TurkJBot 28(2004)29-37 ©TÜB‹TAK TheGenusCrataegus L. (Rosaceae)withSpecialReferenceto HybridisationandBiodiversityinTurkey AliA.DÖNMEZ HacettepeUniversity,FacultyofScience,DepartmentofBiology,06532-Beytepe/AnkaraTURKEY Received:23.09.2002 Accepted:24.02.2003 Abstract: ThegenusCrataegus L.hasapproximately200speciesworldwide.However,thenumberofthespeciesisincreasedto 1200speciesbysometaxonomists.Traditionally,thegenusisplacedinthesubfamilyMaloideae anditiscloselyrelatedtothegenera Pyracantha M.Roem.,Mespilus L. andHesperomeles Lindl. Thelatestphylogeneticanalysisofthegenusshowsthatitisclosestto thegenusOsteomeles Lind. Crataegus isdividedinto40sectionsbyvarioustaxonomists.Thegeneticdiversitycentreofthesection Crataegus rangesfromTurkeytoIran.AccordingtofieldobservationsinTurkey,newdiversitycentresareproposed.Thelatest studiesonTurkishCrataegus speciesshowthattherearemorespeciesthanhithertoknown.Hybridisationiscommonbetweenthe Crataegus speciesinTurkeyandthebiogeographicdistributionsofthemandtheirparentsaregiven.Somefeaturesofthegenus, includingleafandpyrenesmorphology,pyrenesnumberandfruitcolours,arepolymorphic.Polymorphismandhybridisationarethe mainreasonforabundanceofthesynonymsofthe Crataegus species. KeyWords: Crataegus,Rosaceae,taxonomy,Turkey Crataegus L.’un(Rosaceae)Türkiye’dekiDurumu: BiyoçeflitlilikveMelezlenme Özet: Crataegus L. cinsiyeryüzünde200kadartüresahiptir.Bununlabirliktebusayıbazıtaksonomistlertarafından1200’ekadar çıkarılmıfltır.Bucinseskidenberi Maloideae altfamilyasıiçinekonmufl, Pyracantha M.Roem.,Mespilus L. veHesperomeles Lindl.’in yakıncinsleriolarakde¤erlendirilmifltir.Ensonyapılanfilogenetikanalizleregöre,Osteomeles Lindl. cinsinedahayakınoldu¤uortaya konmufltur.Crataegus çeflitliarafltırıcılartarafından40seksiyonabölünmüfltür.Buseksiyonlararasında Crataegus seksiyonunun genetikçeflitlenmemerkeziTürkiye’den‹ran’ado¤ruuzanır.Arazigözlemlerimizegöre,Türkiye’debazıyerlerinbucinsinçeflitlenme merkeziolaca¤ıdüflünülmektedir.Yaptı¤ımızarafltırmalarbucinsinTürkiye’debilinendendahafazlatürüoldu¤unugöstermektedir. Türkiye’deki Crataegus’lardamelezlenmeyaygınbirdurumdur.Tümtürlerinbiyoco¤rafikda¤ılıflıvemeleztürlerinataları belirtilerek,tartıflılmıfltır. Cinsinyaprakveçekirdekmorfolojisi,çekirdeksayısı,meyverengigibibazıözellikleripolimorf tur. Polimorfizimvemelezlenme,Crataegus türlerindebukadarçoksinonimolmasınınnedenlerindendir. AnahtarSözcükler: Crataegus,Rosaceae,taksonomi,Türkiye Introduction FloraCaucasica(Grossheim,1934)andtheFloraof Linnaeusrecognisedthegenera Crataegus L. (nine Turkey(Browicz,1972). species), Mespilus L. (sevenspecies), Pyrus L. (four Ihaveperformedaliteraturesurvey,conductedfield species)and Sorbus L. (twospecies)inthesubfamily studiesandexaminedherbariumspecimensandhaveseen Maloideae (Linnaeus,1754;Robertsonetal.,1991). thatthetaxonomyof Crataegus isproblematicdueto Intragenericclassificationofthemaloidgenerahasbeen polymorphiccharacters,hybridisation,insufficient studiedindetailbyPhipps(1983a),Phippsetal.(1991) collectionsandlackoffieldobservations. andCampbelletal.