Additions to Orophea subgenus Sphaerocarpon (Annonaceae): revision and transfer of Mezzettiopsis

23 Pages·2001·3.9 MB·English

by  Leonardía, A A P

BLUMEA 46 (2001) 141-163
Additions to Oropheasubgenus Sphaerocarpon
(Annonaceae): revision and transfer
of Mezzettiopsis
A.A.P.Leonardía & P.J.A. Keßler
Summary
The genus MezzettiopsisRidl. (Annonaceae)isrevised andtransferred toOropheaBlumesubgenus
SphaerocarponKeßler.PhylogeneticanalysisofsubgenusSphaerocarponandMezzettiopsisplaces
Mezzettiopsiswithinthesubgenus.Furthermore,theprevalenceofcommonoroverlappingcharacter
statesandthepaucity ofuniquecharacterstatesnecessitate thetransfer. Twospeciesweretransferred
from Mezzettiopsis. Ofthese,onespecies ofOrophea isreinstated and onenewcombination is
made.Threespecies aredescribed asnew.Intotal,ninespeciesarefullydescribed.Sphaerocarpon
is asubgenusofsmall,tropical,lowland treesdistributedfrom Sulawesiwestwards toSouthIndia.
Itis characterised by havingsepals andouterpetals thataresessile ornearlysessile and moreor
less ovate, with outerpetals thatare twice the size ofthe sepals, inner petals that areclawed,
roughlydiamond-shapedandconnivent duringdevelopmentattheedgesofthelamina,bymiliusoid
stamens,by stipitate,globosetooblongatecarpidia, andbyglabroustopubescentleaves.Akeyto
thefloweringmaterial ispresented.
Key words.Annonaceae,Mezzettiopsis, Orophea, Sphaerocarpon,phylogeny.
Introduction
TheAnnonaceaerepresentabout130generaand2300speciesofpantropical, lowland,
rarelymontane,shrubs, treesorclimbers.KeBler(1988)places thegenusMezzettiopsis
Ridl. inthetribe MiliuseaeHook.f.& Thomson(as Saccopetaleae) together withthe
generaAlphonsea, Miliusa, Orophea, Phoenicanthus, and Platymitra. In this tribe,
only Miliusanowremains toberevised.
The genericdelimitationbetween Mezzettiopsis Ridl.andOrophea Blumehasal-
ways beendifficult.Mezzettiopsis was establishedin 1912by Ridley withthepubli-
cationofthetypespecies,M. creaghii,describedfromaspecimen collectedinTinkayo,
SabahandnamedinhonourofGovernorCreagh. Confusionfirstarose whenin 1913
thesamespecies was describedandpublished asOropheapalawanensis byElmer.In
a treatment ofthe AnnonaceaeofPeninsular Malaysia, Sinclair (1955) informally
pronounced M.creaghiianOrophea,indicatingitscloseaffinitywithO.palawanensis.
WithhisrevisionofOrophea in 1988,KeBlerreaffirmedthegeneric statusofMezzet-
tiopsis, citinginnerpetal-,stamen-,carpel-,ovule-,andseedintegument charactersto
distinguish Mezzettiopsis fromSphaerocarpon, hisnewsubgenus ofOrophea. KeBler's
1) Institute ofBiology, U.P Diliman, 1101 Diliman,QuezonCity, Philippines;
e-mail: [email protected]
2) Nationaal Herbarium Nederland,UniversiteitLeidenbranch,P.O.Box 9514,2300RALeiden,
The Netherlands; e-mail: [email protected]
142 BLUMEA Vol. 46, No. 1,2001
revisionresultedinthesynonymy ofO. palawanensis andM.creaghiiandthesuggest-
edtransferto Mezzettiopsis offivespeciesofOrophea, namely: O. hastata,O. mono-
sperma, O. salacifolia, O. siamensis,and O. torulosa. In 1990,Goel transferred O.
torulosa to Mezzettiopsis in an articleofthe recent rediscovery and recollectionof
thisspecies intheAndamanIslands.
