NEW AN TENUIOR (ROSACEAE)— INTRIGUING SPECIES CRATAEGUS OKANAGAN AND WASHINGTON COLUMBIA FROM THE OF BRITISH NEW AND A OF OKANAGANENSIS VARIETY C. Phipps J.B. CANADA N6A London, Ontario, 5B7, ABSTRACT RESUMEN INTRODUCTION revealed dispersal recent my prior to its of accumulated duplicate collection axmrnmyofPacificNorthwestCrnmcgusTteCrmaegmdescriWforthatregioncanbedmded™^^^^ fruitandlxttharerepresent^^^^^^^ ripe on color of their fully orpurple-toblackJruitedgroupsbased ThispaperdescribesCrataegustennior,spnov.(reddruited).andC.oknnag<memrivarJ<mg.sp,nn™^ n arra^ e l*-fruitcd),ensuringthattheveryrichvariationencounteredbytheauthormttegenem lun*ia01m„aganisLlyaccount«ifor,atleastasfarasdistinctiveandrq«atedlyoccnrr.nglortnstsc^^^^^ ^“d™fof wererevisedhyPhi^^^^^ Northwest the Pacific OhnaganheingintwLries.ser.Ro,„-nh/olineandser.M«r«m,.hne.The^^^^ MTMandCshcila-phippsiacandwasrevisedfmtherepo^^J^i'n witn ^ Macracmtlm, Senes plane-sided and glandular petioles. nutlets P=Uoles,ishererepresenteLnlyhyC.-m.crncamhn.Crma,*»sch,,s«nrpaandC.marrncand.aarev^ ^ Okanagan the “ngingspecies to occurring from the mucracamhu v ry ance, The new and, at first g local species, C. tenuior, is with white anthers, mmmcanAa oocccciudefnetMaolims, var. Cmlaegus ex Uud. Kruschke. occUeulalis (Britt) var. Journal of the Botanical Research Institute of Texas 7(1) is the most usual form of C. macracmtha west of the Rockies and is the only variety of the species so far record- ed from British Columbia. Crataegus tenuior was collected by myself and O’Kennon BRIT first of R.J. in Sep- tember 1993 in the Shuswap area of British Columbia but, in spite of the similarities alluded to above, several differences from C. macracantha var. occidentalis soon became apparent. These included: often more rounded and leaf apices, generally smaller less coriaceous leaves, small pink anthers, generally more slender thorns, and more delicate plant habit Moreover, (Figs. 10, 11, 12). C. tenuior, like C. chrysocarpa, but unlike C. macracantha, has the sides of the nutlets ± plane or, occasionally, only irregularly and shallowly pitted, as well as glandular, sometimes strongly so, petioles. In addition, C. tenuior shares with sympatric forms of both macracantha C. var. occidentalis and C. chrysocarpa Ashe var. chrysocarpa, the most common form of the latter species, the of trait ripening fruit passing through an apricot to salmon or orange phase on the way to ripening to a bright red (see C. macracantha - Figs. 13a, 12b), different from most other forms both of species, particularly those to the east of the Rockies, the which fruit of normally changes from directly green However, to red. in spite of sharing important charactenstics with C. chrysocarpa, with it is a slender form of C. macracantha (Fig. 12b), sympatric with C. tenuior, that C. tenuior most is easily confused. Crataegus tenuior proves have to a quite limited distribution predominantly in the northern Okanaganof southern Bntish Columbia, mainly from somewhat north Armstrong of to the vicinity of Enderby in the north Shuswap central area where common. From is locally it this region extends thinly into the generally drier it Okanogan Washington of where Crataegus as a whole peters out. Mainly collected in the mid 1990s and 2002, C. tenuior remained undescribed now until because of the need to assess C. macracantha throughout large its range, the greatest of any North American Crataegus, prior to deciding whether should be included in that it specms. Although C. macracantha has proven be to very much variable through of range, especially so in its Quebec and Montana, has, except it for a population in northwest Montana, a strikingly eglandular petioles, trait usually thought of as species-specific. Consequently, a morphometric Northwest study of relevant Pacific was material conducted to investigate these distinctions. study.