(1990).Acomprehensive The Crataegus accountintheFloraofTurkey monographicstudyoftheAsiaticandEuropeanCrataegus recognises17species,onesubspeciesandthreevarieties specieswascarriedoutbyChristensen(1992)and (Browicz,1972).Thetaxonomictreatmentofthe includedtheTurkish Crataegus species.Thetaxonomic TurkishCrataegus speciesbyChristensen(1992)differs treatmentsontheTurkishtaxawerebasedonherbarium fromthatofBrowicz.Heaccepted19species,one specimensandsomepreviousliteraturerecords,suchas subspecies,andtwovarieties.Inaddition,hereported5 29 TheGenusCrataegus L.(Rosaceae)withSpecialReferencetoHybridisationandBiodiversityinTurkey speciesasnewrecordsforTurkeyandhehasalso Habit:Treeorpolycormicshrubs,andmostlyarmed. recentlydescribedtwonewspecies.Hechangedthe Therearetwotypesofthorn;thefirstoneissmalland statusof11speciesrecognisedbyBrowiczintheFloraof lacksleafandinflorescence,andthesecondoneisstout Turkeyeitherbyreducingintosynonymyorbymaking andbearsinflorescence.Shortthornpossiblyturnsto newcombinationsorstatus. stoutovertime.Bothtypesofthornareonalltreesof Thegenus Crataegus hasbeendividedintomany somespecies.Toprepareanaccurateherbariumsheet sectionsandsubsectionsbyvariousauthorsinthelast bearingthorns,thornybranchesoftreesshouldbe century.Thesectionsweretreatedtaxonomicallyandall pruned. werereorganisedbyPhipps(1983a);oneofwhich,the Indumentum:Fivekindsofindumentumareseenon section Orientales ZabelexC.K.Schneid. sensu the Crataegus specimensinTurkey:(1)villous,(2) Christensen,ispresentinTurkey.Thenothosection lanate,(3)sericeous,(4)tomentoseand(5)appressed- Crataeguineae K.I.Chr. hasrecentlybeendescribedby pubescent.Hairsoftheplantsusuallydecreaseduring Christensen(1992),thoughitisnotrepresentedin transitionfromfloweringstagetofruitingstage. Turkey. Leafandstipule:Bothleafandstipulecharactersare frequentlyusedinthetaxonomyofthegenusandthere MaterialsandMethods arethreetypesofleaves(Smith&Phipps,1984; Dickinson,1986). Leavesofelongatedshoots :these Theplantspecimensusedinthisstudywerecollected developfromtheaxisofmainbranchesororiginatefrom fromdifferentpartsofTurkeyover8years.Materials theapexofpreviousyears’branchesonthesameplant. fromvariousherbariainbothTurkey(HUB,GAZI,VAN Leavesoftheelongateshootsarelargestononeofthe andANK)andEurope(P,JE,PR)werealsoexamined.To plantsandthereisnofloweronit.Similarly,stipulesof collectbothfloweringandfruitingmaterials,mostofthe theelongatedshootsarealsothemostrobustamongall localitieswerevisitedtwice.Observationsonhabitat,life stipulesofthesameplant.Theincisionisdeeperthanthat form,phenologyandsomemorphologicalfeatureswere oftheothers.Leavesofshortshoots:leavesappearbelow notedandphotographed. theinflorescenceandtheyshowvariationinshapeand TaxonomictreatmentsbyBrowicz(1972)and size.Thisvariationiscalledleafheteroplasty(Smith& Christensen(1992)ontheTurkish Crataegus taxaare Phipps,1984;Dickinson&Phipps,1986).Leafvariation quitedifferentfromoneanother.Inthisstudy,forthe isalsopresentonleavesofsterileshootsoftheshort taxonomyofCrataegus,thechangesmadebyChristensen shoot.Subterminalleavesoftheshortshootarea areexplainedundertherelatedspecies.Thetaxonomic characteristicandusefultaxonomiccharacter.Numberof treatmentofthegenusbasedonChristensen’srevision lobepairs,depthofdivision,andnumberofteetharealso andthesynonymsofthetaxaarenotgivenhere. commonlyused. Leavesofsterileshoots :similarto floweringshootsinbothshapeandsize,buttheseare clusteredonthetopoftheshortshoots. Results Inflorescence:Corymbs,rarelyumbel,terminalon Adetailedexaminationofthecollectedspecimens, floweringshoots.Thenumberofflowersinthe herbariummaterial,andliteraturesurveysshowthat21 inflorescencevariesfromafewto50. Crataegus Crataegus speciesgrownaturallyandtwospeciesgrowin pentagyna Waldst.andKit.exWilld. ischaracterisedby cultivationinTurkey.Allthespeciespresentinherbaria manyflowersamongtheotherCrataegus species.Bracts: orcollectedfromthefieldhavebeenexaminedandthe theirserration,glandsanddurationarefrequentlyused acceptedtaxonomicranksaregivenhere.Diagnostic fortaxonomicpurposes.Thebractsusuallyfalloffafter featuresofthespeciesandcomparisonsbetweenthe anthesis.Hence,theyshouldbeobservedinthefieldand revisionsbyBrowicz,Christensenandmyown keptinanenvelope. C.tanacetifolia (Poir.)Pers. is suggestionsarediscussedundertherelatedtaxa. characterisedbypersistentbractsandprominentglands. Thefollowingcharactersareusedindelimiting Flower:Five-merous,however,afewfruitswith4,6 species: and7sepalsofC.ambigua C.A.Mey.exBacker havebeen 30 A.A.DÖNMEZ collectedfromCentralAnatolia.Sepalsareindurateinthe Table1.Chromosomenumbersofthe Crataegus taxainTurkey. TurkishCrataegus speciesandarecommonlyreflexedor patent.Erectsepalsarecharacteristicfor C.microphylla Taxa Chromosomenumbers(x=17) C.Koch.C.orientalis M.Bieb. showsvariationintheerect C.tanacetifolia ? patentorreflexedsepals.However,theother Crataegus C.azarolus var.aronia 2n(2x)=34 speciesusuallyhaveone-positionedsepals.Itsshape C.pontica 2n(4x)=68 changesslightlyfromspeciestospecies,butitisnot C.orientalis 2n(4x)=68 characteristicformanyspecies.Moreover,variousshapes C.pentagyna 2n(2x)=34 ofsepalareseeninthesameflowerandinflorescence. C.davisii ? Themaincolourforpetalsiswhite,butpinkishpetalsare C.meyeri 2n(3x)=51;2n(4x)=68 seeninC.rhipidophylla Gand.,whilecreamandgreenish C.caucasica 2n(3x)=51 petalsareseenin C.pentagyna.Stylenumberisclosely C.ambigua 2n(3x)=51 relatedtopyrenenumber,bothofwhicharevery C.heterophylloides ? importantforthetaxonomyoftheCrataegus speciesand C.longipes ? whichshouldbenotedinthefield.Thebasalpartofthe C.microphylla 2n=32;2n(2x)=34 styleishairyonfewspeciesofthegenus.Apicesofstyle C.rhipidophylla 2n(2x)=34;2n(3x)=51; haveenlargedsurfacesforpollination. 2n(4x)=68 Fruit:Thecolourofmaturefruitsisusedfor C.pseudoheterophylla 2n(3x)=51;2n(4x)=68 taxonomicpurposes.Yellow,redandblackarebasic C.monogyna 2n(2x)=34 coloursformaturefruits.However,therearealsomany C. xbornmuelleri ? intermediatecolours. C.azarolus L.,C.tanecetifolia and C.x yosgatica ? C. x bornmuelleri Zabel havevarioustonesofyellow C.x sinaica ? while C.pentagyna,C.davisii Browicz and C.caucasica C. x rubrinervis ? C.Koch areblack-fruitedspeciesinTurkey.Thefruitsof C. x browicziana ? therestofthespeciesareeitherredoryellowish-redor C.x kyrtostyla 2n(2x)=34;2n(4x)=68 varioustonesofred. Flesh:theediblepartoffruit betweentheskinandpyreneissometimesusedforthe identificationofsomespecies.Itisusuallypaleyellow,but HybridisationintheTurkishCrataegus species ifthefruitisblack,thenthefleshismostlyredanddark- red-veined.