This present revision of Mezzettiopsis transfers the genus to Orophea subgenus
Sphaerocarpon resulting in anewspecies combination,the description ofthreenew
species, andthereinstatementofO. torulosa.Phylogenetic analysis corroboratesthe
transfer. This revision is made as a contributiontothe FloraMalesianatreatment of
theAnnonaceae.
VEGETATIVEMORPHOLOGY
Orophea subgenus Sphaerocarpon consistsofshrubs to smalltrees. Theleaves are
subchartaceousto subcoriaceous.The leaftips areoftenfoldedtoonesideator near
theirbases. Hairs, ifpresent,are simple, usually 1 mmor shorterbutinO. hirsuta and
O. malayana they may reach 2 mm in length. The indumentis usually sparsely to
denselypubescent onleavesand youngtwigs,oftenglabrescent,butinO. hirsuta itis
hirsuteandpersistentonevenoldtwigs. ThelowersurfaceoftheleavesofO. creaghii
andO. malayana characteristically dry darkreddishbrown.Leafvenationiseucampto-
dromous:"withasingle primary vein,thesecondary veinscurvedupwardandgradually
diminishing distallywithin themarginandinterconnectedby aseriesofcross-veins
withoutforming conspicuous marginal loops"following thesystem ofHickey (1973)
andRadfordetal.(1974). Klucking (1986) notes twoleafvenationtypes pertinentto
thesubgenus: 'EarlyPhase' venationdominantwithsmallamounts of'MiddlePhase'
andoccasional'LatePhase' venation,and 'MiddlePhase' dominantwithsmallamounts
of'EarlyPhase'venationinthebaseoftheleafandsometimes 'LatePhase' venation
attheupperthirdoftheleaf.Klucking's findings mayprovetobeinaccurate because
ofthe questionable identitiesofsomeofthespecimens hestudied.
INFLORESCENCES AND FLOWERS
The flowersare arranged in 1-3-, 1-6-,or morethan6-flowered, densely bracteate,
cymose, axillary inflorescences. Supra-axillary inflorescencesare sometimesfound
in O. hastata, while ramiflorousinflorescences are unique to O. monosperma.The
pedicels arearticulatedatthebase.Thebractsareusuallycymbiform butinO. malay-
ana triangular. Triangular bracteoles are found atthe middleofthe pedicels. The
flowerbuds are always ovoid but may be surmountedby theovergrown tips of the
innerpetals. The sepals are valvate, sessile, connate atthebase, deltoid-ovate.The
outer petalsare likewisevalvate, sessileornearly so, free,ovate toslightly obcordate.
The inner petals are valvate, clawed, conniventbefore anthesis at the lateral and
terminalbordersofthelamina,forming aroofoverthe androeciumand gynoecium.
Thelaminaareroughly diamond-shaped withthebordersclearlymarkedby aninverted
V-shaped ridge covered by hairs.The innerpetal tips maybeshortly to largely pro-
tracted, straight to strongly recurved, smoothto very warty. The margins ofthetips
mayalsobeincurved.Inspecies withgreatlyprotracted innerpetal tips, theconnivent
conditionoftheinner petals maynot be very apparent sothat theinner petals may
A.A.P. Leonardfa & P.J.A.Keßler: Revision and transfer ofMezzettiopsis 143
mistakenly appearfree. The innerpetal tips areoften keeledon theirinner surface,
thus making them triquetrous; however,the degree towhichthis featureis manifest
varies. Nectaries are foundatthe laminaofthe innerpetals and range from paired
simple tohookedholes,to single andvariously shaped (see KeBler, 1988forillustra-
tions).Paired, hookednectariesandsinglemoreor lesstriangularnectaries mayprove
to be variants of one another as thehooks need only merging together to formthe
latter. This condition has in fact been observed in some ofthe specimens studied.