— Fourteen OTUs flowering and 54 OTUs fruiting (Table constituting all the VWO 1), ffomCT (now TRT) specimens of pmative Crataegus lenulor and macracaMha from Columbia C. Brilish Washington, were The used. flowering specimens were scored lor seven characteristics mean length of three largest leaves, petiole glanduUrity (Figs. 9a, median 9c), thorn length, median thorn thickness, sepal irargin serration (Figs. 9b, 9d), anther color (Fig. and 11), style number. The specimens used the same fruiting c aracters except that number nutlet replaced number style is usually the same) and pitting (it nutlet lateral placed anther coteMissing data, in than 10% if less of OTUs, was meansol repUced by column (character) b^est the OTUs subset of Le considered, u pmteriuri, unequivocally the taxL more than 10% missing if Podanfs cluster analyses were performed using ^5YNN-TTAAXX ZMO*™! 2000. The data were sets ranged before The analysis. results are as follows: The PoCA ordtnattons (Fig and MDS 3) (Fig. of 4) the euclidean sepan distance matrix siraibrly Cru,ueg„s,e„„.mandC.macr«cu„,hucleatdy.Thefirstaxisoftheprin7p:if^^^^^^^^^^^^^^ ““ 62% for of the variation f'"* group alv comprising le s m - 4) small, the ( concordance of very different methods emphasizes tf 2 1 277 and'TCB'=l Phipps, RZila,'WP'=Wm 3 (6923), 4 (8370), 5 (8389), 6 (8400), 7 (8401 8 (6902), 9 (6903), 10 (6963), 1 1 (8304), 1 ), ), 4 (701 5 (701 6 (7171), 7 (8265), 8 (8274), 9 (8451), 10 (8452), 1 1 (8453), 12 (8454), 1), 3), 18 (6792), 19 (8275), 20 (8473), 21 (6484), 22 (7103), 23 (7025), 24 (6783), 25 (8457), J), 32 33 (6787), 34 (6793), 35 (81 76), 36 (6782), 37 (8275a), 38 (6976), 31 (6790), (6781), 4), 42 (TCB4041 43 (8463), 44 (8464), 45 (8478), 46 (6789), 47 (6867), 48 (6871 ), 49 (Z1 7891 ), 50 (Z1 7907), 5 ), , 54 52 7904), 53 (Z22423), (7265). (Z20320), (1 or 4 short margins, 3 styles, sepal „Hr^tnflpnticulate , , :.entxr TheCrataegus OTUs had pink anthers, tenuior all OTUs occidentalis wh _ mm) (mean = 41.66 Jwms (mean = mm) 40.14 and short leaves 1 ^ g l>^whiteorivoryanthers,moredeeplyserratesepalmargms,2or3styles,shortto Aoms mm). and (mean = 47.00 short to longer leaves •^Fnutiiigii ::;emeisolani(OTl]22)inFigure5whichis uch an extr larger nght-hand There also group. is the identifies •^edOTU two larger of and adds on fusion of the all after groups , „Jf"STUs two ffaaiirrllyy llaajr-ggee ei in clustering ^ ‘blaster of OTUs ‘'P‘'“l*“nd“’’;::;u^dLoT.?s22andM.SL,alsono<shown,isvery^^^^ 278 UPGMA except that its left-hand group also includes OTU 39 and group lacks several the large right-hand OTUs which are added only OTU after the fusion of the two big groups. Here also 22 is added last, further emphasizing its isolated position. Finally, ISS (Fig. 6) divides all 54 OTUs into a left-hand group identical to UPGMA that in and OTU the remainder. OTU In this, as in CL, 22 not is as isolated as 54. Summarizing the cluster analyses, there is a consistently occurring group of OTUs 1-16 plus 19 and 26. At least the great majority of may these be have a lovf referred to Crataegus tenuior as they lack, or occasionally evd of nutlet pitting or scarring. any- Similarly, the remainder, comprising a right-hand group, outliers, if large to e left-hand group «- t of C. tenuior, plus may part be outliers, if any, to the combined main groups, in large terrri,„C.macrac»«haasd.evWh-.H„^„.