Revision of Alyxia (Apocynaceae). Part 2: Pacific Islands and Australia

93 Pages·2002·22.3 MB·English

by  Middleton, David J

BLUMEA 47 (2002) I- 93
Revision of Alyxia (Apocynaceae).
Part 2: Pacific Islands and Australia
David+ J. Middleton
Nationaal Herbarium Nederland,Universiteit Leidenbranch,P.O.Box 9514,
2300 RALeiden,TheNetherlands
& Harvard University Herbaria,22Divinity Avenue,
Cambridge,MA02138,USA1
Summary
The genusAlyxia is revised forAustralia and the islands ofthe Pacific Oceanas the second and
finalpart ofacompleterevision ofthegenus.39 speciesarerecognisedforthis areaofwhichthree
are newtoscience, twoare newcombinations and oneis anew name. 14 species arefound in
Australia and its offshore territories,21 species in New Caledonia and theLoyalty Islands,and
sevenspecies in the otherislands ofthe Pacific. Thereis relatively littleoverlapbetween regions:
two speciesarefoundinMalesia andAustralia;onespeciesis foundinNewGuinea andtheSolomon
Islands; onespecies in Vanuatu and New Caledonia;and A.stellata is found in Australia, New
Caledonia andinto the Pacific asfaras Hawaii andHenderson Island. Keys anddescriptions for
the species aregiven.Alyxia stellata and A. tisserantii are particularly variable and in need of
further study. Thenewspecies areA.evansii D.J.Middleton,A..solomonensis D.J.Middleton and
A.veilloniiD.J.Middleton;thenewnameisA.mucronataD.J.Middleton;andthenewcombinations
areA.poyaensis (Boiteau)D.J.Middleton andA. tropica(P.I.Forst.) D.J.Middleton.
Key words:Alyxia,Apocynaceae,Australia,New Caledonia,PacificIslands,taxonomic revision.
Introduction
Thispaperis thesecondpartofatwopartrevisionofAlyxia R.Br.intheApocynaceae.
Adiscussionofthe historyofthegenusandofthecharactershasbeengiveninPart 1
ofthisrevision (Middleton, 2000). Inthat paperthe species fromAsia andMalesia
were presented. This part ofthe revision includes allthe species inAustralia, New
Caledoniaandtheother islandsof thePacificOcean.Thislastareaextendsfromthe
SolomonIslands andtheMarianasnorthwardstoHawaii,southwards toLordHowe
Islandandeastwards to HendersonIsland.
NewCaledoniastands out as anislandwith aparticularlyhigh numberofspecies
inarelatively small area.Allbut two species fromNewCaledoniaandthe Loyalty
Islandsareendemic. The islandis also notableforthe large variationinvegetative
charactersforanumberofspecies.TherehavebeenseveralpapersonthegenusAlyxia
inNewCaledonia:somehavesimplybeendescriptionsofnewspecies orenumerations
ofspecies ineitherAlyxiaorthegeneric synonymGynopogon (VanHeurck &Miiller-
Argoviensis, 1870, 1871;Baillon, 1889b;Planchon, 1894;Schlechter, 1906;Guil-
laumin,1911;Moore,1921;Daniker, 1933;Guillaumin, 1957),othershavebeensome-
1) Current address.
2 BLUMEA Vol. 47, No. 1,2002
whatmore extensive (Guillaumin, 1941, 1948;Boiteau& Allorge, 1979). Someof
the combinationsinGuillaumin(1911) are not validly transferred fromGynopogon
to Alyxia andthe correct combinationswere madein laterpapers. Themostrecent
andextensivework on theApocynaceae ofNewCaledoniawasby Boiteau(1981)in
which 31 species ofAlyxia were recognised. He also described a numberofnew
sectionsinthegenuswhichhavenotbeenmaintainedhere(see discussioninMiddleton,
2000). In my work I have synonymised many of the species he recognised, and
somewhatalteredthestatusofother taxa,resulting intherecognition of21 species.
Brooks et al. (1981) discuss the manganeseconcentration ofseveral species of
Alyxia inNewCaledoniaandsuggest thatsomespecies canbecharacterisedby their
highor low accumulationoftheelement.Alsointhisarea alarge numberof flowers
from severaldifferentspecies have beenparasitised by a gall forming insect. Ihave
not attempted to identify theinsect concernedor whetherdifferentspecies ofinsect
parasitise differentspeciesofAlyxia butthis couldprove interesting forfuture study.
Ihaveincludedmanyofthespecies recognised byBoiteau(1981)inA.tisserantii.
