Notes on the Genus Pandanus (Pandanaceae) in Western Australia
Noteson the genusPandanus (Pandanaceae) in WesternAustralia
By Benjamin C. Stone{'
Abstract Tbe Ee[lusPatldanus in Westem Australia is known so far to occur only in the northern sector,in the Kimberley District, south to about 18. lat. Only one subgenus,Subg. Parlarrr, is represented,but there are two distinct Sectionj, and possiblylhfee. SecL. Pd dant6 lpo..ibly),Sect. AutroA(ura thereinraised to sectionalrank from thal of subsection),and Sect. Scnikerrra. The classi- fication of species_,especially in Sect. Au:troke ta, is not as )et sarisfaqory. The occurrenceol P. spiralis is docuntented,and a review of the soeciesnrosL similarro il included.resulring in comene\'| synonlnls and neu combrnationq at inliaspecificrank. Two new varieties(var. f{ammeus, yar. multimammillatu$a) are p_roposed.The variability of this speciesis discussedin relation to prc_ viously kno\\n and ne\'"lyobtained specimensfrom both WesternAustraliaand the Northem Territory. Three other speciesof subg.Pandanus, p. darwinensis, P. oblatut and,P. semiatmatus,are also reported. Finally a new suite ol coll lectiors ol P. aquaticus (syn. P. kimberleyanus) is reported.
Introiluction
- Knowledge of Pandanus in Australia goes back a long way,-two to the ex- ploratory activity of Robert Brown. Brown described only species, "Prodromus P. spiralis and P. pedunculatas, in his Florae Novae Hollandiae et Insulaevan Diemen" publishedin 1810. Brown's collectionswere studied r€centlyby H. St. John, who publishedan account of these(St. John, 1968). According to St. John, Brown's collection no. 5799 was a mixture,.and he selectedas lectotype a portion ofthis gathering to typify P. spiralis,.the material (one.fruit phalange only) is illustrated in his fig. 26j. flso assignedto the \pecre\blll not consideredto be pan ol rhelectorype \\a\ a slaminatespecimen. This rs correct, as all pandans ire dioecious. fwo phalangeswere ixcluded from the three which rnade up the type materials ofP. spiralis, and served as type for _(holotype) St. John's new-fhe species Pandanus broinii; these phalanges are illustrat€d ,l _h_olig 270. lectotype of P. ;pirulis was obtained by Brown in the Gulf of Carpentaria on Allen Island, (callei "Island Wellesley-from Group by Brown C"). The staminate inflorescence came t-he same locality. The phalangesdiscrininated as P. brownii have an unknown pro- yenence (except that they are assuredly from Brown's Australian travels). It cannot be_assumed that they came from Allen Island, nor can it be proved that they did not. . Brown's second species v,lasP. pedunculat J,.this has good locality data, showing_that_.it 1as collectedat tha north end of Great Sandy Istand (now Fraser Island) off the E. coast of Queensland. The lectotypi, also in the British Muserrm, was collected on 31 July 1802,and Jsgiven the-number 5299A. This species,which in the opinion of Domin was a variely of pandonLlstectonLs, and in_this writer's opinion is certainlypart of the p. tectoriuscomplex, has never been found in Western Australia; it is common however along the Queenslandcoast, especiallyon the oflshore islands. It is known curiently under a large number of different names, but that subject is dealt with elsewhere. Further contributions to the knowledge of Australian Pandanaceae, especially Pandanus, have come from Martelli and more recently from St. John and Stone. Some of these contributr'onshave dealt with collections
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Genus PANDANUS
SubgenusPANDANUS (sensuStone, 1974b).