„,„„...... outliers™* 22 and 54. nouble re Ainong the PCoA ordinations, (Fig. 7) yields a left-hand group of OTUs 1-16 plus 19 and 22 and a tag! group of the remainder. Here 41% of total picked up by MDS yields a top-r»i^‘ the first axis. (Fig. 8) mT “ In remainder. “P “ bottom-left group of the s 6 and 54 are peripheral. This strong OTU MDS UPGMA As isolation of 22 situation. in the reflects the duster ^lyses, a oral discrete Crataegus tenuior group may by a lack of, be found characterized that is And nutlets ^‘^^rri^g OTUs with four as in the flowering analyses, the all in lie the putative u differe^ ^’^cept for the OTUs two very isolated 22 and the 54. Significantly, ordination methods identify ‘thhee s&ame main those patterns groups to of and variation very similar also detect the w cluster analyses ^™‘^o’^istendesregardingOTUs22and26.Thefruitinganalysesarethussoin^ „ mumalfy ™ilar of than the perceuug! flowering analyses, an lower observation with the in line vanance accountedforbyaxis 1 of the PCoA. 19.Inth^ OTUsd1--16 + nilar core grouping of ^ tenuiorsp. Crataegus I Phipps, ^ P"** - wwtittnh ssppeecciimmeennss "'1^, contams. trhhoossee analyses, the core tenuior group glanduUrpeHotejl«m^w^^^^ ona 0-1 of three-poin. scale), InU^ cbtactercouldstillberelUblyinlerpreted.nfru«,a^s^^^^^^ Hl5;andOTUslO,16, 19, 13,0 OTUs splits into three subgroups containing: 4, 5, 7, 9, ’ ’ ’ ’ j the significance tlir.iiah seco«lsubgro„p.^^^^ LcUve ^ularily the than l^taaad higher petiole las. of ^ they Figure 6 (ISS) ^ of found in diisis not Similar subgroups are clear. ^ »"3(Figs.7,8).,ti^ofcourse,afeatureofglandstba.d.eyn«y^^ “tat^aybeaumreconsuntcharacteratanthesiscanbelessajelWeE^^^^^^^ nntte^r S®t^g»d,h ‘ht.thasa.mM.ng ^ the tauiorcore fruitinganaly- ofdK«m _ Cr<nnegnsmncrncan,ln.isrepresen.edbya g - macracanthagroup,plusOTU m 18 70 54 OTUs tte ^except OTU the 2^ ^ All 22, see below, interpretation. 8^ We suox^ bc ^ nutlet sides significantly (score of 2) to ) ^ UX i. ) 1.25, ’^hich have (respectively middling nutlet side scores macracmtha -t.in.esc.nster,oUwi.b.becote.em,iorgmuptbou^^^^^ Wit Ponp but group. in the part closest to the tenuior comprises the smaUer- ^3 29,30,32,K^^^^^^ ^-dchaster OTUs containing 20,21, 17, 18, tenumrThe^^^^ ^v^specimens more likeC. with dimensions thevanousanalyses,afactparticularlyseeninbolh ^^^^bly fimitingsam^^ the that A reason major CLdendrogram not shown. no doubt due to the that is andAe^^^ this diverse, ^ l^gerthanthe one(540TUscompared to 14), flowering be collect can . specimens longer which adequate time-frame over Journal of the Botanical Research Institute of Texas 7(1) Thef mentioned above- differences as However ^ OTUs ruiting more variation is present and two mutually groups do exclusive xr *^'*‘^’^g^;Neverthele^,agroupofl-16 + 19isconsistentlyfoundwhichisCrataegusfenMiorInfruit,thebest seems interpretation to be accept to the presence of plane-sided nutlets as and con- decisive for this species to oHowrnutlivemr'^'TT''1I'1^^^^^ '* haps Vbt^d betw^n chTiX was L '"*r<tcanlha. However, not set the sntdy up minimal test this and, wtl^ldr" ^'XHtM^XT*' an oddity, perhaps cuItivatedXm the Okanagan. This macracant clearly a C. is form svith eglandular and its petioles pitted nutlet.; cnn,hnindividn*abintheXdy,as,.rt:Xh^‘:“““ parts, again Some quite of unlike its the others studied. more distinct features did nnt aet f“n''«inore> meXenXs tog thornless, achieved it thorns of toaanahtyntotedistinctthantheatalysessLandXtaXr^ g u-h!tn>mTonasket,Washmgton(Fig Mhlhisindwidualappearsthentost new and a variety of okanaganensis V. C. m OTU PcoA. the only and peripheral It is extreme UPGMA MDS, most extreme in ISS is of in and second all is Unlike C. tenuior, and deeply pitted nutlet sides. in the study combine highly glandular petioles, to four nutlets, tohaveadistmctively 68 studied specimen of the only the ithasessentiallyglandlessleaf-teethatxlO.Also,it is a very rare This, suspect, is chrysocarpa. 