Thesespecies arefromhisseries Globuliferae, alongwithA. celastrineafromseries
Reinwardtianae.Itwould appearthatBoiteaurecognised manyofthesetaxa based
on characterssuch as peduncle length, leafshape andsize, fruitshape andsize, and
thepubescence or lack ofit. However, all ofthesecharacters are highly plastic and
variable.Thereis alarge variationinleafshape withthenarrower-leavedformsbeing
includedinspecies such asA. tisserantii,A.pseudoserpentina, A.spathulata, A.micro-
carpa, A. microbuxus, A. breviflora andA. discolorandthelarger-leaved variations
generally being includedinA. celastrinea.The nameA. obovata Seem,from New
Caledoniapredates alloftheseexceptforA. tisserantiiandwas overlookedduetothe
fact that itwaspublished inaworkotherwiseaboutFijian plants. Itencompassesthe
sortof variationwhichhas generally beenreferredtoA. microbuxus.Many ofthere-
ported differencesinfruitshape andsize(Boiteau, 1981)areduetothefactthatmost
specimens werecollectedbeforethefruitwas ripe. Whenripe thearticlesarecovered
in athin flesh and are usually black or almost black. Immaturefruits are usually
slightly ribbed, greenish andmore globosethan themature ones, although not exclu-
sively.
There hasbeen some confusionover the correct authority for several names of
NewCaledonianspecies ofAlyxia. Guillaumin(1911)published alistofthespecies,
includingthespecies whichhadoriginallybeendescribedinGynopogon. Unfortunately
hedidnotactuallymakethenew combinations.InsteadGuillauminhimselfandBoiteau
(1981) ascribedthecorrect publication ofthesecombinationstoGuillaumin(1941).
However,Daniker(1933) discussedseveralofthesespecies and,whilstascribing the
names toGuillaumin'spaperfrom1911,actually madethesecombinationsinhispaper.
Fortunately theslightly earlierpublicationofthesenames thanwas previously appre-
ciateddoesnotchange thenamestobeusedforthespecies involved,onlythecitation
oftheauthority.Boiteau(1981) typified severalspecies ofAlyxiafromNewCaledonia
withtheindicationholo-,P!InmostcasestheseParisspecimens canatbestbedescribed
as lectotypes, oftenas theoriginal description containedseveralsyntypes. However,
thisindicationis takenas alectotypificationofthePariselementandthetypification
is given hereas 'lectotype' where appropriate. Underarticle9.15 oftheSaintLouis
Code (Greuteretal., 2000) whereBoiteauindicatedParisbut didnot indicatewhich
D. J.Middleton: Revision ofAlyxia. Part 2: Pacific Islands and Australia 3
specimen oftwoor more duplicates wasintendedIhave narroweditdownasastep2
lectotypificationofone specimen inParis.
TherehavebeenseveralpapersontheAlyxiaspecies ofAustralia,including many
Floraaccounts. Earlierworks includethoseby Brown(1810),whodescribedthethree
mostwidespread species A.spicata, A.ruscifoliaand.A.buxifolia,andBentham(1869),
whodidthefirstFloraaccount ofthegenus.Morerecent accounts includeStanley &
Ross(1986) forsouth-easternQueensland, Dashorst&Jessop(1990) fortheAdelaide
region, Harden& Williams(1992) forNewSouthWales,andWheeler(1992) forthe
Kimberley region. The mostrecent paperson the Australianspecies are by Forster
(1992)andCranfield(1995)withaFloraaccount forallofAustraliainForster(1996).
Thespeciesrecognised inthisrevisiondiffersomewhatfromtheFloraofAustralia
account. I was unableto findany significant differencesbetweenA. ruscifolia and
A.sharpeiandinsteadfoundthatthecharacterofinrolledleafwascontinuously variable
fromoneextreme totheother,as is theindumentumontheleaf.On theotherhandthe
two subspecies withinA. ruscifolia thatForster describedin 1992andmaintained in
1996,subspp. tropica and major,appeartometobe differentenough towarrant spe-
cificstatusand are certainly as differentfromA. ruscifolia as<A. gynopogon.I have
raisedbothtospecificlevelasA.tropica (P.I.Forst.)D.J. MiddletonandA.oblongata
DominandnotedthatitisA.oblongata whichalsooccurs intheLesserSundaIslands
andNewGuineaandnotA. tropica as was suggested by Forster(1992) whentreated
asasubspecies. Alyxia tropica is highly distinctivewithits dense pubescence onthe
outsideofthecorollatube.Theaffinitiesofthenewspecies A. evansiiaresomewhat
obscure withinthisA.ruscifolia group.Itoccurs intheNorthernTerritorieswithinthe
rangeofA. tropica buthas aglabrous corollatubeandanovary whichhashairsonly
inatuftbetweenthecarpels. ItappearsclosertoA.oblongata but differsintheovary
andtheshorterobtusesepals. Unfortunatelyitisonly knownfromthetypecollection.
