An analysis of the variation within Cratoxylum arborescens (Clusiaceae) in Malesia

9 Pages·1997·1.1 MB·English

by  Church, A C

BLUMEA 42 (1997) 397-405
An analysis of the variationwithinCratoxylum
arborescens (Clusiaceae) in Malesia
A.C.Church & P.F. Stevens
The ArnoldArboretumofHarvardUniversity, 22Divinity Avenue,Cambridge,MA02138,U.S.A
Summary
The variation within thewidespreadWestMalesian speciesCratoxylumarborescens(Vahl)Blume
wasanalyzed andthree distinctivevariants were found. Theseare recognizedformally asvarieties,
var.arborescens,var.miqueliiKing, and var.borneense A.C.Church & P.F.Stevens. Akey and
full descriptionsaregiven.The characters employedto delimitthe three taxa arediscussed and the
selection ofvarietalrank isjustified.
Introduction
Themostrecent revisions ofCratoxylum Blumeare by Gogelein (1967) andRobson
(1974). Gogelein recognized six species, and observed substantialvariationin the
leafmorphology ofC. arborescens(Vahl) Blume.He notedthatC. cuneatum Miq.,
laterreducedby King to C.arborescens var. miquelii,was only "anextreme para-
morph connectedwith the averagepopulation by many intermediates"(Gogelein,
1967:473), and was marked by its ratherslenderpetioles, elliptic leavesand long
acumen. Corner(1939) had earliersuggested thatfoliarvariationin C.arborescens
was correlatedwithhabitat:arobustformgrewin lowlandswamps inIndochinaand
WestMalaysiagenerally, whileamore gracile form, var. miquelii,was foundonhill-
sidesandmountainridges in Sumatra, Malaya, andBorneo. Althoughhe foundthat
thetwo forms couldbewellcharacterizedby featuresoftheleaves, includingpetiole
widthandshape ofthelaminaapex,hebelievedthattherewereintermediatesandthat
inbothflowerandsizeofthetree therewere no differences.
TwomodesofgrowthofC. arborescens thatseemedto correlatewith thevaria-
tionjustmentionedwere notedinthecourse ofstudying growth patterns intheClus-
iaceae, andthispromptedfurtherexaminationofthe problem. In somespecimens all
theinternodesimmediately belowtheinflorescenceareapproximately equal inlength
andhaveassociatedleavesthatatmostbecomegradually reducedinsize towards the
inflorescence(Fig. la), andthelowestbranchesoftheinflorescenceareoftensub-
tendedby almostfull-sizedleaves. Otherspecimens haveoneormore very shortin-
ternodesimmediately belowtheinflorescence; theselack associated, expanded leaves
(Fig. lb), although there maybereduced leavessubtending the lowestbranchesof
theinflorescence.This distinctioncanbephrased inaratherdifferentway: inthefirst
formtheinnovationconsistsofaninflorescencewhichisborne terminally onaleafy
axis, whilein the second theinnovationbearing theinflorescenceconsists only of
scaleleaves and then amore or less branched, flower-bearing axis. Elsewhereon
plantsofbothformsthere areterminalbuds with scales. Although Ruthvan Crevel
398 BLUMEA Vol. 42, No. 2, 1997
Fig. 1.Cratoxylum arborescens (Vahl)Blume. Flowering twigshowing internode lengthand bud
scales. — a. C. arborescens var.miquelii(FRI 1139),lacking scars below the inflorescence; b. c.
arborescens var. arborescens(SAN91289),with bud scalesbelow the inflorescence. Arrow head=
lowestbranch ofinflorescence;arrows=budsin theaxilsofdeciduousreduced leaves (scales).
(in Gogelein, 1967: fig. 7aand inRobson, 1974: fig. 8a) illustratedC. arborescens
with scale leaves immediately below the inflorescence(see also Stevens, 1990),
Robson himselfmade no mentionofbudscales in his account;andthe drawing ac-
companying the original description of Hypericum arborescens Vahl omits such
scars (seeVahl, 1791:t. 43); such variationhas not been usedtaxonomically before.
All species havebuds withscales, as doestheclosely relatedmonotypic genusEliea
Cambess., fromMadagascar (Baas, 1970).
RESULTS
To understandthe variation, herbariummaterial from theArnold Arboretum (A),
Gray Herbarium(GH), HerbariumBogoriense (BO), and Rijksherbarium (L) was
examinedand 169particularly intactspecimens werestudied inmore detail.Charac-
ters fromtheflower andfruit, takenfromrehydrated herbariummaterial, wereexam-
ined,as were several vegetative characters- petiole width, laminalength andwidth,
acumen length, and venationprominence.