Sect.Austrokeura (B. C. Stone)B. C. Stone,.r/al.ror,. (Pandanus Sect. Pandanu,cSubsect. luslro,(srr.4 Stone, Bot. Jahrb. Syst. 94 : 517 (1914)Basiorrym). (a) SeriesAustrokeura. Type'. Pa dqnussolms-laubachii. Phalanges laterally conspicuously sulcate, wilhin the cephalium the phalangesinterlocked by the groove-and-ridgesystem formed by these sulci. Phalangesvarious, from rotund to obovoid or suboblong. Mostly Queensland species,but someprobably also in the Northern Territory. (b) SeriesSpirales Stone, ser. nov. Carpellisphalangiorum fere esulcatis; phalangibus rotundatis. fype'. Pantlauutrliralrs R. Br. Mainly WesternAustralia and the Northern Territory. This differenceis not hard ald fast but on the whole seemsto reflecttwo 'forms' real lrends. Using this conception a group of appears to cluster rou.ndP. spiralis. This group also showsthese additional phalange characters: (1) broad, rounded shape of the phalange,and (2) the conrparativelylarge number of carpelsper phalange. It becomesevident that the conspicuouslyridged-and-grooved phalanges characteristicof Set. Austrckeuraare particularly common among Queensland species,becoming scarceralong an east-to-westgradient; while the smoother plralanges of the Ser. Spirales are most common in Westerl Australia and the Northern Territory, but scarceto absent in Queensland,especially eastern North However, there are enough exceptionsto suggestthat Queensland."hybrid there is possibly a belt" of forms bridging the W. Queensland E. Northern Territory populations,and that the ridge-and-groovecharacter has "flowed" in a westerlydirection. The specieswhich would be assignedto thesegroups are, if one simpJy followed the classificatiorrand speciesconcept of St. John (which I do not accept) (Those marked * appear sonewhat intermediate between the two serres): 238 (/961 -euots ""9o"tatun1nqored sledr€cJo requnN (l ) : sJelJ?ftqt 3ur,to11o3eql e^loAul eqJ so^rlElel lse.rEeusI ptrE sllD.tlds 2'jo se8utlerld uI suol]?llE^ ul"u 'sllErJds'd salJadspalslar puB saPolJB^sl! '(9161'euo1g eas)sDpt,lD,t,{lo3el?J eql esn ol tlolslJePeq} JJtlJq :ex€l lBlJuJrlde eq Aeu 'esuos ("ru sleqlo tnq p:ot8o1orqrelcrrls o,1jui'sairodsqns :q o1 :ro:d es'qlJo euroS sarlalrel oleq EX"l Jlxtllploqns Jo lslsuoJ.o] peuieepsI Pell"J ir,ru:'ui.r'lo'tIPJJ\es 'qJrq'nn ',{pnis put uorssncsipe}eltlttoJ 01 se tijn*'te roJ serc'ds eq} xjldruoc r sEstlD.tds g pruSa: o1 leJeJdI ,{qA\pu" ,lroq ^\oqs ol aleq luelul reg o1 sr tr pu? . ,rl,!?./arllrrp 7 "1rtp"*relur ,tlrudrecide eqt 01 pue sap;rd5 redtd slql u1 peua''ssee,leq I ,ictq,meioql uo pessLrool5q 11t,truot1ual1e.ra,l:'toq :ornn1o.,l"'ny' 'ut \uq 1.ig serjedsriezop drlt ueqt :.roru pasodord uqof lS ',{1e.rrin1cxe 'srstq 'sJe)JeLU lsoup slql uO tlurouoxel s" slln{ uo {c?q llP'J '1ect1s'lrls'apqns er€ '(eql 'lstx3 e^ ,{j'sdereu;g le.,iudqae pue eq ol elqull serueiegrpg1- -rtyirursflSurpeecxe Illuelequ! ele ,{eql :secuecseJoUurel"urLu"ls eqt 'oslB 'oS se^?elaql ul punoJ uJJq J^cLIsJrJJds elElltt;r4lp ol luerc$ns p3zuelc€l"rl} tra1.n:nq" foo8 ou'ie; os pi,t'*ro1 ieal ped{loerets{lJ\ E Iq 'sar:rods Suolllv esn re,\e,roqsr un loJlstlv.1ca5;o e1o,1'nlql uetlerlsny aq1 'setcads op secuel{ Jo 3q ^elu srelJ"r"r{J el€utu"}s ue^e pu" uee,trleqlslxe -jrp e8erlog:lle,r s? peuruExe eq lsnu $ln{ ueq} loqlo se.rnlJnrls"ro^e'\\oH 'rrorlEreprsuoosnoues qlto,'!