Thesefourspecies areclosetoA.orophila,A.ilicifolia,A. gynopogonandA. magnifolia
andany moredetailedstudy ofthegroupshouldincludeall thesespecies.
Forster (1992, 1996)synonymised A. obtusifolia underA. spicata andincluded
A. thozetiiinalistofexcludednames as asynonym ofA. stellata, noting that itwas
probably introducedby abirdandwasnot native.Bothspecies areactually synonyms
ofA.stellata,aspecies knownfromthroughoutthePacificbutnotpreviously acknowl-
edged to be nativeto Australia.Several other collectionsofthis species have also
beenfoundfromAustralia, mostlyinPortCurtisDistrict.ItdiffersfromA. spicatain
thesimplefew-floweredinflorescenceandinalmostalways havingclearly pedicellate
flowers.InAustraliathisspecies alsoalways hasa glabrous inflorescence compared
tothepubescent inflorescenceofA. spicata.
TheAustralianoffshoreislandsofLordHoweIslandandNorfolkIslandhave two
andonespecies respectively. Thetwo species onLordHoweIslandare A. ruscifolia,
alsofoundontheAustralianmainland,andA.squamulosa, aspecies whose affinities
wouldappeartobemorewithspecies inNewCaledonia.Theonespecies on Norfolk
Island,A. gynopogon,isclearly closelyrelatedtoA. ruscifolia, differingfromitmost
noticeablyinthelack ofalong sharp mucronate leafapexalthough some specimens
doshow arudimentary one.
SpeciesofAlyxia extendout intothePacificasfarasHendersonIslandintheEast
andHawaiiintheNorth.Itis very noticeable,however,thatthe diversity inspecies is
4 BLUMEA Vol. 47, No. 1,2002
very much lowerthan in Malesia,New CaledoniaandAustraliaandin this account
only seven species arerecognised inthisarea(notincluding theAustralianterritories
ofNorfolkIslandandLordHoweIsland). Oneoftheseis anewspecies, A.solomon-
ensis, fortheSolomonIslands. There have been a numberofworks ontheAlyxia
species ofMicronesiaand eastern andnorthernPolynesia (including Hawaii) butas
theseconcernonlythe.A.stellatacomplex theyarediscussedfurtherunderthatspecies.
Themost detailedwork inthe western Pacifichasbeenfor Fiji, beginning withthe
workofSeemann(1866) andmost thoroughly withSmith(1988). Smithrecognised
five species and anumberofvarieties, several ofwhichhavenow been reduced to
synonymy. The Alyxia species of Samoahave been extensively written about by
Rechinger (1910),Setchell(1924) andChristophersen (1935). InVanuatuGuillaumin
(1932) describedthenewspecies A. efatensis andlisted anunknownAlyxia which I
have includedin A.podocarpa, the only species from mainlandNew Caledoniato
alsobefoundelsewhere.Alyxia stellataandA. bracteolosahavealsobeen foundin
Vanuatu.In Tonga Yuncker (1959) enumeratedjust three species ofwhichone has
beensynonymised. Ifone doesnot includeNew Caledoniathereis a veryrapid drop
offinnumbersofspecies going eastwards:inNewGuineathere are 30species, inthe
neighbouring SolomonIslandstherearejustfourspecies, inVanuatufourspecies, in
Fiji threespecies, inTonga andSamoatwo species andintherest ofthePacific only
onespecies, A. stellata.
InPart 1ofthisrevisionproblematic species complexes werehighlightedforfuture
work withanadmissionthatastudy ofthissort cannothope to solvealltheproblems
insuchalarge genus.ThesameistrueinPart2withparticularlyproblematiccomplexes
around A.stellata, A. tisserantiiand,A. ruscifolia. Further discussions can befound
underthose species.
Collections
Thereare manyareaswhereinsufficientcollectingmayhavegivenfalseimpressions
of the distributionsofspecies andmayeven mask intermediatesbetween species. I
haveseennoAlyxiacollectionsfromKiribati, theMarshallIslands, thePhoenixIslands
ortheLineIslandsalthoughtherearecollectionsfromtheislandgroupsinalldirections
fromthese.Thereareafewfruiting collectionsfromVanuatuofmaterialIhaveidenti-
fied as A.podocarpa, aspecies otherwisefoundonly inNew Caledonia.Itwouldbe
interesting to collectflowering materialof this species toconfirm, orotherwise,that
itisconspecific withA.podocarpa fromNewCaledonia.Alsotherelationshipbetween
A. bracteolosa andA. efatensis mightbe clarifiedby furthercollectinginVanuatu.
SYSTEMATICTREATMENT
ALYXIA
Alyxia R.Br.(1810)469,nom. cons.; Roem. & Schult. (1819)439;Spreng. (1817)494;G.Don
(1838)96;A.DC.(1844)345;Benth.&Hook.f.(1876)697;Pichon(1948) 164.—Alyxia sect.
