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A New Genus and Species in Cupressaceae (Coniferales) from Northern Vietnam, Xanthocyparis vietnamensis Author(s): A Farjon, Nguyen Tien Hiep, D K Harder, Phan Ke Loc and L Averyanov Source: Novon, Vol 12, No (Summer, 2002), pp 179-189 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/3392953 Accessed: 15/06/2014 04:52 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive We use information technology and tools to increase productivity and facilitate new forms of scholarship For more information about JSTOR, please contact support@jstor.org Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Novon http://www.jstor.org This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions A New Genus and Species in Cupressaceae (Coniferales) from Northern Vietnam, Xanthocyparis vietnamensis A Farjon Royal Botanic Gardens, Kew, Richmond, Surrey,TW9 3AB, United Kingdom a.farjon@rbgkew.org.uk Nguyen Tien Hiep Instituteof Ecology and Biological Resources of the National Center for Natural Sciences and Technology of Vietnam, Nghia Do, Cau Giay, Hanoi, Vietnam ntienhiep@hn.vnn.vn D K Harder Arboretum,UC Santa Cruz, 1156 High Street, Santa Cruz, California 95064, U.S.A dkharder@cats.ucsc.edu Phan Ke Loc Departmentof Botany,Vietnam National University,Hanoi, Vietnam pkeloc@yahoo.com L Averyanov Komarov Botanical Instituteof the Russian Academy of Sciences, Prof Popov Street 2, St Petersburg, 197376, Russia av@herb.bin.ras.spb.ru ABSTRACT Botanical explorationsof heretofore timeeven thoughtheyare a small groupof plants This dispoorly inventoriedlimestoneformationsin the withno morethan650 species worldwide because northern Vietnameseborderprovinceof Ha Giang coveryhas become even moreinteresting have yieldeda new taxonof coniferthatis to be detailedcomparisons and subsequentanalysishave classifiedin a cupressoidclade ofCupressaceae.It revealedit to be relatedto a species of whichthe It has has foliage and ovuliferouscone charactersthat genericplacementhas been controversial nootkatensis comparecloselyto Chamaecyparis (D thus notonlyenlargedour knowledgeof biodiverofphyDon) Spach, as well as a featurepeculiarto it and sity,but also increasedour understanding rarein conifers:thesimultaneous of the conifersin thewider occurrenceofju- logeneticrelationships venile and matureleaves on normalplagiotropic contextofthefamilyCupressaceaeto whichit bebranchingsystemsof maturetrees.It is proposed longs Implied in these relationshipsis a biogeoin thispaperto uniteC nootkatensis withthenewly graphic historyencompassingeastern Asia and discoveredspecies in a new genusXanthocyparis; NorthAmericathathas been knownfromnumerous ofthisnewgenusand species a fulldescription and previously discoveredrelationships amongplantsof illustrations are provided.Additionalcontributions both continents.The new conifer,its characters, to this paper include observationson its habitat, and taxonomicrelationships are presentedbelow conservation and a taxonomic discussion status, Key words: conifers,conservation,Cupressa- Xanthocyparis Farjon & Hiep, gen nov TYPE: Vietnam,Xanthoceae, NorthAmerica,taxonomy, vietnamensis Farjon& Hiep Xanthocyparis cyparis in superA genereChamaecypari stomatibus utrinque lateralium foliorum maturorum ficiebus disposiequaliter The discoveryofa newlivingconiferthatis suf- tis,foliismaturiis lateralibus apicesliberos(nonadpres4 (raro6) strobilisbracteis-squamis ficientlydistinctso as not to be accommodated sos) gerentibus, minoribus a genereCupresso strobilis bracteissuffultis; in known an is comfortably any genus exceptional minusnumerosis diagnoscenda eventwhichhas onlyoccurreda fewtimesin the squamisseminibusque last 50 years.The conifershave receivedmuchatSmall or largeevergreentrees,monoecious,with tentionfromtaxonomistsand othersover a long fibrousbarkexfoliating in longitudinal strips;heart NovoN 12: 179-189 This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 2002 4:* Missouri Botanical Garden vietnamensis(Cupressaceae), Xanthocyparis a new genus and species (See page 179.) A JournalforBotanicalNomenclature V0LUM E12 N BER UM This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 2002 180 Novon inconspicuaplerumqueadaxialia,pauca in parwood yellowish,slow growing.Foliage branches adultorum te abaxialiae Folia juveniliain verproximali superficiei formspreadingin plagiotropic spraysor drooping, ticillisquadrifoliidisposita,basin versusdecurrentia, paa or ing pyramidal,conical, irregularflat-topped tentia,15-20 mm