Single H-shapedandhippocrepiform nectariesalsoseemtoberelated:hippocrepiform
nectaries withpaired lateralgougesattheircurvedendhavebeenobservedandseem
to be intermediary between hippocrepiform and H-shaped nectaries. The rounded
torus isdensely hairybetweentheinnerpetalsand thestamens. Thealways miliusoid
stamens are sessile or nearly soand number6 ina single whorl, or9 in two whorls
(6+3), or 12 intwo whorls (6+6). Whetheror not thestamens are disposed in series
seems to be partly dependent on thenumberof carpels oftheflower. Staminodes
whichare commonly foundin Orophea subgenus Orophea arealways absentin sub-
genusSphaerocarpon. Stamenconnectiveprolongations neverconcealthetheca.They
are acute, sometimesretuse, erect or recurved. These states vary within the same
species, oftenwithinthesameflower andare seemingly dependentontheproximity
ofthestamen tothegynoecium. Carpel numbervariesfrom3to 18inmultiples of3
andmaybe sparsely hairy to completely glabrous. Thestigma is sessile, globoseto
oblongate,and hairy,sometimesappearing likeapom-pom.An extra-gynoecial com-
pitum(Endress, 1982)appearing likeajelly-likemassabovethestigmas isoftenfound.
Ovules arealmost always 2, but among the species with broad, incurved, strongly
recurved, innerpetal tips, atendency to having morethan2ovules(up to6 inO. laui)
isobserved.
CARPIDIA
ThefruitsofOrophea subgenus Sphaerocarpon areall globoseto shortly oblongate,
andstipitate monocarpswhichvary inthelengths of thestipeandfruiting stalks, the
diameterofthetorus,thesizeofthemonocarps,andnumberofmonocarpsthatreach
maturity. Thefruitingstalks tend tobe longer andthicker thantheflowering stalks.
The sepals are persistent in fruit, not caducous as reported by Van Setten& Koek-
Noorman(1992) butremainattachedeven afterthemonocarpshave fallenoff. The
monocarpsareall smoothbutare oftenfoundshrivelledand thus slightly rugose, a
probable artefactofthedryingprocess. Orophea hirsutais unique inthesubgenus in
having hirsutemonocarps, a curiousconditionconsidering thatthecarpels are com-
pletelyglabrous. Theremainderofthe subgenus has glabrous monocarps.The fruits
ofO. glabraandO. rubraseemtobepeculiarto thesetwo species. At maturity,they
are paired, relatively large monocarps(up to3 cm diam.) withproportionately long
stipes onasmalltorus.
The seeds are globose, half-globose to thick-discoid, filling themonocarps. Van
Setten & Koek-Noorman (1992) reported the following additional fruitand seed
features:fruitwallcrustaceous, c.0.5 mmthick, seedswitharaphethatisflatorshal-
lowly grooved, hilumcircular, ruminationsspiniform to flattenedpegs towards the
raphe, endosperm very hard,glass-like or stony. Christmann(1986, 1987)observeda
rare middle(third) integumentinO. creaghii (sub Mezzettiopsiscreaghii) tobeabsent
144 BLUMEA Vol. 46, No. 1, 2001
inthespecies ofOrophea hestudied,afindingthatinfluencedKeBlertore-affirmthe
identityofMezzettiopsisas agenus. However,Christmannincludedonly onespecies
ofOrophea subgenus Sphaerocarpon in his studiesandthemiddleintegument may
well befoundinother species ofthesubgenus.