GynopogonPichon (1948) 165,nom.illeg.[itincluded the typespecies ofAlyxia],—Alyxia
ser.Alyxia Markgr.—Typespecies: Alyxiaspicata R.Br.
Pulassarium [Rumph. (1747)430, nom.inval.];Kuntze (1891)416,nom.illeg
GynopogonJ.R.Forst.& G.Forst.(1775)35,nom. rejic.; K.Schum.(1895) 151.—Typespecies:
Gynopogonstellata J R.Forst.& G.Forst.
D.J. Middleton: Revision ofAlyxia. Part 2:Pacific Islandsand Australia 5
AlexiaWight(1848)t. 1293,orth.var.
Paralstonia Baill. (1888)750. —Typespecies:Paralstonia clusiacea Baill
Discalyxia Markgr. (1926)282.—Alyxia ser.Discalyxia (Markgr.) Markgr. (1977)410;Boiteau
(1981)98.—Typespecies: Discalyxiaridleyana(Wernham)Markgr.
Alyxia ser.Reinwardtianae Markgr. (1977)380;Boiteau (1981)100.—Alyxiaser.Reinwardtianae
subser.Reinwardtianae Markgr. (1977)386. —Typespecies: Alyxiareinwardtii Blume.
Alyxiaser.Reinwardtianae subser. Clusiaceae Markgr.(1977)380.—Typespecies:Alyxiaclusiacea
(Baill.)Pichon.
Alyxia ser.Reinwardtianae subser. Pilosae Markgr. (1977) 382. — Typespecies: Alyxiapilosa
Miq.
Alyxia ser. DefoliataeMarkgr. (1977)398. —Typespecies: Alyxia defoliataMarkgr.
Alyxia ser.Floribundae Markgr. (1977)394. —Typespecies: Alyxia maluensis Markgr.
Alyxiaser. GlobuliferaeMarkgr.(1977)392;Boiteau(1981)98.—Typespecies:Alyxiaconcatenata
(Blanco)Merr.
Alyxia ser. Laurinae Markgr. (1977)391.—Typespecies: Alyxia laurina Gaudich.
Alyxiaser.LaxifloraeMarkgr. (1977)406.—Typespecies:Alyxia laxiflora Merr.
Alyxiaser.MegalocarpaeMarkgr. (1977)393. —Typespecies: AlyxiascortechiniiKing&Gamble.
Alyxia ser.MicrophyllaeMarkgr. (1977)404. —Typespecies: Alyxia microphyllaMarkgr.
Alyxia ser.RuscifoliaeMarkgr. (1977)412. —Typespecies: Alyxia ruscifoliaR.Br.
Alyxia ser.SubalpinaeMarkgr. (1977)402. —Typespecies: Alyxia subalpinaMarkgr.
Alyxia ser.Baillonianae Boiteau in Boiteau& L.Allorge(1979)444;Boiteau (1981)100.—Type
species: Alyxia baillonii Guillaumin
Alyxia ser.Suaves Boiteau in Boiteau & L.Allorge (1979) 444; Boiteau (1981) 100. —Type
species: Alyxiasuavis (Baill.) Schltr.
Alyxia ser.CylindrocarpaeBoiteau in Boiteau & L.Allorge(1979)445;Boiteau (1981) 101.—
Typespecies:Alyxia cylindrocarpaGuillaumin.
Alyxia ser.Bracteolosae A.C.Sm.(1988)56.—Typespecies:Alyxia bracteolosa A.Gray.
Alyxia sect.MonospermaeTsiang&P.T.Li (1990)27—Type species:Alyxia balansae Pit.