longa, 1.5-2 mm lata, monomorphia, crown Foliage of three types: juvenile linear linearia,eglandulifera, margineintegra,apice acuto; stoabaxiali juveniliumin vittis2 in superficie leaves,transitional leaves,and maturescale leaves matafoliorum Juvenileleaves presenton seedlingsonlyor also in solumdisposita.Amentamasculasolitaria,in ramulisfolia mature trees, radially disposed in alternating adulta gerentibusterminalia,2.5-3.5 mm longa, 2-2.5 mmlata; microsporophylla 10-12, peltata;microsporangia whorlsof four;foliagebranchletswiththistypeof 2(-3), abaxialia Strobilifemineisolitares,in ramulisfolia leaves alwayssterile.Transitional leaves presenton adultagerentibusterminalesvel subterminales, postduos seedlingsonlyor also in maturetrees,decussate, annos maturi,subglobosi,postdehiscentia9-11 mmlondividedin facialsand lateralsofnearlyequal size, gi, 10-12 mmlati; bracteae-squamae2(-3) paribus,opthe laterals weakly disposed in a plane; foliage positae,decussatae,pare superomajoreconnatovel recluso valvatovel subpeltato,distaliterlatiores,rugosaeet branchletswiththis type usually sterile.Mature recurvo-umbonatae Semina ca 8-9, ovoidea vel irreguleaves presentin maturetrees,decussate,dimor- laria,4.5-6 mmlonga,4-5 mmlata, testain alas 0.5-1 phic in shape and size withthe lateralsstrongly mmlatas expansa flattenedand disposed in a plane; foliagebranchSmall to medium-sized tree to 10-15 m; trunk lets withthistypeoftenfertile.Leaves in whorlsof terete,up to 50 cm diam.; bark smooth monopodial, fouror decussate.Pollen cones terminaland soliand on thin branches, purplish to red-brown,extaryon lateralbranchletswithsmall scale leaves, 2.5-5 x 2-2.5 mm;microsporophylls 10 to 16, de- foliating in thin flakes and strips, on the trunk of trees becoming soft and fibrous, brown to cussate, peltate,bearing2 (or 3) relativelylarge larger gray-brown, exfoliating in numerous thin strips Seed cones terminaland solitary microsporangia Branches long, spreading ?+ horizontally; foliage on lateralbranchletswithunmodified scale leaves, branches numerous, spreading mostlyin plagiotroX to 7-11 10-12 wider mm, opening maturing or Bract-scalecomplexesin (sometimes 3) decussate pic overlapping sprays slightlydrooping,forming in a crown trees but a spreading, young pyramidal pairs, fused at base, the upper pair(s) connate, or flat-toppedcrown in old trees Foliage irregular wide to release subthe valvate to seeds, spreading centralumbo.Centralcol- in mature trees predominantlywith mature leaves, peltate,witha prominent umellapresentor absent,small Ovules axillaryto also withjuvenile leaves, oftenalso withtransitionbracts,1 to per bract;seeds usuallyfewer,con- al leaves Foliage sprays withjuvenile leaves bushy, centratedon the upperpair (if pairs) or middle sparsely branched, ultimate branchlets 20-50 mm withtwothin long, not flattened Foliage sprays with mature pair (if pairs) of scales, flattened, lateralwings.Seedlingswith2 cotyledons, followed leaves flattened,with rounded outline; leading foliage branches quadrangular to terete, with ca by juvenilelinearleaves in whorlsoffour orders of branching, still covered with green leaves in 3rd to 4th year of growth;subultimatebranchlets pinnate, ultimate branchlets of unequal and irregular length and spreading at 300-450, 5-20 x 1.51 Xanthocyparis vietnamensis Farjon& Hiep, mm,distinctlyflattened.Juvenile leaves in whorls sp nov.TYPE: Vietnam.Ha Giang:Quan Ba, of four, decurrent, monomorphic, the distal part Bat Dai Son, Bat Dai Son ProvincialProtected spreading at nearly 90', the proximal decurrentpart Area, 10 Feb 2001, D K Harder,N T Hiep, 4-5 mm long, the distal free parts 15-20 X 1.5-2 R K Loc, L V Averyanov, G E Schatz & S mm, linear, margins entire,tapering to a fine point Bodine DKH 6091 (holotype,HN; isotypes, Stomata in two whitish bands on the abaxial side HN, K, LE, MO) Figure1 only, in each band ca 6-7 irregularrows, oriented Arbor10-15-metralis, monoica;cortex parallel to the leaf axis Transitional leaves similar sempervirens, fibrosus Ramilonges,horizontaliter ramulorumto mature leaves but longer (5-7 mm), lanceolate, dispositi; frondes foliisadultisetjuvenilibus dimorphae praeditae the laterals spreading at 450 Mature leaves decusFolia adulta decussata, inconspicueglandulifera; ea ramulorum 1.5-3 mmlonga,1-1.3 mmlata, sate, short decurrent, imbricate, dimorphic, on (sub)ultimorum dimorpha:foliadorsi-ventralia (anglice"thefacials")ad- (sub)ultimate branchlets 1.5-3 X 1-1.3 mm (the vel rhombica,in partedis- laterals slightlylonger than the