PHYLOGENY
A cladisticanalysis wasperformed toelucidatethephylogenetic relationship between
Mezzettiopsis andOrophea subgenus Sphaerocarpon. Alphonsea ventricosaandPlaty-
mitra macrocarpa,both species withinthe sametribeMiliuseae, were usedas out-
groups.Threespecies of Orophea subgenus Orophea (O. hexandra,O. merrillii, and
O.enneandra)werealsoincludedintheanalysis.Threespeciesofsubgenus Sphaero-
carpon, namely, O. multiflora, O. hainanensis, and O. thorelii, were not seen and
characterstatesforthesewerescoredfromliterature(KeBler, 1988) orwereleftambig-
uous. The analysis was executedwith the following characters andstates(for data
matrix, seeTable 1):
1. Maturetwigs: 0= hairy; 1 = glabrous.
2. Petiolecross section:0 =terete; 1 = grooved.
3. Leafbase:0=rounded; I =cordate;2 =cuneate.
4. Leaftip length: 0=< 10mm; 1 => 10mm.
5. Leafmidribuppersurface: 0= grooved; 1 =raised.
6. Flowersper inflorescence:0= <3; 1 =4-6;2 =>6.
7. Outer petal vs. innerpetal shape: 0 =identical; 1 = different.
8. Totallength of innerpetal / clawlength: 0=0; 1 =<0.4; 2=>0.4.
9. Length ofinnerpetal tip /laminalength: 0 =<0.25; 1 =>0.25.
10. Innerpetal tip: 0= short; 1 =protracted.
11. Inner petal tip surface: 0=smooth; 1 =warty.
12. Margins ofinnerpetal tip: 0= flat; 1 =incurved.
13. Inner petal tiprecurvation: 0=absent; 1 =presentin budor atanthesis;
2 =always present.
14. Nectaries: 0=absent; 1 =W-shaped; 2 =paired, horizontalholes;3=single, tri-
angularor paired, hooked;4=single, undulating margin;5 =H-shaped to hippo-
crepiform; 6 = pairedslits orpunctureholes.
15. Stamen number:0=>20; 1 =6; 2=9; 3= 12.
16. Carpel number:0=2 or8; 1 =3;2=6; 3=9; 4= 12;5 = 15;6 = 18.
17. Carpel indument:0= very hairy; 1 =sparsely hairy; 2 = completely glabrous.
18. Stigma shape: 0= globoseto ellipsoid; 1 =oblongate.
19. Ovulenumber:0= >2; 1= 2.
20. Monocarp shape: 0=cylindrical; 1 =spherical tooblongate
Thecladistic analysis was performed withPAUP version3.1.1 (Swofford, 1993).All
characters were treated as unorderedand equally weighted. Multiple states were
interpreted as polymorphic. Aheuristic search was performed (MULPARS option in
effect, additionsequencerandom). Decay indices werealso determined.
Forty-two mostparsimonious cladograms were foundby theanalysis. Theclado-
grams are72 steps long with polymorphic characters accounting for 14 steps(CI =
0.632, RI = 0.851). The strict consensus of themost parsimonious cladograms is
A.A.P. Leonardfa & P.J.A.Keßler: Revision and transferofMezzettiopsis 145
Table 1.Data matrixused for cladistic analysis.