Climbers, scramblers or shrubs.Branchlets mostly strongly or weakly angled when
young,becoming mostlyterete withage;lenticellateornot;pubescent ornot.Leaves
opposite orinwhorlsof3-7, moreor less equal insize, entire; colleterspresent inthe
axils; secondary venationusually only distinguishable withdifficultyfromthetertiary
venationornotdistinguishable atall,tertiaryvenationgenerallyparallel tothesecond-
ary venationor somewhatreticulate,oftenwith an intramarginal vein.Inflorescence
ofsolitaryflowers, orsimplepleiochasia, orcompound pleiochasia andthensometimes
forminglarge terminalpanicles; peduncle delicateorrobust,rarely moreorlessabsent,
pubescent orglabrous;bracts usually small, sometimesratherleafy, persistent orcadu-
cous;bracteolesabsentorwith one, two or severalbracteolesonthepedicel. Flowers
5-merous(oneMalesianspecies sometimes4-merous). Sepals erect,rarely reflexed,
rarely somewhat fleshy andrarely ofwidely varying sizes, ovate to linear.Corolla
actinomorphic; lobessinistrorsely contortedinbud;tubecylindric, somewhatinflated
aroundstamens;lobeserect,spreading or reflexed;outsideandinsideglabrousorpu-
bescent. Stamens inserted mostly in theupperhalfofthecorolla tube, more rarely
aroundorjustbeneaththemiddle,notexsertedfromcorollathroat;filamentsstraight,
short and thin; anthers ovate, fertilefor most oflength; freefrom pistil head. Disc
absent.Ovaryoftwo separatecarpelsunitedintoacommonstyle; glabrous, withtufts
ofhairbetweenthetwo carpels, pubescent inaring aroundthebase oftheovary, or
pubescent allover;style glabrous; pistilheadsmall, pubescent. Ovules several.Fruit
apairof drupes from eachflowerconsisting ofone or more articles withone seed,
6 BLUMEA Vol. 47, No. 1,2002
whenmorethanone thenforming amoniliformchain;articles globose, ellipsoidor
elongated, symmetricalorsomewhatcurvedespecially intheelongated articles;endo-
carpmostlythinandpaperybutoccasionally muchthickerandbecoming tough, meso-
carpfleshy, oftenvery thinly so, exocarp thinand coloured.Seedssimple; ruminate
or withlongitudinal ridges. Embryo with flatto strongly undulatecotyledons.
106species foundfromNorth-EastIndiathrough SouthernChinatoTaiwanandsouth-
wardsthroughSouth-EastAsiatoAustraliaandeastwardsthroughtheSolomonIslands
out intothePacificwest asfarasHendersonIslandandnorthtoHawaii.Part 1ofthis
revisionsuggestedtherewere 108speciesbutsubsequent workonthespecies included
in Part 2 has revised that figure slightly downwards. In Australiaand its offshore
islandsthereare 14species, inNewCaledonia21 andintheremaining PacificIslands
7 species.
KEY TO THE SPECIESIN AUSTRALIA (INCLUDINGTHE AUSTRALIAN
TERRITORIES OFNORFOLK ISLAND AND LORD HOWE ISLAND)
la. Leafapexclearly mucronate;shrubs 2
b.Leafapexnot clearly mucronate;shrubs or climbers 13
2a. Leafmargin toothed 14.A. ilicifolia
b. Leafmargin not toothed 3
3a. Outsideofcorolla densely pubescent. — NorthernTerritories ... 38.A.tropica
b.Outsideofcorolla glabrous or withjustafew hairs aroundthemiddleor top of
tube.— Widespread 4
4a. Leafmargins strongly inrolled 5
b. Leafmargins only weakly inrolledor not atall 7
5a. Inflorescencesaxillary, robust, flowerssolitary; plantsofhigh altitude
24.A.orophila
b. Inflorescencesterminalor appearing terminal, mostly delicate, flowers solitary
or incondensedcymes;plantsoflowlands or submontane 6
6a. Inflorescenceaxessparselyto densely puberulent;sepals densely puberulent, apex
acuminate;corolla tube>4times aslong as sepals. — WesternAustralia ....
35.A. tetanifolia
b. Inflorescenceaxes glabrous; sepals glabrous tosparsely puberulent, apexobtuse
to acute; corolla tube < 4 times as long as sepals. — Queensland, New South
Wales 29.A.ruscifolia
7a. Mucronateleafapex shortandnot sharply pointed andoftenabruptly apiculate/
mucronatefrom anobtuseoreven retuse apex. —WesternAustralia,SouthAus-
tralia,Victoria, Tasmania, NorfolkIsland 8
b. Mucronateleafapex sharply pointed from an acute or acuminateleafapex. —
NewSouth Wales,Queensland, NorthernTerritories; LordHowe Island 9
....
8a. Venationfrequentlyobscure;flowerspedicellate; antherapex< 1mmfromcorolla
throat;ovary atleastpartly puberulent. — MainlandAustraliaandTasmania .