Belowweshowhowthecharactersof leafscar,internodelength,petiole width,
andlaminaapex canbeused tocircumscribe threeinfraspecific entitieswithin Crat-
oxylum arborescens. Within the area from Myanmar to the Banka Archipelago the
two formsobserved by Cornercanberecognized, whileintheislandofBorneothere
A.C.Church & P.F. Stevens: Variation within Cratoxylum arborescens 399
arealso two forms, the 'lowlandMalayan' formofComerandanundescribedform
which combines characters ofCorner's two forms.The 'lowlandMalayan' form,
whichwerecognize belowas C. arborescens var. arborescens, isrobust inhabitand
canberecognized byits coriaceouslamina,winged petioles andshortenedinternodes
belowtheinflorescence.Itis distributedfromMyanmartoSumatraandBorneo.The
othertwo entitiesare gracileandhavea chartaceouslaminaandslenderpetioles. They
differin growth pattern, ecology and geography. One, corresponding to Corner's
gracile form, isrecognized below as C. arborescens var. miquelii;itgrows athigh
altitudesalong mountainridges and hillsides, andisrestrictedto SumatraandPenin-
sularMalaysia. Theother, C. arborescens var. borneense, grows ata variety ofalti-
tudesandisknown only fromBorneo.
Themostobvious quantitative characters separating thesethreevarietiesare inter-
nodelength,petiolewidth,andthelengthoftheacumenonthelamina(Fig. 2).When
thesecharactersareexamined inthecontextof geography, andthe divisionofthe
specimens intoa groupwith shortenedinternodesimmediately below theinflores-
cenceanda grouplacking suchinternodes, astriking setofcorrelationsis seen (Fig.
2-5). Although atonelevel- thatis, whengeographical considerationsareexcluded
- thereappearstobea continuumofvariation(Fig. 2),thegroupofspecimens with
broadpetioles, short acumen, and scars (var. arborescens) formsa coherentgroup
easily delimitedfromtheothers(Fig. 3-5). They arealso distinguishedby less easi-
ly quantifiable differences- theleaves generally have inconspicuous finevenation
andthelowersurfaceofthelaminaisoftenglaucous. Thesespecimens, foundfrom
Fig. 2. Variation inlamina apex lengthandpetiolewidth inCratoxylum arborescens (Vahl)Blume:
var.arborescens(+), var.borneense(￿),var.miquelii(□).
400 BLUMEA Vol. 42, No.2, 1997
Fig.3.Variation in petiolewidth in Bornean specimensofCratoxylum arborescens var.borneense
(■) andvar.arborescens (□).
Fig.4.Variation in petiolewidthin MalaysianandSumatran specimensofCratoxylum arborescens
var. miquelii(■) and var.arborescens(□).
A.C. Church & P.F.Stevens: Variation within Cratoxylum arborescens 401
Fig. 5.Variation in petiolewidthin the varieties ofCratoxylumarborescenswith bud scalespresent
immediatelybelowtheinflorescence;var.borneense (■), var. arborescens (□).
MyanmartoBorneo, tendtogrow almostexclusively atlow altitudesandareoften
recordedasbeing acomponentofpeat-swampforests. However,S20124(1070 m),
S34844(1100 m), and Jacobs5555 (750-900 m), all from Northwest Borneo,
werecollected athigher altitudes, yet were not recordedas growing in kerangas or
otheracid vegetation types. Specimens otherwiseassignable to var. arborescens may
very occasionally lack scars, e.g. King 6152,8610, fromtheMalay Peninsula.
Thespecimens withnarrower petioles and longer acumen canbesubdividedinto
two groups. Cratoxylum arborescensvar. borneense isknownonly fromBorneo. It
growsoverawidealtitudinalrangeandhasshortenedinternodesimmediately below
theinflorescence(Hallier1065, apoorspecimencollectedinKalimantan, lacks these
internodes). The othertaxon, Cratoxylum arborescens var. miquelii, is known only
fromSumatraand theMalay Peninsulaand grows atmoderateelevations; itlacks
these shortenedinternodes. Otherthaninternodelength and geographic distribution,
thetwoare indistinguishable (Fig. 2).Thereare, however,two Sumatranspecimens,
bb5770(collected at 880 m)andbb 2929(800 m), whichboth havebudscalesbe-
lowtheinflorescence.However,they differfromvar.arborescens intheirvenation
andintheabsenceofa glaucous undersidetothe lamina(as well as inthealtitudeat
whichthey were growing), andthey seem tobecorrectly assigned to var. miquelii.
