). The habitat type ȍScrub and loZ IoreVt vegetation oI Q. alnifolia (9390)Ȏ iV a priority habitat oI Annex ,, Directive 9243EE&. ThereIore, large IoreVt expanVeV have been propoVed Ior incluVion in the Natura 2000 netZork, to protect the endemic IoreVtV oI golden oak. Q. alnifolia aV a VpecieV iV protected by &ypruV IoreVt laZ. The Red List of Oaks (2ldʏeld EaVtZood, 2007) liVtV Q. alnifolia aV a vulnerable VpecieV >9u %1 ab(ii)@ Vaying that ȍhabitat degradation by ʏre, gra]ing, Zood cutting and developmentȎ threatenV the VpecieV. Q. alnifolia iV the 1ational Tree oI &ypruV. Drummond, Zho viVited &ypruV in 1754, VpeakV oI it aV ȍȓa kind oI alder, the leaveV oI Zhich Vhine like a green orange the backV oI them, Zhen young, are yelloZ, but aV they groZ old, they turn broZn.Ȏ (&hapman, 1945). The AuVtrian botaniVt 3/ Quercus alnifolia with the author.
31 Theodor .otVchy collected thiV oak at the end oI 2ctober in 1840 ȍȓZith ripe acornV, on the eaVtern VlopeV oI 2lympoV.Ȏ (.otVchy, 1862). He Vent thiV material to hiV colleague -oVeph AloiV Poech in 9ienna, Zho deVcribed Q. alnifolia in 1842 aV Zell aV a couple oI the other plantV oI &ypruV collected by .otVchy. Already a couple oI yearV earlier, in 1834, the )rench botaniVt Pierre 0artin 5©mi Aucher-loy collected ȍinVuIʏcient material Irom Vhrubby Vpecimen, Zith undeveloped acornVȎ (.otVchy, 1862). 2ne year aIter PoechȊV deVcription oI Q. alnifolia, the )rench botaniVtV Hippolyte )ran§oiV &omt©
4/ The elegant acorns of Quercus alnifolia. de -aubert and Edouard Spach publiVhed in 1843 a deVcription oI the oak collected by Aucher-loy under the name Q. cypria -aub. Spach. The draZingV oI Q. alnifolia in &amuV (1934) are made Irom the original material oI Poech and alVo oI Aucher-Eloy 1o. 2861, and the not Iully developed acorn Irom the latter collection iV clearly viVible. .otVchy (1862) VtateV that the VpecieV may be hardy in Vouthern Europe aV it iV covered more than a month Zith VnoZ in itV natural habitat. The VpecieV ZaV introduced into .eZ in 1885, Zhere it proved to be ȍȓperIectly hardy, but VloZ groZingȓthe yelloZ indumentum oI the leaveV iV only Vlightly developed Zith a more grayiVh underVurIace.Ȏ (%ean, 1976). Semi-deciduouV treeV groZn Irom Zild-collected provenanceV have proven their hardineVV in central WeVtern Europe (Germany: %lack )oreVt, 850 m2,788 It, HardineVV =one 7a), groZing VloZly but Zithout maMor IroVt damage in theVe climateV Zhere Zinter temperatureV drop beloZ -20 p& (-4 p)), under VnoZ cover. Quercus alnifolia Poech var. argentea HadMik. Hand (2005) ,n 2005, HadMikyriatou and Hand deVcribed a neZ variety oI Q. alnifolia. They obVerved a number oI populationV oI Q. alnifolia aV Vmall groupV oI plantV Zith ʐattiVh leaveV Zith Vlightly revolute marginV and a Vilvery-tomentoVe loZer VurIace. The authorV conVider thiV conVtant variation aV remarkable enough to Vupport the creation oI a varietal level taxon (Hand, 2006), deVcribing it aV ȍQuerco alnifolia var. alnifolia foliis adultis