3.A.buxifolia
b. Venationweakly prominent above; flowerssessile or subsessile; antherapex> 1
mmfromcorollathroat; ovary glabrous. — NorfolkIsland . . 12.A. gynopogon
D.J.Middleton: Revision ofAlyxia. Part 2: Pacific Islands and Australia 7
9a.Flowers clearly pedicellate, generally in a branchedcyme; largest leaves 2.7-
14.7 cm long 10
b.Flowers sessile or subsessile, generally solitary or in very condensed cymes;
largest leaves 1.4-7.8cm long 11
10a.Stamens inserted at0.31-0.54of corollatubelength; sepals 0.9-2.2times as
long as wide; secondary veins at65-75°from midrib 14.A.ilicifolia
b.Stamensinsertedat0.66-0.72ofcorollatubelength; sepals 2-2.6timesas long
as wide; secondary veins at20-65° frommidrib 19.A. magnifolia
11a. Antherapex>2 mmfromcorollamouth;stamens insertedat0.37-0.58ofcorolla
tube length; corollatube5.8-9.5 mmlong 12
b.Antherapex<2 mmfromcorollamouth;stamens insertedat0.51-0.73ofcorolla
tube length; corollatube4.3-7.2mmlong 29.A. ruscifolia
12a. Sepals 1.8-2.6mmlong, 1.4-2.6timesaslong aswide,apex acute toacuminate;
ovary pubescent allover. —Queensland 22.A.oblongata
b.Sepals c. 1.6mmlong, 1.1 timesas long as wide,apex obtuse;ovary pubescent
only in atuftbetweenthecarpels.— Northern Territories 9.A.evansii
13a.Rigid erectshrubs; leafbladescoriaceous 14
b.Climbersor scandentshrubs; leafblades coriaceoustopapery 15
14a. Venationfrequentlyobscure; flowerspedicellate; antherapex<1mmfromcorolla
throat; ovary atleastpartly puberulent. — MainlandAustraliaandTasmania .
3.A.buxifolia
b.Venationweaklyprominentabove;flowerssessileor subsessile; antherapex> 1
mm fromcorollathroat;ovary glabrous. — NorfolkIsland . 12.A. gynopogon
15a.Flowerpedicels mostlywithnumerousbracteoles, somepedicels withonly 2but
thenbracts large andleafy or lanceolate.—LordHoweIsland
33.A.squamulosa
b.Flowerpedicels mostlywithoutbracteoles,bracts generallysmallandinconspic-
uous.— MainlandAustralia 16
16a. Inflorescencemostlyasimplepleiochasium, 2-4(-6)-flowered,glabrous (inAus-
tralia);pedicels to 23mmlong 34.A.stellata
b.Inflorescencegenerally morecomplex, 8-15-flowered,sparsely to denselypubes-
cent; pedicels to 1.1 mmlong 17
17a.Leafapex mostlysharp acuminate,rarely toobtuse;secondary venationdistinct
above;fruitarticles 12-15by 11.6-13.4mm 11.A. grandis
b.Leafapexroundedto bluntacuminate; secondary venationmostly only slightly
visibleorobscureabove(in Australia); fruitarticles 7.6-13 by 6.9-11 mm ..
32.A.spicata
KEY TOTHE SPECIESIN THEISLANDS OF THEPACIFIC OCEAN
(EXCEPTNEWCALEDONIA AND THE LOYALTY ISLANDS)
la. Leaves sessile. —Fiji 8.A. erythrosperma
b.Leaves petiolate. — Widespread 2
2a. Branchlets sparsely to densely puberulent 3
b. Branchletsglabrous 5
8 BLUMEA Vol. 47, No. 1,2002
3a. Leaves darkgreenanddullabove,venationobscurebeneath,usuallyalsoobscure
above;fruitwith2-8 articles, denselypubescent. —Vanuatu 26.A. podocarpa
b.Leaves variablebut generally shiny above,venationobscure to distinctbeneath,
notusually obscure onbothsurfaces; fruitwith1-3articles, glabroustoonly very
sparsely pubescent. —Widespread 4
4a. Inflorescencepuberulent, 5- or6-flowered; corollatube5-7mmlong
31.A.solomonensis
b.Inflorescenceusuallyglabrous, ifpuberulent thenfewer-floweredandwithshorter
corolla tube 34.A. stellata
5a. Bracteolespresent onall pedicels 6
b.Bracteolesabsentorpresenton some,but not onall, pedicels ofaninflorescence
7
6a. Bracteolesusually morethantwo perpedicel, sometimesreducedtooneonjusta
few flowers; corolla tube3.5-12.4mmlong 2.A.bracteolosa
b.Bracteoles one perpedicel immediately beneath thecalyx; corolla tubec. 12.6
mmlong 7.A.efatensis
7a. Inflorescencerobust; all pedicels <2 mmlong; corolla tube8.1-11.2mm long;
leaves obovateto elliptic. — Bougainville andtheSolomonIslands
16.A. kwalotabaa
b.Inflorescencedelicate; atleast somepedicels in an inflorescence> 2 mm long;
corollatube 1.8-10.1mmlong; leaves only rarely obovate.—Widespread . . .