Othercharacterswereexaminedto seewhatvariationthey showedwithin Crat-
oxylum arborescens.Petal appendages, staminodial fascicles, and seedshape have
been previously usedto distinguish betweenotherspecies inthegenus. Quantitative
datawere taken from 32specimens to see ifthefirst two characters, and also seed
size, showedany correlationwiththepresenceorabsence ofbud scales. Variationin
thesecharacters didnot correlatewiththatoftheothercharactersdiscussed.
402 BLUMEA Vol. 42, No. 2, 1997
CONCLUSION
Ourconclusionsarebasically inline withCorner'sobservationsmadeover 50 years
ago. We can provide more detailedjustification forour position, partly because of
thediscovery ofvariationinthetimingofinflorescencegrowth. Thedecisionto em-
ploy infraspecific rank forthe threetaxa was madebecausethevariationwithin C.
arborescens isratherrestrictedand not entirely discrete. Although thereare strong
geographical, altitudinalandecological components to thevariation, we haveseen
thatalltaxa show exceptions andoverlap here,too. Noparticular significance should
bereadinto our choiceof variety(as againstsubspecies) fortherankofthetaxa we
recognize (see Hamilton& Reichard, 1992, for asummary ofthe usage ofthese
ranks). Indeeda question moreinteresting than 'whatrank arethesetaxa?',andone
whichwould helpclarify the variationpattern wehave described,is, 'are thesetaxa
eversympatric?'.
Fromfieldlabelsthereis noindicationthatthethreevarietiesdifferin featureslike
tree sizeorbark.Nevertheless,although werecognize thethreetaxa as varietiesonly,
collectionsfrom Borneocomeclose tosuggesting thatthetwoBornean varieties, at
least, may bereproductively isolated.Kostermansmadetwo collectionsonNunukan
Island, NortheastBorneo,whichrepresentboththesevarietiesandwereobtainedat
lowaltitudes:Kostermans 9016(var. borneense) andKostermans9196(var.arbor-
escens). Fieldstudiesare clearly needed.
OfthethreevarietiesofCratoxylum arborescens, var. arborescensis mostsimilar
in vegetative and inflorescence characters, and also in habitat,to C. glaucum, the
otherspecies inthesectionIsopterygium, whichalsohasshortenedinternodesimme-
diatelybelowtheinflorescence.Futurestudieson C. arborescenswill needtoinclude
C. glaucum in theirpurview ifweare tounderstandthesystematics andrelationships
of thispartofCratoxylum.
Thereis onefinalpoint. Discontinuitiesin variation,likecharacterstates,are not
absolute; theirexistence and detection depends onthecontext inwhich variationis
analyzed. For our problem, itisperfectly appropriate torestrictthe geographical ex-
tentofindividualanalyses; discontinuitiesthatappearatalocalscaleareofpotential
biological interest, even ifthey disappear atanalyses thatencompass abroadergeo-
graphical scale.
KEY TO THE VARIETIES OFCRATOXYLUM ARBORESCENS
la. Internodesimmediately belowinflorescencemuchshortened 2
b. Internodesimmediatelybelowinflorescencenot shortened.[Petiole 0.8-1.3mm
wide, rarely winged; laminachartaceous,elliptic;apex acuteto acuminate, acu-
men6.0-17.0mm long. Throughout PeninsularMalaysia andSumatra.]
c. var. miquelii
2a. Petiole 1.3-2.6mmwide, oftenalmost winged; laminacoriaceous, broadly ob-
ovate-oblong or obovate-elliptic; apex shortly cuspidate to acuminate, acumen
1.0-10.0mm long. [Myanmar (Tenasserim) toSumatra, Malaysia andBorneo.]
a. var. arborescens
A.C. Church & P.F.Stevens: Variation within Cratoxylum arborescens 403
b. Petiole0.8-1.4mm wide,rarely winged; laminachartaceous, obovate-oblong to
elliptic; apexacute-acuminate, acumen 4.5-17.0mm long. [Restricted to Bor-
neo.] b. var. borneense
Cratoxylum arborescens(Vahl) Blume
Cratoxylumarborescens (Vahl)Blume,Mus. Bot. Lugd.Bat. 2(1852)17.—Hypericum arbores-
censVahl,Symb. 2(1791)86, t.4 3. — Type: Koenig s.n., 1778 (cf.: Fl. Males. I, 1, 1950,
288)(holoC,n.v.),Malaya; seeRobson,Fl.Males. I,8(1974) 11,for synonymy.
a. var. arborescens
Internodesimmediately below theinflorescencemuchshortened, budscales present
there;petiole 1.3-2.6mmwide,oftenalmostwinged; laminabroadly obovate-oblong
or obovate-elliptic, coriaceous, drying yellowish brownto darkbrown; apex shortly
cuspidate toacuminate,acumen 1.0-10.0mm;reticulationabove barely visible.