32 Oaks of Cyprus complanatis argenteis glanduloso-pilosis infra differt.Ȏ AV the leaveV oI the golden oak are Tuite polymorphic, the above obVerved variation may not prove VuIʏcient to maintain thiV varietal level taxon. Etymology: argentea Vilvery Quercus ×campitica HadMik. Hand (Quercus campitica nothoVubVp. hylatis HadMik. Hand) >Q. alnifolia Q. coccifera VubVp. calliprinos@. 5ecently, hybridi]ation betZeen tZo VpecieV oI &ypruV oakV haV been deVcribed: The tZo evergreen &yprian oak VpecieV, Q. alnifolia and Q. coccifera VubVp. calliprinos, Iorm mixed populationV around the TroodoV mountain range. ,t ZaV knoZn Ior Tuite a long time that the local inhabitantV oI the village oI .ampoV in the northZeVt part oI the TroodoV range had obVerved that Vome oakV Zithin theVe mixed populationV bear characteriVticV oI both taxa. They adopted Ior them the vernacular name “Lakopernia” (Irom the &yprian expreVVionV Ior Q. alnifolia = lakia, and Q. coccifera VubVp. calliprinos = pernia >prinos@). ,n 1999 Veveral VpecimenV Zere collected and the holotype ZaV deVcribed in 2005 (Hand, 2006) aV Q. campitica nothoVubVp. hylatis HadMik. Hand, Zhich ZaV contrary to Art. 34.2 ,&%1 (2006). ,n 2007 Greuter 5auV publiVhed the name validly aV: Quercus campitica HadMik. Hand (Q. alnifolia Q. coccifera VubVp. calliprinos) (2005). (holotype: %, iVotype: & 33 in Q. alnifolia, but rare pollination oI Q. coccifera by Q. alnifolia could be inIerred. ThiV coincideV Zith the obVervationV made by Hand (2006). 5eproductive barrierV may be the reaVon Ior the limited genetic introgreVVion betZeen the tZo VpecieV. An allo]yme characteri]ation oI Iour 0editerranean evergreen oak VpecieV baVed on cladiVtic analyViV (Toumi Lumaret, 2001) haV VhoZn a cluVter oI Iour VpecieV (Q. ilex L., Q. coccifera and Q. alnifolia, Zith the Tibetan Q. aquifolioides 5ehd. WilV., ZhereaV Q. suber L. and the Himalayan Q. semecarpifolia Sm. in A. 5eeV Zere both genetically very diVtinct). According to Toumi Lumaret (2001) both Q. alnifolia and Q. coccifera Vhould be claVViʏed in the Vame cluVter oI 0editerranean oakV that correVpondV Zith Section Cerris VenVu 1ixon (VubgenuV Sclerophyllodrys VenVu SchZart]). Introduced oak species ,n addition to the indigenouV oakV, Veveral VpecieV have been introduced, either Ior economical or ornamental reaVonV. TheVe include Q. ilex L., Q. ithaburensis Decne. VubVp. macrolepis Hedge Conclusion The ʐora oI &ypruV VhoZV a high diverVity, Zith a total oI 1,750 plant VpecieV knoZn, due to the edaphically, climatic and topographic variation. The three indigenouV oakV Q. infectoria VubVp. veneris, Q. coccifera VubVp. calliprinos and the endemic Q. alnifolia are deVcribed, aV are the recently deVcribed hybrid Q. campitica and a purported variety Q. alnifolia var. argentea. Their taxonomic VtatuV and the poVVibility oI occurrence iV VubMect to diVcuVVion. Photographers. Title page: %©atrice &haVV© (Quercus alnifolia). PhotoV 1-4: Eike -ablonVki. Bibliography AnonymouV. 2005. Promoting Protection and Sustainable Management of the Oak Trees of Cyprus. European &ommiVVion, &ypriot &ivil Society in Action ,, Grant Program (Europeaid127215LA&T&<). %arton, G.A. 2002. Empire Forestry and the Origins of Environmentalism. 1eZ 34