facials); the facials pressa,angusteangulato-ovata tall carinata,margineminutedenticulatapraeterapicem narrowlyovate-rhombic,keeled, + appressed, marversusintegra,apice acuto; folialateraliamoderatelonentire towardthe acute giora, conduplicata,basin versus decurrentia,recta vel gins minutelydenticulate to falcata,margineminutedenticulatapraeterapicemversus or acuminate and free apex; the laterals condupliintegra,apice liberoacuto vel pungenti;stomatafoliorum cate, the proximal part decurrent, the distal part Eponymy From Greek xantho = yellow,the colorofthe wood,and cyparis= cypress This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions Farjonet al NewVietnameseGenus Volume12, Number2 2002 181 HERBARI MISSOURI GARDEN NT 519464 _BOTANICAL o "A" (H N) of Qw Vi0 rnip stennsl,44 Fu A a,,9 nn M i rowr; ; f , Det A Lr 2001 Kew) Farjon (RBG I 04 ~;'::~::VIETNAM Ha Siang a Quan Ba1 Ba Distrct, Forest cultivatilons Possiblyore-, a so, only ca.25 years apo sma7l Bat ,re h ,westof:" Da : S , ;n b re few and fire tolerant herbs) trees 04 6064 T Schat~cz e; S Bdn69 very??,anew,?;G.E ted by Naataonal Supprkrrn Sci~ence F'ound)ation~ a nd tMISSOURI M' S rSOU,.RI National Geograp?hiclGARDEN BOTANICAL BOlT AN.I CAL, GARDEN S.%ociety HERBA:RI:UM HElRBAR IUM (MO) ("I vietnamensis of the holotypeofXanthocyparis Farjon& Hiep, D K Harder,N T Hiep,P K Figure1 Photograph G E Schatz& S BodineDKH 6091(HN) Loc, L V Averyanov, This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 182 Novon undertheca 20 m spreadingfree fromthe leaf above at ca 300, Zuccarini.In a second stratum orfalcate,marginsminutely denticulateex- tall canopyspecies ofElaeocarpus,Eriobotrya, Sorstraight and manyothersfrequently occur cept towardthe acute or pungentapex Stomataon bus, Schefflera, matureleaves inconspicuous,mostlyadaxial,a few Shrubsand herbsabound;amongthelatterare nuas well scatteredon the proximalabaxial faces, covered merousspecies of Orchidaceae,terrestrial witha layerof cuticularwax Glands inconspicu- as epiphytic,sometimesdetermining the aspect of ous, in a depressionbelowthekeeled distalpartof the groundcovervegetation.Ferns and especially some of the facials.Pollen cones 2.5-3.5 x 2-2.5 bryophytes are similarlyabundantboth as litho10 to 12, ca phytesand as epiphytes.The limestoneridgeson mm,oval-terete;microsporophylls X mm,witherose-denticulate occursare extremely eroded, marginsand mu- whichXanthocyparis cronate apex, green turningyellow-brown; each composed of resistant,marble-likerock outcrops withthinsoil pockets.The climateis bearingabaxiallytwolarge,subgloboseyellowmi- interspersed crosporangiacontainingspherical pollen Seed subtropicalbut damp and wetmuchofthe year cones sparse but sometimesgroupedwith2 or Conservation NewlydiscoveredXanthocyparis restrictedto a fewlocalities in at the outermarginsor nearerthe base of is vietnamensis together with mature consistclose leaves, foliagesprays mostlynow in inaccessible sites initially proximity, ing of the upper pairs of greenleaves (bracts) on steep limestoneridges.Logginghas increased withaxillaryovules Maturecones developingin in recentyears and is estimatedto have caused twoyears,green,turning darkor dull brown,subgl- seriousdecline in numbersoflarger,well-growing obose,9-11 x 10-12 mmwhenopen,some + per- trees.This practicemayhave had negativeeffects sistentafterseed dispersal.Bract-scalecomplexes on geneticdiversity.Regenerationis poor due to in (sometimes3) decussatepairsin normally in remainingpopulations.This de- heavycompetition veloped cones (irregularor underdevelopedcones species is CriticallyEndangeredunderthe IUCN are found),valvateto subpeltate(the thirdpair if Red List CategoriesVersion3.1 (IUCN, 2001): CR present+ peltateand 4-5-angled),the lowerpair (B2a-c) Uses This speciesproducesfine,yellow-brown, oblong,all widestdistally,withroundedbut irregular uppermargin;outersurfacesmooth,becoming veryhard,fragrant timber.The superbqualityof in or from a the furrowed wood, 1conjunctionwiththe widespreadderugose radiately prominent, uses ofcupressaceouswoodin traditional 2.5 mmlongumbo(includingthebractapex); inner sirability surfacered-brown markedproximally withwhiteor ofmanykindsin easternAsia, combinedwithslow has made it a highlyprizedtimber.Due to grayseed scars;a smallcolumellapresentorabsent growth, facilitiesand otherfactors,most at the shootapex Ovules to per fertilebract lack of transport has of been tradedlocally the timber in bracts 6-scaled cones seeds max sterile); (upper or per cone, ovoid or irregular, flattened (1.5Paratypes.VIETNAM.Ha Giang:QuanBa, BatDai mmthick),4.5-6 x 4-5 mmincludingtwolateral Protected Area,10Feb.2001, Son,BatDai SonProvincial wings,bodyofseed ? pustulate,lightbrownorred- D K Harderet al DKH 6090 (HN,MO,LE), 12 Feb et al DKH 6224 (HN,K, MO,LE); brown,withwhitehilum at base and micropylar 2001,D K Harder TienHiep, beak oftenpersistentat the apex; seed wings0.5- Can Ty,SingXuoiHo, 12 Oct.1999,Nguyen & P J CribbNTH3594 (HN,MO,LE, V Averyanov L mm wide, thinlymembranous,lightercolored K) Seedlingsnotseen In October1999 a coniferwas foundin North Distribution Vietnam: North Vietnam, Ha Vietnam(N T Hiepet al NTH 3594) thatappeared characterssuggestinga reGiang Province,verylocally in the Bat Dai Son to have morphological mountainsystemnear the Chinese border;altitu- lationshipwithspecies in the cupressoidclade of dinal range1060-1180 m Cupressaceaes.l (Gadek et al., 2000) The specicloud men comparedin particularwith Chamaecyparis Ecology In mixed angiosperm-conifer forestwith the conifersAmentotaxus argotaenia and Cupressus, yetshowedsometraitsnotfoundin (Hance) Pilger,Nageia wallichiana(C Presl)Kun- any knownspecies in theseor relatedgenera,imtze,PseudotsugasinensisDode var.brevifolia (W.C mediatelysuggestingthata new taxoncould have Cheng & L K Fu) Farjon & Silba, Podocarpus been found.The onlyknownspecies in thatgroup and Taxuschinensis(Pilger)Reh- thatare certaintobe indigenousin thispartofAsia pilgeriFoxworthy, der Dominantamongangiospermsare species of (includingmost of Myanmar,NorthLaos, South Acer,Carpinus,Lithocarpus,Quercus,and Ulmus; Yunnan,Guangxi,Guangdong, and Hainan [China]) J.Poissonex are CalocedrusmacrolepisKurzand Fokieniahodgfrequentare Pistacea weinmannifolia Franchet and Platycarya strobilaceaSiebold & insii (Dunn) A Henry& H Thomas.The new co- This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions Volume12, Number2 2002 Farjonet al NewVietnameseGenus 183 C'Tl OL m Fir Ab vietnamensis at Bat Dai Son (photograph -B BranchofX Figure2 -A TreeofXanthocyparis by L Averyanov) vietnamensis withtwodifferent with foliagetypes(photograph by D K Harder).-C Branchletof X vietnamensis matureseed cones (photograph withpollencones (photograph by D K Harder).-D BranchletofX vietnamensis by D K Harder) nifer(Figs 2, 4) appearedto occurverylocallyon limestonekarstridgesknownto have yieldedother narrowendemics,some of whichhave turnedout to be new species, e.g., orchids.Such habitatsare also frequently the refugesof relictconifertaxa, theseconifershave adaptedto because presumably the poor growingconditionsassociated withkarst broad-leaftreespecies could not wherecompetitor follow.The mostconspicuousfeatureof this new conifertaxonis the occurrenceofjuvenile leaves, transitional leaves,and matureleaves in thefoliage of crownsof maturetrees(Figs 2B, 4) These juleaves 15 x 1.8 mmcan alvenile,monomorphic ternateon a branchwithmaturedimorphicleaves 2.5-3 x 1.2-1.3 mm (D K Harderet al 6224) This traitis reminiscentof the foliagein mature trees of Callitrismacleyana F Mueller;however, unlike that species, Xanthocyparisvietnamensis does not producecones on branchletswiththese juvenile leaves In Widdringtonia nodiflora(L.) E This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 184 Novon -j low ?-APO Figure3 nr withmatureseed cones (photograph BranchletofX nootkatensis by A Farjon) Powrie, coppiced (fire-damaged)plants produce distinctspecies; however,its coexistencewithmathe life historyof the tree similarjuvenile leaves usuallyon basal reiterated turefoliagethroughout ofnee- is rarein conifers branches(Pauw& Linder,1997) A mixture Carefulcomparisonofthemorphology ofthenew dle-likeand scale-likeleaves is commonin Juniin where the needle leaves Vietnam with that of moreor chinensis other, discovery Roxburgh, perus representthe juvenile stage because such leaves less similar,species in Cupressaceaeleads to Chaare thefirstto appear afterthecotyledons.Indeed, maecyparis nootkatensis (Fig 3) as themostsimilar in all species of Cupressaceae s.l (Farjon,1998, species This taxon has verysimilarseed cones fromjuvenile (via transitional with4(to 6) bract-scalecomplexes("-12" is cer2001) thistransition in Taylor& Szikforms)to matureleaves takes place in the seed- tainlyan errorforC nootkatensis lings,oftenaccompaniedby a changein phyllotax- lai, 1976), whichare quite distinctfromboth its withwhich is Normally,in most cupressoid taxa, juvenile congenersand fromthoseof Cupressus, has been unitedfromtimeto time leaves notreoccur;whentheyare presentthese