Distribution—Myanmar, Thailand, PeninsularMalaysia, Sumatra, andBorneo.
Ecology —Peatandfreshwaterswampforests, kerangas; 10—100(—1100) m alti-
tude.
b. var. borneenseA.C. Church& P.F.Stevens, var. nov.
Avarietatibus aliis C. arborescentiincicatricibus infrainflorescentesproximepraeditiset
petiolisgracilibus0.8-1.4mm latis,differt.—Typus:Kostermans 4335(holoA;iso BO),
Kalimantan,Sg. Wain region,N ofBalikpapan, 10m,Oct. 1950.
Internodesimmediately belowtheinflorescencemuchshortened, bud scales present
there;petiole 0.8-1.4mmwide, seldomwinged; laminaobovate-oblong toelliptic,
chartaceous, drying golden yellowto greenish brown; apex acute to acuminate, acu-
men4.5-17.0mm;reticulationaboveslightly visibletoprominent.
Distribution—Borneo.
Ecology —Primary forest, including hillsidesandridges ofmountains, sometimes
peatandfreshwaterswampforests, 10-1200(-1900) m altitude.
Note—Afew specimens withaxillary inflorescenceareknown, e.g.D. G. Frodin
& O. Ismawi2066, Sarawak. Clemens40463was collectedfrom 1830mon theMt.
Kinabalumassif.
c. var. miquelii King
Cratoxylum arborescens var.miquelii King, J.Asiat. Soc.Bengal 59,ii (1890) 146.—Cratoxy-
lum cuneatum Miq.,Fl. Ind.Bat. I, 2(1859)517. —Type: Teijsmann HB 636 (holoU, n.v.;
iso L,fragm.),Sumatra,nearLoeboe,Sikkeppeng.
Internodesimmediately belowinflorescencenot congested, bud scalesabsent there;
petiole0.8-1.3mmwide,seldomwinged; laminaelliptic, chartaceous, dryinggolden
yellow togreenish brown; apex acutetoacuminate, acumen6.0-17.0mm;reticula-
tionaboveandbelowraised.
Distribution—PeninsularMalaysia, Sumatra.
Ecology —Primary forestonhillsidesandmountainridges, 300-1500maltitude.
404 BLUMEA Vol. 42, No. 2, 1997
ACKNOWLEDGEMENTS
Wewould like tothank S.J. Davies forassisting with thegraphs and G.Romero for hisexpertise
and aidwith thephotographs.Assistance fromthecurators atBO andLmade itpossible forthede-
tailedstudies needed here.
REFERENCES
Baas,P. 1970.Anatomical contributions toplant taxonomy. I. Floral and vegetative anatomy of
Eliaea from MadagascarandCratoxylumfrom Indo-Malesia (Guttiferae).Blumea 18: 369-391.
Blume,C.L. 1849-56 [-1857]. MuseumBotanicum Lugduno-Batavum.Leiden.
Corner,E.J.H. 1939.Notes onthe systematyanddistribution ofMalayanphanerogams.Gard.Bull.
Straits Settlem. 10: 21-36.
Gogelein, A.J.F. 1967.A revision ofthe genus Cratoxylum Blume (Guttiferae).Blumea 15:471-
473.
Hamilton,C.W.,& S.H.Reichard. 1992.Currentpractices in theuseofsubspecies, variety, and
formain the classification ofwild plants.Taxon 41: 485-498.
King,G. 1890.Materials fora Floraofthe MalayanPeninsula. J.Asiat. Soc.Bengal59, ii: 113—
206.
Miquel,F.A.W.1855-1859.Flora vanNederlandsch Indie.Leipzig.
Robson,N.K.B.1974.Hypericaceae.In:Flora Malesiana I,8: 11-12.
Stevens, P.F. 1990. Nomenclatural stability, taxonomic instinct, and flora writing- arecipe for
disaster? In:P.Baas etal. (eds.), ThePlant Diversity ofMalesia: 387-410. Dordrecht,Nether-
lands.