1 C. resemblances to non-macracanthoid shape, one with clear leaf C between chryso- ^ple an hybrid or clone thereof, of in the study perhaps unique in collections, area, ‘^^^ndC.macracanthavar.occidentalis. names taxonomic consistent is is groups to and flowering Confirmation of fruiung that the calibration being from the and JBP 8401) 6814 (JBP tenuior by two flower™ sDecimens attributed to Crataegus JBP 6903 (flowering) C occidentalis macraeantlia var. while for plants asJBP 7171 and JBP 8454 (fruiting) comes from the same plant asJBP 6871 (fruiting). 282 m Bus 2.5-4.0 es, multi-stemmed: tall, l-year old twigs tan 3-5 to reddish brown, older gray-brown; thorns cmlong straight to slightly recurved, thinnish, glossy black-chestnutwhenyoungAeaves deciduous; peti^^^^ cm 1.25 2.5 long, pubescent only in sulcus, variably cm gland-dotted; blades (2 5-)3-5(-7) long, broadly el- liptic to ovate or rhombovate in general shape; apices subacute, bases broad-cuneate, shallowly 4-5(-^) lobed persidMobesacute to obtuse;margins serrate, teeth with conspicuous black marginal glands (except±entire m basal venation third), craspedodromous with 5(-6) pairs of im- lateral veins, veins adaxially not strikingly pressed at maturity, adaxially densely short-appressed hairy, abaxially thinly pubescent on the veins, other- wise glabrous, texture autumnal thin, color bright yellow. Inflorescences branches tomen- abt. 10-flowered; mbranous, gland-margined 15-17 bracteoles, glands Flowers subsessile. ns glandular-dentic- ± mm); styles 0.6 .^..lalKca. , , ± broad-ellipsoid, t-, < B most easdy distinguished in dower C (Figs. when sympairic 9, 11) diders from it mnrmrandinbyavpety of characters asdisciissed above. In fruit. however.Clenui^ tmguish from smaller-leaved specimens ofC macroconthfl Without fh dissect Th because the ' petiolar glandularity character has some overlap, the m sepal margin cham^m'ayTrracnU d* discern, numllr™'^' vithout and dissection, the nutlet l-f^eoueml'^f™!,''' irequently 't found, no‘t''a At that stage, is diagnostic. autumnal although leaf color (yellowish ii new and a Crataegus tenuior sp. nov. variety of C. o Phipps, 54fruitingspedinensofs fit7.StandardizedPCoA of eudideandistance matrix of group is C macmcantha; OTUs 22, 54 exduded. OTUs 22, 26 and 54 are starred. — wtth the and texture J* leaf thorn, i»y twig, the smaller leaves, and greater delicacy of habit, m chamctem several sharing of otC. mncracnntha. With regmd d,rysocarpa, in spite foUo™^^ to C. The nmcrosc^tcahy. separated easily h«ior,seefcey,ithassttchadifferentfaciesthat is it C mneracn«.l.ovar. prizes C. between tenuior, critical distinctions oe Wclirysocarpainthecontextofallthered-fmitedCratacgiofthcOkatiagan. it is best to check several fruits for nutlet pitting. Journal of the Botanical Research Institute of Texas 7(1) macracantha C. C ''*™**' macraamlha» to mT'" uch'more'norifei^r 1®’ overlap'^ “'“'ifT ‘i"™* n smaller-leaved and smaller innorescence tormsof maeracmrt.. C. " suggested '’'TTf origin for the sporadic "“"T*'' ® C. lotnemimm (OuleT «Tr “““,f ,m- ’’ named form u Minnesota in and perhans a th & Great Plains (Phipps O’Kcnnon 2007). Tis nombkT!'l Stffl '"“"'“'t T'!^'!'r"‘‘^'^*7^“^”‘^^«reater01mnava„.^C ^ this paper). This Okanagan en- being the so, distinctive