species~~-~~^_ 11 22 33 44 55 66 77 88 99 1100 1111 1122 1133 1144 1155 1166 1177 1188 1199 2200
AA..vveennttrriiccoossaa 00 n0(I0000011 00 00 00 00 00 00 00 0I) 00 00 00 00 0000
PP.. mmaaccrrooccaarrppaa 11 00(10000011 00 00 00 00 00 00 00 0(1 00 00 00 00 0000
0O..hheexxaannddrraa 00 0000000011 1100 00 00 00 00 11 11 1 00 00 0000
0O..mmeerrrriilllliiii 00 00000000 00 110(1 00 00 00 00 11 11 1 00 00 0000
0O..eennnneeaannddrraa 00 (0) 00000() 00 1100 11 00 00 00 22 22 1 00 1100
0O..hhiirrssuuttaa 00 11111 00 00 1100 00 00 00 00 33 11 1 22 111
0O..mmuullttiifflloorraa 1100 7? 0011 1 100 00 00 77 00 33 22 1 77 111
0O..ssaallaacciiffoolliiaa 11000011 001100 00 00 00 00 33 22 55 22 0(1 11
0O.. zzeeyyllaanniiccaa 1100000000 1100 00 00 00 00 33 11 22,,33 22 1 11
0O..tthhoorreelliiii 11227700001100 00 00 77 00 44 11 22 77 111
0O..hhaaiinnaanneennssiiss 112277(0) 00 1100 00 00 77 00 33 11 22 77 111
0O..ppoollyyccaarrppaa I 1 00,,22 00 00 00 1 1 00 00 00 0(1 00 33 11 44 1 00.,11 1 1
0O..uunniifflloorraa II 00,.22 00 0000 1100 00 00 00 00 33 33 22 1111
0O..mmaallaayyaannaa 110010000 11 22 00 00 00 00 00 33 33 11.,22 11 11
0O..tthhoormnssoonniiii 112210000 11 22 00 00 0(1 0(1 00 22 33 22 22 111
0O..rruubbrraa 112210011 1100 00 00 00 0(1 33 33 22 100 11
0O.. ggllaabbrraa 112210011 1100 00 00 00 0(1 33 33 22 100 11
0O.. ddooddeeccaannddrraa 112210000 1100 0() 00 00 00 33 33 44,,66 100 1 1
0O..kkeerrrriiii 1122100001100 100 00 11 53 33 44 100 11
0O.. ssiiaammeennssiiss 1122100001100 100 00 11 55 11,,33 44 100 11
0O..hhaassttaattaa 1 1 22 1 0(1 00 1 1 1 1 1 00 11 66 22,,33 44 100 11
0O..kkiinnggiiaannaa 1 1 00,,22 1 00 11 11 22 1 1 1 00 11 66 33 66 100 11
0O..ccrreeaagghhiiii 110,02,2 100 22 11 221 11 122 6633 66 100 00 1
OO..llaauuii 1122 100 22 11 22 1 11 1 22 66 22,,33 33,,44,,55,,66 100 00 1
0O..llaaoottiiccaa 1122111 2211 22 111122 66 22 55 100 001
0O..mmoonnoossppeerrmmaa 1 1 00,,22 1 11 00 11 11,,22 1 1 1 1 22 66 22 55 22 00 00 1
0O..ttoorruulloossaa 1100111 0011 22 111122 66 22 55 22 00 001
showninFigure 1.Thecharactertransformationseriesandthe decay indicesarealso
presented.
Theanalysis showsthatOrophea subgenusSphaerocarpon isamonophyletic group.
Orophea hirsuta, which shares several character states with Orophea subgenus
Orophea, is shown tobethe sister species oftherest of subgenus Sphaerocarpon. A
polytomy isfoundwheretherestofsubgenus Sphaerocarpon splitsofffromacommon
ancestor with O. zeylanica. The uncertainrelationship of the groups in this clade,
namely, O. thorelii, O. hainanensis,O. polycarpa, andthesistergroupformedbythe
remaining species ofsubgenus Sphaerocarpon furtheralong thetree,exists because
several characterstateswereleftambiguous forO. thoreliiandO. hainanensis.These
species couldnot beexaminedandwereonly scored fromKeBler's(1988) description.
Orophea malayana and O. thomsoniiform sister species and are vegetatively very
similar, both having oblong leaves with long-acuminate leaftips. Moreover, these
two species have relatively short inner petal claws and share the same numberof
stamens andcarpels. Orophea rubra andO. glabraalso form sister species andare
nearlyindistinguishable fromeachotherinbothvegetativeandreproductive characters.