Vahl,M. 1790-1794. SymbolaeBotanicae. Copenhagen.
List of collections
Numbers between bracketsrefertothe threeacceptedvarieties ofCi ratoxylumarborescens: C. arbor-
escensvar.borneense (1),var.miquelii(2)and var.arborescens (3).
A 361 (1),368,490, 1035, 1305, 1331, 1345, 1608,4073 (3)—Afriastini,J.J. 1009 (1), 9989
(3) —Anderson 4238 (3),4353, 12579(1), 13257 (3)—Ashton 21346(3).
bb 2866,2897,2919,2929,3986,5166,5679,5770 (2),6188 (3), 6339,6398 (1),6530(2),7062,
7090, 7131 (1), 7389 (2), 10231, 10483 (1), 13925, 13945 (3), 15799,16057,16058, 16064,
16474 (1), 17778 (1), 17830,18185 (3), 18204(1),18266(3), 18366(2), 18640,20609,20722,
21169, 21194,21233 (1),23001, 23002,23003,23020,23023, 23880(3), 24627, 24667 (1),
25252, 25786 (3), 26166, 27011 (1), 27592 (3), 27725 (1), 28533, 28678, 29156,30085,
30179,33094, 33096 (3), 35356(1) —Beguin304,484 (3) — BNB,see SAN—BRUN 639
(3)—Buwalda6387,6742 (3),7691 (1).
Clemens 26882,40463 (1)—Coode,M.J.E.6817 (1).
Forman 444 (1)—FRI 0722 (3), 1139, 1285 (2) 2803 (3),6175 (2), 6663, 8902 (3), 14243 (2),
15615, 15865, 16875 (3), 17084 (2), 18436 (3), 20423 (2), 24875 (3)—Frodin & Ismawi,
2066 (1) —Fuchs 21290(2), 21346(3).
Grashoff46(3)—Griffith 839(3).
Hallier 2975 (1) —Hansen 1308 (3)—Hou 569(1).
Jacobs 5555 (3).
Kadim & Noor 155,312 (3)—Kandalis 9045 (1)—Kasim 748 (2)—KEP5248,24875,30494,
32561 (3), 35937 (2),36387, 36480,36990, 51940 (3), 76498 (2),80178, 80230 (3), 80424
(1), 98391,99440 (3)—KeBler et al. 524 (1) —King 3041,5251, 6072, 6152,8610, 10105
(3)—Kirkup et al. 276 (1)—KL 3016 (2)—Kostermans 17(1), 92, 112(3),4335,6484,
6683,7105,9016 (1), 9196, 10354, 12889 (3)—Kostermans & Anta 385, 569, 590, 592 (3)
—Kostermans& Sabana 9(2).
A.C. Church & P.F. Stevens: Variation within Cratoxylum arborescens 405
Lorzing 17223(2).
Maxwell 87-532 (3)—Melegrito1566,2602 (3),3301 (1)—Mikil 37751 (1).
Native Collector 378,1132,5141 (3)—Nedi 749 (1)—Niyom 829(3).
Ogata 10155(3).
Rahmat si Boeea 7976,8037,9218, 9276(2)—Rahmat si Toroes 2012, 2143,3015,4859 (3) —
Ridley 3611a(3).
S 12841 (3), 14989 (1), 20124 (3),23921,28155,28441, 30369 (1),33085,34517,34844,43013
(3)—SAN 2602, 15855,15868, 19192,23821 (3), 25594,25821 (1), 27305,27731, 28170
(3),28790,30283(1),31260(3),32272(1),34564 (3),35159,40214,40356(1),50661,51729,
55280 (3),55764 (1), 61320 (3), 65113 (1),65329,72307,72491,72536,80760,80814,84062,
84247,84278,84326 (3), 84800,89495,89499, 90091 (1),91289 (3)—SFN 3470 (3),3705,
11392 (2), 19289 (3), 21428 (2), 34002,34759 (3),35679 (1),36280,36387, 36754, 37347
(3)—Shah 891 (2) —Shea 26259, 27541 —Soepadmo 127 (3)—vanSteenis 9984 (2)—
Stone9601, 13757 (2)—Stone & Weber 15954 (2).
Tandum2826 (3)—Teijsmann HB 636 (2)—Thorenaar 9T1910,9T 1P 177 (3).
vanValkenburg 1239(3)—deVoogd 1132(2).
Whitmore3316 (2)—deWilde 15138 (2)— Winkler 2919(1)— Wong918 (3).