Chamaecyparis This (see, e.g., Camus, 1914, fora monographic treatbranchesare oftenthe resultof reiteration seems to be thecase in somespecimensofX viet- ment) There has been debate concerningthe in eithergenus,with namensis(e.g.,N T Hiep et al NTH 3594), but in placementof C nootkatensis others(e.g.,D K Harderetal DKH 6091 holotype, some authorsarguingfor inclusion in Cupressus D K Harderet al DKH 6224) thebranchingpat- based on characters of the ovuliferouscones ternof bothfoliagetypesshowsa normalalterna- (Frankis,1993; Jagel& Sttitzel, 2001) RecentclaIn the knownspe- distic evidence based on moleculardata (matK tionnot suggestingreiteration cies of Chamaecyparisand Cupressusjuvenile gene) gives onlyweak supportforits inclusionin totheseedlingstage.However, Cupressus(Gadeket al., 2000); similarevidenceusleaves are restricted juvenileleafcharactershave been retainedin some ing a combineddata set (matK + non-molecular It is possiblya neotenic data) does not and places it as a sistergroupto cultivarsofChamaecyparis withstronger trait controlledby certain genes that can be Cupressus+ Juniperus bootstrap supin port(Gadeket al., 2000) InclusionofthenewViet"switchedon or off"at different stagesofgrowth severaltaxa By itself,this would be a somewhat namese species in a phylogenetic analysisof Cudata (a full doubtfulcharacterto indicatethe existenceof a pressaceae s.l based on morphological This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions Volume12, Number2 2002 Farjonet al NewVietnameseGenus 185 accountof whichwill be publishedlater)resulted a small columelladevelopsat theshootapex of4in a separateclade forC nootkatensis + X viet- scaled cones,sometimesconsistingoftwoparts.As namensiswithina majorclade distinctfromboth in all Cupressaceaes.l (incl.Taxodiaceae)theovuand Cupressus(Fig 5) Sequencing liferouscone startswithovule developmentborne Chamaecyparis ofDNA ofX vietnamensis has yettobe undertaken on, subtendedby,or surrounded byleaves (bracts); growth Apartfromthemarkedlydifferent ages ofplants the scale formsby secondary(intercalary) is morepronouncedon between C nootkatensisand X vietnamensis in In bothspecies thisgrowth whichjuvenileleaves stilloccur,theleafmorphol- theadaxial side ofthebractsand morerapidin the ogyof bothis verysimilar,bothofjuvenile,tran- second pairof the bract-scalecomplexes,resulting sitionaland mature(Figs 2D, 3, 4) leaves In both in semi-valvatecone scales that remain largely The onlyothertaxonin the cuspecies the matureleaves are markedlydimorphic fusedproximally on facials and shaped laterals) plagio- pressoidclade (Gadek et al., 2000) withsomewhat (differently thistypeis is Tetraclinis; This is commonin all Cu- similarcone development tropicfoliagebranchlets pressaceaewithsuch branchingsystems,causinga morefullydevelopedin thecallitroidclade (Gadek of the leaf-coveredbranchlets et al., 2000) withwhichbothfossil(e.g.,Engelhardt bilateralflattening Growthof branches profoundlydeterminesleaf & Kinkelin,1908) and Recent species (Li, 1953) shape in Cupressaceae (Daguillon,1899; Rouane, of Tetraclinishave erroneouslybeen identified 1973; Offler,1984) Unlike othertaxa with this Whereas in Tetraclinisthe numberof bract-scale markedleaf dimorphism(e.g., Chamaecypariss complexes(2 pairs) seems primaryin origin,in str.,Calocedrus,Fokienia,Platycladus,Thuja,Thu- Chamaecyparisnootkatensisand Xanthocyparis ofabaxial stomata vietnamensis it is almostcertainlyreduced,as evijopsis),in whichthe distribution is asymmetrical, i.e., largelyconfinedto the phys- denced by the occasional thirdpair thatis much iologicalundersideofthelateralleaves and theab- smallerand neverassociatedwiththeovules.Pooraxial side of facials on thatside of the branchlet, ly developed,smallercones ofCupressuslusitanica bothChamaecyparis and Xanthocypar- Millersometimeshave only2 pairs(insteadof3 to nootkatensis is vietnamensis have onlya weaklydevelopedsto- 4, rarely5 pairs) but theyare morepeltate.The In thisrespecttheyresemblecer- ontogenyof the bract-scale complexes and the matalasymmetry and Chatain species of Asian Cupressuswith (weakly) placementof ovules of X vietnamensis be similar but to are adult leaves on nootkatensis branchlets likely (C dimorphic pendulous maecyparis cashmerianaRoyle ex Carriere,C funebrisEnd- were only studiedin detail forthe latterspecies and in Cupressaceaeis strong- (Jagel& Sttitzel, 2001) Seedlingdevelopment licher).Leafdimorphism remain be X to of vietnamensis with moist correlated climates characters ly (i.e., adaptive) seedling