\u3ql sl suleuJl se Fltltqet:ll,l qJnS pol€ululll? slrn{ pozlltueJun eq lsn,u'aseql-'sielcurtqr1Ie'510'Sirrfrp ut 53e1u'lqs'1et::1cruodtrun Jo slq"llnsun el" Jo",{lrnotnqo selnl€eJ eltue,lnf lsetcedsSutlttttutrcstp loJ iro'1e':u,r.go spuil euos leql ,{esot ll JJUJnS sl;q pJlcade:aq lou 'qtget 'euo1g1slrodar snotr:ld ur qlStrrl lE qll't\ ll€ep pieu pur (Stef h19i1 'ltLeld i,""q inq tiiff siiiScis e-uii eql jo s1uu1d1uo-rogrp urorJ Ptru et'"s oql Luorj seduiliqO ro slrnrj SuoutepunoJ eq o1 uotlerrn^Jo \ptrr) '(unutale areql '{pn}S ]"r11 u,{\oqs s?q snaDPuttJJo l"r-Iel€ufJo lunolLl€ elqvleplsuoJ " Jo 'sorJeds 'lunpl^lpul ,{uBolloJ ol 3uo alqeuo E JuolBlal tlDul{llllqEU"A Jo uollou (roprsuoc ol qanous {tfLeq' t.8;ni;U; ou] xq',t1uo pelueserdersr srql pu? 'f-rr:luauBurl s,u'torg o1 linrj eq1 ,{1uo',(1Iuarsr ireql Suroq uerutceds.ed'{1 '10^e'\oH 1ar1 ,(1ilqirre,r.eq] seop uroleqr'd:st uotlsanb 1u€curu?ls3t{1 'serreds(p:qrrrsepun {11cnsn1 l:q}o o} pauBtssleq lsnru 'tlclBLu 'alques 's!lD.tdyd (ad'(]otJrl)uJurJf,ds s,u,t\otB ^llenll? lou oP lnq Jo 'uqot ',t.rel^ -ai F13uor1"q.rrq,rr suepued leqt sepnpuoJ ^[leln]"u ]S puoces pelcrJlseJ srLll.lo luJUodord aq1 erluluodrt3 Jo JlnD eq] ur pu"lsl uellv ol lir,rod. stql .,trlapua1e.o1 u tou alqnr.rune sltltttrds al leql'\l leql;JJdEd Jo oql luoun6le Jql \ulrol Inql uorldunsseeql \JullJJpunJJJq 6ulPPlq uJ\ sllBrlds 'd u! ,tllllqBlrs^
'uqof ]s sr?l'''r' 'uqof 'tS '.uqol 'd 'uLIo['lS -Irr.r 'd' slsuapi.taLaosJ lS snlolclo sllDwraql J.* 'uqo1 'd' iutlof 'JB J5 ,rnxJ,uo.r :lS ta1av! d u r'/1r'lirlt d :saPttds \eS 'JOLulJ JO rIeTJEll souDao J ,{lqeqordpul'areq s8uoleq€nd?dJo,fuJod d' ll0n r.rss,.r.1d.'oslv uqot lS 'd 'uqot 'uqol '1,5-la/Dtpr.i s!.tD\,\t.t pu" lS sls-uaDPuD.tDld ;r'uqof 73 sn)!uDul iuqog 'uLlof 'uqof -ssru g lS stlsollldDd J 1S sn1tntlltol d 1S taJtuolols d 'utfof ruqol '1S lur{of 'd 'utfoI lS sntDpund d sn|olnBuor'tr1dd 1S sntDw.tDxa luqof 'u{of 'lS slfif)ry.o 1S stsuaut$.lDp dx 1,5stwrautqns d-uqot rS d/ 'uqor 'tlqof 'uqof stiuatninoajtuu lS snLrDtuowtrral d lS snJo)utt.tl d ls 7'uqof 'lS 'rlletJ"t! 'uqof stsuatlDrtsnDJ s^niDfELtDJ !!1on d lS s??aJ"rl?r 'J 'uqof '1S 'IIIeuBhJ 'd 'JaS stsuaLuaqu,lt)d lluuDululDp :DJnalotlsnv (2) Size of phalange, particularly length "Rotundity" "complanation" (3) versus of the phalangeapex (4) Relative convexity of each component carpel apex (5) Presenceor absenceof lateral sulci (in extremecases forming a dense seriesof Iongitudinalridges and grooves) (6) Size of stigmas
. Chiefly using these characters,St. John has discrirnjnatedthree species (P. convexus,P, integer, P, thermalis), which were considered as close relitives o-| P. but which here are regarded as subordinate taxa of that species. lpiralis, ',closest St. John in his original diagnosesoften indicatesa relative', but this seemsto be basedchiefly on the position of such taxa in his unpublishedkeys to species. The:e species,and sevenothers which on morphologicalgrounis and geographical distribution are thought also 'closest to be iniplicaiiA, r.iith tle indicated relative'according