DSpace at VNU: A new genus and species in Cupressaceae (Coniferales) from northern Vietnam, Xanthocyparis vietnamensis

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. 2 (Summer, 2002), pp. 179-189
Published by: Missouri Botanical Garden Press
Stable URL: .
Accessed: 15/06/2014 04:52
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
/>
.
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

.

Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to
Novon.



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.

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.
D. K. Harder
Arboretum,UC Santa Cruz, 1156 High Street, Santa Cruz, California 95064, U.S.A.

Phan Ke Loc
Departmentof Botany,Vietnam National University,Hanoi, Vietnam.
L. Averyanov
Komarov Botanical Instituteof the Russian Academy of Sciences, Prof. Popov Street 2, St.
Petersburg, 197376, Russia.

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 2 (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 5 per bract;seeds usuallyfewer,con- al leaves. Foliage sprays withjuvenile leaves bushy,
centratedon the upperpair (if 2 pairs) or middle sparsely branched, ultimate branchlets 20-50 mm
withtwothin long, not flattened. Foliage sprays with mature
pair (if 3 pairs) of scales, flattened,
lateralwings.Seedlingswith2 cotyledons,
followed leaves flattened,with rounded outline; leading foliage branches quadrangular to terete, with ca. 4
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, 3
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

.4

.

"A"
(H
N)
of
Qw.
Vi0
rnip
stennsl,44
......

Fu

A

a,,9

nn.

M

i rowr;

8

;

f..

.

,

Det
A.
Lr
2001
Kew)
Farjon
(RBG

.1

.

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. 1 phytesand as epiphytes.The limestoneridgeson
mm,oval-terete;microsporophylls
X 1 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 3
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 2 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 2 (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 1 to 3 per fertilebract lack of transport
has

of
been tradedlocally.
the
timber
in
bracts
6-scaled
cones
seeds
max.
sterile);
(upper
8 or 9 per cone, ovoid or irregular,
flattened
(1.5Paratypes.VIETNAM.Ha Giang:QuanBa, BatDai
2 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.
1 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 do 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

1 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: 3. 1937;Roy.Bot.Gard.KewHand-list
ed. 4:
Conif.,
37. 1938.

2. Xanthocyparis nootkatensis (D. Don) Farjon

3. 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: 3. 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. 7. 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.: 4. 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- 5.
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