monomorstudied Fokienia, (e.g., Chamaecyparis, Thuja); phismof matureleaves resultingin quadrangular Consideringthe evidencebased on morphology, to teretebranchletsis correlatedwith (seasonal) it is verylikelythatthereis a closer relationship withChamaecyparis vietnamensis aridity(e.g., Cupressusarizonica,Juniperussect ofXanthocyparis thanwithany otherRecent taxonin Sabina) Glands on the facial matureleaves of X nootkatensis vietnamensisare less conspicuousthan those of Cupressaceae.We appear to have founda veryinneithernormallypro- teresting relicttaxonon the westernmarginof the nootkatensis; Chamaecyparis duce dropletsof resin Pacificthatbelongsto a lineagedistinctfromboth and Cupressusand is possiblymore Pollen cones ofbothspecies are verysimilar(in Chamaecyparis withthe C nootkatensis slightly larger)with2(or3) relative- ancientthaneitherofthese.A comparison in Chamaecypariss str Late CretaceoustaxonMesocyparis McIver& Basly large microsporangia; North and especially Cupressusthe latterare more nu- inger,whichhas been foundin northwestern Asia (McIver& Basinsmaller(Figs 2D, 4) Americaand northeastern merous(3 to 6) and relatively vietnamensis Seed cones ofXanthocyparis (Fig 2C) ger,1987; McIver& Aulenback,1994) is ofinterand Chamaecyparisnootkatensis (Fig 3) are also est In thisfossilgenusshortdimorphicleaves apsimilar;the moststrikingfeatureis the predomi- pear on opposite(pen)ultimatelateral branchlets, nance of only two decussate pairs of bract-scale and long dimorphicleaves followa series of short complexesin the maturecones In bothspecies an leaves on leading shoots Althoughthe opposite occasionalthirddistalpair develops(or sometimes branchingpatterncontrastswith the alternate onlyone ofthese);althoughthesamplingofX viet- branchingin Recent cupressoidgenera,the alternamensisstudiedis still somewhatlimited(ca 30 nationof twoleaf shapes is also foundin Xanthocones seen) it seems thatthis is somewhatmore cyparisvietnamensis, but withmonomorphic (juvefrequentin thisspecies (Figs 2C, 4) In bothtaxa nile) longerleaves The seed cones ofMesocyparis This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 186 Novon 1\6 cm F cm 1/3 JJ S1/8cm cm 1/3 cm 1cm sF-cjo,, AL' dpC cm 1/2 LVIL vietnamensis Figure4 Xanthocyparis Farjon& Hiep -A Branchletwithscale leaves and seed cones.-B Branchlet withscale leaves and needle leaves -C Branchletwithscale leaves -D Branchletwithneedle leaves -E Detail ofupperand lowerside ofneedleleaf.-F Branchletwithpollencone -G Microsporophyll withtwomicrosporangia -H, I Seed cones, closed and open -J Seeds [A, C, F-J = Harderet al 6091 (HN, K); B, D, E = Harderet al 6224 (K) Illustration by Aljos Farjon This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions Volume12, Number2 2002 Farjonet al NewVietnameseGenus 187 Callitris drummondii Callitris oblonga Callitris macleana nodiflora Widdringtonia Actinostrobus pyramidalis Neocallitropsis pancheri cupressoides Fitzroya Diselmaarcheri chilensis Austrocedrus Libocedrus plumosa Papuacedruspapuana uviferum Pilgerodendron nootkatensis Chamaecyparis vietnamensis Xanthocyparis Microbiota decussata orientalis Platycladus articulata Tetraclinis Calocedrusdecurrens Cupressusbakeri californica Juniperus drupacea Juniperus Juniperus oxycedrus indica Juniperus recurva Juniperus Juniperus phoenicea virginiana Juniperus Cupressusfunebris lawsoniana Chamaecyparis Fokieniahodginsii Thujaplicata Thujopsisdolobrata Metasequoiaglyptostroboides Cryptomeria japonica Glyptostrobus pensilis distichum Taxodium Sequoia semprevirens Sequoiadendron giganteum lanceolata Cunninghamia Taiwaniacryptomerioides "Athrotaxis laxifolia verticillata Sciadopitys Pinussylvestris treesof 523 stepsfoundfroma heuristicsearchofmorphoFigure5 Strictconsensusof 144 equallyparsimonious as logical and anatomicaldata (53 characters)in Cupressaceaes.l and Sciadopitysverticillata, usingPinus sylvestris outgroup.CI = 0.2428; RI = 0.3077; RC = 0.0747 [A fullaccountof thecharactersand the data set will be given of Cupressaceae.] by Farjonin his monograph umbonataMcIver& Basingerare similarto those growthafterpollinationaltersthe relativesizes of of Chamaecyparis nootkatensis and X vietnamensis bractand scale, considerablyinfluencing the size withtwo paired bract-scalecomplexes.However, and shape of these bract-scalecomplexesat mathe positionof seed cones on the foliagebranches turity The seeds offossilMesocyparis and of Chaof Mesocyparisumbonatais quite different and Xanthocyparis are verysimilar.Anespe- maecyparis cially fromC nootkatensis