Orophea siamensis and O. kerriiare also shown to be closely related. These two
species seemtodifferonlyinthekindofnectariesthatthey possess andintheprevalent
sizesoftheirleaves,bothcharacterswhichtendtooverlap.Allbutone([iO. salacifolia)
ofthespecies thatKeBlerrecommendedbetransferredtoMezzettiopsis, togetherwith
146 BLUMEA Vol. 46, No. 1,2001
shown.
torulosa O. parsimonious
monosperma O. also
are
laoticaO. most
reversals.
indices
laui O. of
cladogramDecay
creaghiiO. parallel
kingianaO. indicated.
consensus
hastata O. reversals; synapom==orphies;x ￿
are
siamensis O.
strict
analysis
=
kerrii O.
X
Sphaerocarpon:the
dodecandraO.
in
developments;
glabraO. used
rubra O. characters
subgenus
parallel
thomsonii O.
of
malayana O. Oropheaseries
uniflora O.
of
Transformation
polycarpa O. Phyloge1ny.
hainanensis O.
thorelii O. Fig. trees. =
•
zeylanica O.
salacifolia O.
multifloraO.
hirsuta O.
enneandraO.
merrillii O.
hexandra O.
macrocarpa P.
ventricosa A.
A.A.P. Leonardfa & P.J.A.KcGler: Revision and transferofMezzettiopsis 147
O. kerrii, form asister groupto therest ofsubgenus Sphaerocarpon. This groupis
supported by innerpetal characterstates such as protracted andrecurved tips.Further
downtheclademoreinnerpetalcharactersbecomeinformative:areductioninlamina
size, wartinessofthetips,andtheappearanceofsmall, slit-likenectariesare foundin
thecladeofO. hastataanditssister group.Shortening oftheclaw,andeven greater
compaction ofthelaminaare foundinthesistergroupofO. hastata. Broad,incurved,
strongly recurved innerpetaltips as well as atendency to having morethan 2ovules
percarpel, a novelty inthe subgenus andanapparentreversal (a characterstatealso
foundinOrophea subgenus Orophea),, supports thesistergroupofO.kingiana. Oro-
phea lauiispolymorphic instamen andcarpel numbers,hencetheunresolvedrelation-
ship ofthegroupsofits clade.Thelastthreespecies share similarstamen andcarpel
numbersandallhaveleaveswithmidribsraisedontheuppersurface.Orophea torulosa
andO. monospermaare supported as sisterspecies by theirhavingroundedleafbases
andcompletely glabrous carpels.
Thestrictconsensus cladogramexhibitsagreatdegreeofhomoplasy:elevenparallel
developments, threereversals andthreeparallel reversals arefound.Despite this, the
cladogramshowstherelationship betweenMezzettiopsis andsubgenus Sphaerocarpon
andhowtheprevious andpreviously purported membersofMezzettiopsiscompletely
fallwithin subgenus Sphaerocarpon.
KEY AND DESCRIPTIONS
OROPHEAsubgenusSPHAEROCARPON
OropheaBlume subgenusSphaerocarponKeGler, Blumea 33 (1988)62; 34 (1990) 505. —Type
species:Orophea polycarpaA.DC.
Treesorshrubs. Leaves sparsely pubescent to glabrous onlowersurface, glabrous on
uppersurface(lower surface densely pubescent, uppersurface sparsely pubescent in
O. hirsuta). Inflorescences axillary,rarely supra-axillaryorramiflorous.Sepalsconnate
attheirbases, ovate toslightly obcordate,concave. Outerpetals free,ovate toslightly
obcordate,twiceas largeas the sepals,concaveanderectinbud,flattening andspread-
ing at maturity.Innerpetals divisibleintoclaw, lamina, andtip; inner petal lamina
roughly diamond-shaped, connivent during development; innerpetals long-clawed
withlaterally expanded andflattenedlaminaandshorttipstoshort-clawedwithsmall
andthickenedlaminaandprotracted,flattoincurved,straight torecurved, tips.Necta-
ries present. Stamens allequal in size, nearly sessile, 6 in asingle whorl, 9 in two
whorls(6+3), or 12 intwo whorls (6+6).Staminodesalways absent. Carpels3-18in
multiplesof3, sparsely hairytocompletely glabrous, roughly banana-shaped, angular
incrosssection. Ovulesalmostalways 2(rarely upto3-6inspecies withinnerpetal
tips that are recurved frombud to anthesis). Carpidia globose to shortly oblongate,
stipitate, upto3 cm diam.,glabrous (except O. hirsuta). Seeds 1-3,globosetohemi-
spherical.