34.A.stellata
KEY TO THESPECIES INNEWCALEDONIA AND THELOYALTYISLANDS¹
la. Branchletsstrongly angled; leavesinwhorlsof4or5; largestleaves 8.1-16.4cm
long; inflorescencesbranchedseveraltimeswith>20flowers 17.A.leucogyne
b. Branchletsteretetostrongly angled; leavesopposite or inwhorlsof3(—5);largest
leaves0.9-13cm long;inflorescencesvery simple withfewerthan 10flowersor
large andlaxbutthenwithopposite leaves andwith< 15flowers 2
2a. Leaves distinctlymucronate 3
b. Leaves not distinctlymucronate 7
3a. Branchlets sparsely to densely pubescent; leavespubescent or glabrous beneath;
sparsely pubescent aroundtheoutside ofthecorollatube . . 21.A.mucronata
b. Branchletsand leavesglabrous; corollatubeglabrous outside 4
4a. Inflorescencesvery lax, 4.8-14cm long;pedicels 2.8-22mmlong; fruitarticles
somewhatsickle-shaped, 2.4-5.7cm long 20.A.margaretae
b. Inflorescencesvariable, 1.2-3.8cm long;pedicels0.8-8.5mmlong; fruitarticles
ellipsoid or fusiform,0.7-3.7cm long 5
5a. Leafbladepaperytosubcoriaceous; inflorescence1-3-flowered;corollatube7.7-
10.2mmlong,5.4-8.5timesas long as sepals; fruitarticlescylindrical
6.A.cylindrocarpa
b. Leafbladecoriaceous;inflorescence3-11-flowered;corollatube3.2-7mmlong,
2.3-4.5 timesas long as sepals; fruitarticlesellipsoid 6
1) Thekey usesthe characteristicsfor the widespreadAlyxia stellata only asitis found in New
Caledonia and theLoyaltyIslands.
D.J. Middleton: Revision ofAlyxia. Part 2: Pacific Islands and Australia 9
6a.Inflorescence7-11-flowered; peduncle not strongly flattened;corolla tube 3.2-
4.2mmlong,2.3-2.5timesas long as sepals 28.A. rubricaulis
b. Inflorescence 3-flowered; peduncle strongly flattened; corollatube5.8-7 mm
long, 4.1-4.5timesas long as sepals 27.A.poyaensis
7a.Sepals oblong, leafy, apex roundedto obtuse; inflorescencesconcentratednear
branch ends; corolla tube densely puberulent outside; corolla lobes densely
puberulent outside; fruitarticles globular, black,very fleshy . 30.A. sarasinii
b.Sepals ovateor lanceolate,leafyornot, ifleafythenapex acute toacuminate;in-
florescenceposition variable; corollatube glabrous, puberulent around topof
tube or only sparsely puberulent outside, corolla lobes glabrous outside; fruit
articles various 8
8a. Inflorescence7-11-flowered,with distinctinternodes;branchletsred
28.A. rubricaulis
b.Inflorescencel-6(-7)-flowered,if7-floweredthenwithoutredbranchlets;mostly
withoutdistinctinternodes 9
9a. Flowers terminalandsolitary; leavessmall, 0.5-2.2 by 0.15-0.45cm
4.A. caletioides
b.Flowersininflorescencesof2ormore,ifsolitarythennotterminal;leavesvariable
insizeandshape 10
10a. Sepals3.5-4.2mmlong; bractslanceolateorleafy, 4-11mmlong; leavesthickly
coriaceous, margins mostly strongly inrolled 5.A. clusiophylla
b.Sepals 0.6-3mmlong; bracts mostlydeltoidorovate,rarely lanceolateorleafy,
0.5-2.5 mmlong; leaves variable 11
11a.Leaves darkgreenanddullabove, venationobscurebeneath, usuallyalsoobscure
above;fruitwith 2-8articles, densely pubescent 26.A.podocarpa
b.Leaves variablebut generally shiny above,venationobscuretodistinctbeneath,
not usually obscure onboth surfaces; fruitwith 1-6 articles, glabrous to only
very sparselypubescent 12
12a.Leaf surfaces strongly discolorous, mostly with obscure venationbeneathand
often glaucous, corollatube5.5-9 mmlong; fruitarticles generally fusiform,
6.3-23.5 mmlong 1.A. baillonii
b.Leafsurfacesgenerally onlyslightly discolorous ifatall, venationvariable,only
rarely glaucous butifso then corollatube less than5.5mm long; fruitarticles
variable 13
13a. Stamens insertedat4.3-6.4mm fromcorollabase 14
b.Stamensinsertedat 1-3.9 mmfromcorollabase 16
14a. Petiole0.1-0.2cmlong;leavesreaching 1.8-3.9cmlong; inflorescencedelicate,
peduncle c. 0.3 mmwide; sepals 0.7-1.1 mm long, c. 1 timesas long as wide;
corolla lobes 1.4-1.6by 1.3mmwide,tube3.7-5 timesas long as lobes
....