They are not terminal other Late Cretaceous fossil, Chamaecyparis on foliage sprays but appear lower on foliage corpulenta(Bell) McIver(McIver,1994) fromVanshoots,on opposite,veryshortbranchletsaxillary couverIsland, Canada, has leaves moresimilarto to longleaves A lowerpositiondoes also occurin otherRecent species in Chamaecyparis, but again X vietnamensis, but thesecones are notin decus- verysmall seed cones withfourscales This low sate pairs.Earlydevelopmental stagesobservedin number of bract-scale complexes is apparently fossilseed cone specimensofMesocyparis umbon- primitive, as it occurs in all earliest(Cretaceous) ata are essentiallysimilarto thosein C nootkaten- membersof Cupressaceaethusfarknown(McIver, sis (McIver& Aulenback,1994) but in the fossil 1994) thecones remainnotablysmaller,i.e., development We concludefromthe above evidencethatChaafterpollinationof the ovules lags behind.In both maecyparisnootkatensisshould be placed with C nootkatensisand X vietnamensisseed cone Xanthocyparis vietnamensis in a separate genus This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 188 Novon has retainedseveral charactersor Xanthocyparis traitsthatwereevidentin similarcupressoidmembers of Cupressaceae s.l in the Late Cretaceous, but whichhave been lostor replacedby otherfeaturesin moreadvancedmembersofChamaecyparis and Cupressusduringthe Tertiary The following has to be made new combination as a resultof the new taxonomy here proposed: erationit has been consideredbest practiceto effectas minimala change as possible The necesare givenbelow sarynew combinations XCuprocyparis Farjon,nom.nothogen.nov & A B Jackson, Dallimore Forestry Cupressocyparis 11: 1937;Roy.Bot.Gard.KewHand-list ed 4: Conif., 37 1938 Xanthocyparis nootkatensis (D Don) Farjon XCuprocyparis leylandii (A B Jackson& & Harder,comb nov Basionym:Cupressus Dallimore)Farjon,comb nov Basionym:CuD Don, in Lambert,Descr Pinus nootkatensis leylandiiA B Jackson& Dallimore, pressus 2: 18 1824 Chamaecyparisnootkatensis (D Kew Bull 1926: 114 1926 XCupressocyparis Don) Spach, Hist Nat Veg Phan 11: 333 leylandii(A B Jackson& Dallimore)Dalli1841 Cupressusnutkatensis Hooker,Fl Bor more& A B Jackson,Forestry11: 1937; Amer.2 (10): 165 1838 Chamaecyparis nutRoy Bot Gard Kew Hand-listConif.,ed 4: kaensisLindley& Gordon,J Hort.Soc Lon37 1938 TYPE: England Northumberland: don 5: 207 1850 TYPE: Canada BritishCoHaggerstonCastle (cultivated),26 Nov 1925, lumbia: Hecate Strait, Banks Island E J Leylands.n (specimenmarkedD, with coast of NorthAmerica behind ["northwest cones) (lectotype, designatedhere,K) Bank's Island"], 1787, A Menziess.n (holoformula: MCupressusmacrocarpaHartweg Hybrid type,notseen; isotypes,K, MO) ex Gordonx 9Xanthocyparis nootkatensis (D Mom.Acad.Imp.Sci Saint-PeDon) Farjon& Harder[Cupressusmacrocarpa ThujaexcelsaBongard, ser.6, Sci Math.2: 164 1832.Cupressus tersbourg, Hartwegex Gordonx Cupressusnootkatensis americana Trautvetter, P1.Imag.Descr.Fl Russ.1 D Don; CupressusmacrocarpaHartwegex (1/2):12,t 1844.TYPE: U.S.A.Alaska:Baranof Gordon X Chamaecyparisnootkatensis(D s.n.(nodate)(holotype, Island,Sitka,"Dr.Mertens" notseen;isotype, Don) Spach] MO) borealishort.ex Carriere, TraiteGen.Conif.: xCuprocyparis notabilis (A F Mitchell)FarThujopsis 113 1855 TYPE: Unknown, basedon cultivated jon, comb nov Basionym:XCupressocyparis plant notabilisA F Mitchell,J Roy.Hort.Soc 95 (10): 453 1970 TYPE: England.Hampshire: The species CupressusmacrocarpaHartwegex ForestResearchStation,Alice HoltLodge,31 Gordonis consideredto be the male parentof the July1963, A E Mitchells.n (holotype,K) generichybridXCupressocyparis leylandii(A B formula:9CupressusarizonicaGreenevar Hybrid Jackson& Dallimore)Dallimore& A B Jackson; (Sudworth)Little x glabra Xanthocyparis the femaleparent,fromwhichhybridseeds were nootkatensis & Harder (D Don) Farjon collected,is thoughtto be Chamaecyparis nootka- XCuprocyparis ovensii (A F Mitchell)Fartensis(D Don) Spach (Jackson& Dallimore,1926) jon, comb nov Basionym:XCupressocyparis Reversedparentagewas also reportedto have ocovensii A F Mitchell,J Roy Hort Soc curred(Jackson& Dallimore,1926), but boththe 95(10): 454 1970 TYPE: England Hampprotologueand the type specimensat K indicate shire: Forest Research Station,Alice Holt thatthishybridwas firstraised fromseed obtained K) Lodge, 1970, A E Mitchells.n (holotype, in 1888 fromC nootkatensis Two further nothosformula:9 Cupressuslusitanica Miller X Hybrid pecies withmale parentagefromC nootkatensis, nootkatensis 6Xanthocyparis (D Don) Farjon