Distribution—22 species. SIndia,AndamanIslands,Thailand,Laos,Cambodia,
Vietnam, S China, Hainan; Malesia: Peninsular Malaysia, Borneo, Philippines,
Sulawesi.
148 BLUMEA Vol. 46, No. 1, 2001
Notes—AchromosomecountofOrophea spec,andO. creaghii (sub Mezzettiopsis
creaghii, materialnot seen) by Okada(1993) found2n = 18.Christmann(1986)ob-
servedarare middleintegument inseedsofO. creaghii(sub Mezzettiopsis creaghii)
that was absent in the only other species of subgenus Sphaerocarpon he studied
(O.polycarpa). According toWalker(1971) thepollengrainsofOrophea aresolitary,
medium-sized,globose, apolar, radiosymmetric, inaperturate (nospeciesofsubgenus
Sphaerocarpon were studied).
KEY TOTHE SPECIES OFOROPHEA SUBGENUS SPHAEROCARPON
(Thespecies described here areprintedin bold romantype)
la. Mature twigs, densely hirsute or velvety, leafbasecordate, mature monocarps
hirsute.— Indochina,Hainan,PeninsularMalaysia <O. hirsuta
b.Mature twigs generally glabrous, leafbase cuneate to rounded, never cordate,
mature monocarps glabrous 2
2a. Indumentonbractsandbracteolesstraw yellowtowhitish, 1-2mmlong,flowering
peduncles 10mmlong, flowering pedicels 20 mmlong.— PeninsularThailand,
PeninsularMalaysia O. malayana
b.Indumenton bractsand bracteoles reddish- to golden brown, up to 1 mm long,
floweringpeduncles up to8 mmlong, flowering pedicels upto 12mmlong . 3
3a. Tipsofinnerpetals, incurved,recurved frombud toanthesis, atanthesis double-
bent,laminarelatively small(lessthan25%ofentirepetal), laterallycompressed
4
b. Tipsofinnerpetals flat, erect atbudor atanthesis,ifrecurved atanthesis, lamina
relatively large (morethan25% ofentirepetal), flattened,laterally expanded 8
4a. Midribon upperleafsurfaceflator grooved nearbase ofleaf 5
b. Midribon upperleafsurfaceraisednearbaseofleaf 6
5a. Leaftips short-caudate, dryingdarkreddishbrown.— PeninsularMalaysia, Bor-
neo,Palawan,Luzon, Sulawesi 1.O. creaghii
b. Leaftips acute toshort- tolong-acuminate, drying yellowish- to greenish brown.