23.A. oppositifolia
b. Petiole0.3-1 cm long; leaves reaching 5.3-10 cm long; inflorescence robust,
peduncle 1.2-1.5mmwide; sepals 1.6-2.2mm long, 1.5-2.2times as long as
wide;corollalobes2.3-4.3 by2.3-3.1 mmwide, tube1.6-2.8timesas long as
lobes 15
15a. Secondary veins on leaves distinct beneath; inflorescence sparsely to densely
puberulent allover; sepals densely puberulent; corolla lobes 2.3-3.5mmlong,
10 BLUMEA Vol. 47, No. 1, 2002
tube2.1-2.8 timesas long as lobes; ovary densely pubescent allover
25.A. oubatchensis
b.Secondary veins onleavesonly weakly visiblebeneath; inflorescenceglabrous;
sepals glabrous; corollalobes c. 4.3 mmlong, corollatube 1.6times as long as
lobes; ovary pubescent in tuftbetweencarpels 39.A. veillonii
16a.Branchlets sparsely to densely puberulent 17
b. Branchletsglabrous 19
17a.All inflorescencesterminalor pseudoterminal 15.A. kaalaensis
b.Mostinflorescencesonaplantclearly axillary, occasionally alsowithapseudo-
terminalinflorescence 18
18a.Leafblade generally obovate, thickly coriaceous, often with obscure venation,
often glaucous beneath; corolla tube3.6-5.1mm long; fruitarticles8-16 mm
long 10.A. glaucophylla
b.Leafbladevariablebut mostly notthicklycoriaceousandwithvenationgenerally
visible, not glaucous beneath; corolla tube 1.6—3.8(—4)mm long; fruit articles
3.5-14.5mmlong 36. A.tisserantii
19a. Bracteolespresent, corollabudheadellipsoid 18.A.loesneriana
b.Bracteolesabsentoronlyonpedicel ofterminalflower, corollabudheadglobular
or ovate 20
20a.Fruit articles 14-20by 7-9.5 mm, generally about2 times as long as wide or
more;ovary densely pubescent allover;inflorescence4-6-flowered,lax ...
13.A.hurlimannii
b.Fruitarticles 3.5-14.5by 3.4-9 mm, mostly less than2 timesas long as wide;
ovary pubescence variable; inflorescence 1-5-flowered,variable 21
21a. Peduncle< halflength ofleafblade; fruitarticles ellipsoid, ifglobose > 6 mm
long 22
b.Peduncleabouthalflength ofleafblade; fruit articles globose, 5.2-6mmlong
37.A. torqueata
22a.Corollatube<4mmlong. —NewCaledonia 36.A.tisserantii
b.Corollatube>4mmlong. —Loyalty Islands 34.A.stellata
1.Alyxia bailioniiGuillaumin— Map 1
Alyxiabaillonii Guillaumin (1941)364;Boiteau(1981) 144.—Gynopogonlaurinus Baill. (1889a)
781. —Alyxia laurina (Baill.)Guillaumin [(1911) 194,combination notmade] exDaniker,
(1933)381,nom.illeg.;Guillaumin (1948)292. —Type:B.Balansa 1403 (lectoP, designated
byBoiteau, 1981;iso P)from NewCaledonia,Bourail,Fene
Alyxiacf.leucogyneGuillaumin (1957)79.—BasedonGuillaumin &Baumann-Bodenheim 12582,
12636 and 15753.
Climbers,6-10mhigh.Branchletsweaklyangled, sparsely lenticellateornot,glabrous.
Leaves opposite orinwhorlsof3; petiole0.3-1.4cmlong, glabrous; bladecoriaceous,
elliptic,obovate orspathulate, apexretusetoshortly acuminate,not mucronate,base
acute to cuneate,margin weakly inrolledorflat, weakly undulateornot,pale greenor
glaucous beneath, 2.6-10by 1.1-4.7cm, 1.2-2.9timesaslong aswide;midribsunken
above,intramarginal nerveabsent,secondary veins 16-33pairs,65-75°frommidrib,
weakly prominenttoindistinctabove, obscureto weakly prominentbeneath,tertiary
venationweaklyprominenttoobscureabove; glabrousbeneathandabove,notpunctate