but involvingas femaleparentsCupressus arizonica & Harder Greenevar.glabra (Sudworth) Littleand Cupressus lusitanica Miller, were described by Mitchell Acknowledgments.The authors thank Marc (1970) Under the rules of the Code (Art H6.2; Coode of Kew forhis kind help withthe Latin diGreuteret al., 2000) the name of a nothogeneric agnosis and description,and two anonymousreremarks.Fieldwork hybridis to be combinedfromthe firstpartor the viewersfortheirconstructive wholeof one and the last partor the wholeof the supporting thisdiscoverywas providedby the Naotherparent.The reclassification ofChamaecyparis tional Science Foundation(DEB-9870231 to the nootkatensis in a new genus Xanthocyparis there- MissouriBotanicalGarden,D HarderP.I.) and the forerequiresa namechangeofthehybridgenusof National GeographicSociety (grant# 6733-00, whichit is one of the parents;afterample delib- "Botanical inventory of unexploredareas in Viet This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions Volume12, Number2 2002 Farjonet al NewVietnameseGenus Nam:thenorth"to DKH) This supportis gratefully acknowledged maecyparis Spach und CupressusL (Cupressaceae)und die systematische Stellungvon Cupressusnootkatensis D Don [= Chamaecyparisnootkatensis(D Don) Spach] Feddes Repert.112(3-4): 179-229 of Libocedrusand CuLi, H L 1953 A reclassification pressaceae.J ArnoldArbor.34: 17-34 McIver,E E 1994 An earlyChamaecyparis (Cupressaceae) fromthe Late Cretaceousof VancouverIsland, BritishColumbia,Canada Canad J Bot 72: 17871796 and rela& K R Aulenback.1994 Morphology umbonatasp nov.:Fossil CutionshipsofMesocyparis pressaceae fromthe Late CretaceousofAlberta,Canada Canad J Bot 72: 273-295 & J F Basinger.1987 Mesocyparis borealisgen et sp nov.:Fossil CupressaceaefromtheearlyTertiary of Saskatchewan,Canada Canad J Bot 65: 23382351 Mitchell,A F 1970 A noteon twonewhybridCypresses J Roy.Hort.Soc 95(10): 453-454 C E 1984 Extantand fossilConiferalesof AusOffler, traliaand New Guinea Part 1: A studyoftheexternal of the vegetativeshootsof the extantspemorphology Abt B, Palaeophytol.193: 18-120 cies Palaeontogr Pauw,C A & H P Linder.1997 TropicalAfricancedars Cupressaceae): Systematics,ecology (Widdringtonia, status.Bot J Linn Soc 123: 297and conservation 319 des Rouane, P 1973 Etude compareede la repartition ramifications au coursde l'ontogenesede quelques Cupressacees Tray.Lab Forest.Toulouse T 9(3) 4: 1277 nootkatensis Taylor,S D & Sziklai 1976 Chamaecyparis (D Don) Spach-Yellow Cedar,memberof the family Cupressaceae.Davidsonia7(4): 56-62 Literature Cited Camus,A 1914 Les Cyprns(genreCupressus).[EncycloII.] Paul Lechevalier, p'die 'conomiquede Sylviculture Paris sur les Daguillon,A 1899 Observationsmorphologiques feuillesdes Cupressinees.Rev Gen Bot (Paris) 11: 168-205 H & F Kinkelin.1908 I Oberpliocaine Flora Engelhardt, und Fauna des Untermaintales,insbesondere des Frankfurter Klarbeckens.Abh Senckenberg.Naturf Ges 29(3): [150]151-306 ofCoFarjon,A 1998 WorldChecklistand Bibliography nifers.Royal BotanicGardens,Kew [2nd ed 2001.] or Frankis,M P 1993 NootkaCypress:Chamaecyparis Cupressus?ConiferSoc Austral.Newsl.12: 9-10 Gadek, P A., D L Alpers,M M Heslewood & C J Quinn.2000 RelationshipswithinCupressaceaesensu lato: A combined morphologicaland molecularapproach.Amer.J Bot 87: 1044-1057 Greuter,W., J McNeill, F R Barrie,H M Burdet,V Demoulin,T S Filgueiras,D H Nicolson,P C Silva, J E Skog, P Trehane,N J Turland& D L Hawksworth.2000 International Code of BotanicalNomenclature(St Louis Code) RegnumVeg 138 IUCN 2000 IUCN Red List Categories:Version3.1 Prepared by the IUCN Species Survival Commission and Cambridge,U.K IUCN, Gland,Switzerland, Jackson,A B & W Dallimore.1926 A new hybridconifer.Kew Bull 1926: 113-115 von ChaJagel,A & T Sttitzel.2001 Zur Abgrenzung This content downloaded from 195.78.108.81 on Sun, 15 Jun 2014 04:52:01 AM All use subject to JSTOR Terms and Conditions 189 ... rhombica ,in partedis- laterals slightlylonger than the facials); the facials pressa,angusteangulato-ovata tall carinata,margineminutedenticulatapraeterapicem narrowlyovate-rhombic,keeled, + appressed,... marversusintegra,apice acuto; folialateraliamoderatelonentire towardthe acute giora, conduplicata,basin versus decurrentia,recta vel gins minutelydenticulate to falcata,margineminutedenticulatapraeterapicemversus... juvenile linear linearia,eglandulifera, margineintegra,apice acuto; stoabaxiali juveniliumin vittis2 in superficie leaves,transitional leaves ,and maturescale leaves matafoliorum Juvenileleaves presenton