— Hainan 5. O. laui
6a. Leavesmorethan3.5timesas longaswide,sometimesfalcate,leaftipsnot folded,
inflorescences6- tomore-flowered,carpels sparsely hairy.—Laos
4. O. laotica
b. Leaves less than3.5timesaslongas wide,neverfalcate,leaftips folded,inflores-
cences 1-3-flowered, carpels completely glabrous.—AndamanIslands .... 7
7a. Treelets deciduous,leaves6-10by 3-4cm, inflorescencesramiflorous
6.O. monosperma
b.Treelets not deciduous, leaves 9-16 by 4-7 cm, inflorescences on leafy twigs,
axillary 9. O. torulosa
8a. Stamens9 9
b. Stamens6or 12 11
9a. Laminaofinnerpetals small, compact, less than25%of total length, innerpetal
tips protracted 2. O. hastata
b.Laminaofinnerpetals large, laterally expanded, morethan 30%oftotal length,
innerpetal tips short 10
A.A.P. Leonardia & P.J.A.Keßler: Revision and transfer ofMezzettiopsis 149
10a.Leaves 5-8by 2-3.5cm,carpels 3.— SWChina,Vietnam .. . O. multiflora
b.Leaves 7-11 by 4-5.5cm, carpels 15. —AndamanIslands . 7.O. salacifolia
11a. Carpels 6 or9 12
b.Carpels 12or 18 18
12a. Stamens6 13
b.Stamens12 15
13a.Nectary single withundulating margins. —Vietnam <O. thorelii
b. Nectariespaired, hooked 14
14a.Inflorescences 1-flowered, flowerstalk 1.5cm long, carpels 6.— Hainan . . .
O. hainanensis
b.Inflorescences2- or3-flowered, flowerstalk3 cmlong, carpels 9 (rarely 6).—
S India O. zeylanica
15a.Leaves narrowly ovate, nectariespaired,horizontalslits, carpelscompletely gla-
brous.— SW India O. thomsonii
b.Leaves lanceolateto elliptic, nectary single, horizontally broadly triangular or
paired, hooked,carpels sparsely hairy 16
16a.Leaves 3-6by 1.5-2.5cm,inflorescences 1-3-flowered,stigmas longoblongate.
— SW India O. uniflora
b.Leaves 4-11 by 2.5-4 cm, inflorescences 4-6-flowered, stigmas globose to
ellipsoid 17
17a. Nectary single, horizontally broadly triangular. — Borneo O. rubra
b.Nectariespaired hooks.—Philippines O. glabra
18a.Innerpetal laminasmall, laterally compressed, less than25%of totallength .
3.O. kingiana
b.Innerpetal laminalarge, laterally expanded, greaterthan30% oftotallength
19
19a.Nectariespaired hooks or single and horizontally broadly triangular 20
b.Nectary single, hippocrepiform or H-shaped 21
20a.Leaf tips acute to acutely acuminate.— Sri Lanka, Andaman Islands, Indo-
china,Hainan, PeninsularMalaysia, Lesser SundaIslands(Flores, Sumba) . .
O. polycarpa
b.Leaftips caudate,falcateor foldedto appearfalcate.— Borneo
O. dodecandra
21a.Leaves 8.5-16.5by 3.5-6.75cm, nectary H-shaped. —Thailand
8.O. siamensis
b.Leaves 7-11.5 by 3-4.5 cm, nectary hippocrepiform. — PeninsularThailand
O. kerrii
1. Orophea creaghii (Ridl.) Leonardía& Keβler, comb. nov. — Fig. 2, Map 1
MezzettiopsiscreaghiiRidl.,KewBull. (1912)389;SarawakMus. J. 1 (1913)97;KeBler,Blumea
33(1988)62.—Type:Creaghs.n.(holoK; isoBM,barcode000547317),BritishNorthBorneo,
Tinkayo,April 1895,fl.
OropheapalawanensisElmer,Leafl.Philipp.Bot.5(1913) 1721;Sinclair,Gard. Bull.Straits Settlem.
14(1955)398.—Type:Elmer 12858(hololectoNY, barcode 0026165,designatedhere;isolecto
BM,BO, G, GH,K,L,P,U, US),Palawan,Puerto Princesa, denselyforested slopesalongthe
Balsahan River,3-1911,fl
150 BLUMEA Vol. 46, No. 1, 2001
Fig. 2. Orophea creaghii(Ridl.)Leonardia & Keßler. a.Habit; b. flower;c.sepal; d.outerpetal;
e. inner petal showingnectaries;f. stamen; g.carpel [a: SAN76985 (G. Shea;Aban), L;b-g;
Bogorhort.bot. XX-D-124 collectedby Keßler].