to St. John, are: "Closest P. conyexusSt. John, l96l : Relative": p- spiralis "Closest P. integerSt. John, 1962: Relative". p. conrexus "Closest P. thermalis St. John, 1962: Relatiye": p. latiliuctus "Closest P. Iatifructus St. John, 1962: Relative".. p. medialinermis "Closest P. medialinermiltSt. John, 1962: Relative" ..p. somerserensr "Closest P. somersetensisSt. John, l96l: Relative,':Not indicated "Closest P. darwinensisSt. John, 1962: Relative": p. \rhitei "Closest P. arnhemensisSt. John, 1962: Relative".. p. truncatus "Closest P. truncatusSt. John, l96l: Relative"..p. brookei
From this sequenceit may be seenthat a seriesof forms may be deduced" contexus spiralis*i teget; then another series,thermalis lot ifrictus_mediql_ inermis somerseter,rlJ, then a. pair, darwinensis-v,hitei,. and then another shoft series, qrulrcmensis-truncetu,s-brookei. It is a standard feature ,,closest of the taxonomic papers of Dr. St. John to state such a relative", but it is disputablewiether it is correct to.assumethat a specieshas always-andonly one close relative. In any case the sequencesnoted are instructive. If ref_ erence-is made (prev. page) to the specieslisted under each of the newly pro- posed Series,it is seenthat the speciesofthe first_sequence(cuar,e;ra,r spiiaits_ integer) have phalanges with smooth faces an_dno ridge-and-groove system; the species of the second sequence lthermalb-latifricrus- edialinirmis somersetensis)are mixed with respect to the _ridge-and-groove character, somersetensisand thermalis lacki[g the 1ystem Vhermalis polsessesperhaps i very weak.expression of this charact€r), while latifructui and ^"dialine:rmi show the character strongly developed; darwine sis and b)hitei both show the claracter; and in the last sequence,arnhemensis a]iLd tuncatus strongly show the ridge-and-groove system, bul brookei lacks it entirely.
To these facts must be added the point that as far as can be determined, P. brookei is extremely similar ( -. closely. related ?) lo p. somerserensu,a relatiorship not. indicated by St. John. I do not consider lhat p. truncatus p. should be considered as a close relative of brookei, nor do I acceptp. somer_ Je/ezJ6 as a close relative of P. mecliqlinerml-r. (The latter in fact is most likely a synonym of P. d.rmmqnnii Warb.). Furthermore I would consider P. tr,uncatusas ,a synonym of P. whitei Maft,elli:- this taxonomic interpretation would then link the. isolated parl, clarwinensis-,whitel,with the nna dequence, where_wemay substitute wlitei for tr.uncatus,all these speciesthen linkiig into a singfesequence Jarwinensis whitei arnhemensis. 240 I
r9z 'elaq €x31 eql lle lceJs '1stxe Pol"leunus 'wnl)o'tatu .{lpuno3ord plno^\ JnJco plp rlJ[l^\ 1q8rur snqn[ PttE 'LunJDxo,tDJ lJ Jr s? e.reue8qcns ur punoJ ]€ql Sullqueser uotlnnlls_tlurouoxel B 'os Jo {llllqlssod eql ueql JI Jncco ,{euJ s]Xlurod€ l€ql elqlssod osl" sI 1l lnq 'sarceds suolols uerlurlsny Jeqlo leJe^es pde wottds d ur lslxe ol u,{\ou{ sl 'osJnoc 'lncco sulroJ ,{q uorlcnpo:de; l?uolc JO ,(EuruorlJnpojdsJ Jo lsnxes"'es?' ]?ql (serJodssnuopuDd uerle.rlsny-uou 'Jeqlo uo peseq) slstxe uolcrdsns ? .(ut ri1 -q uotleuStsep-srq1 'ubrlezrplrq.{q sJoord ,{uE Jo lercrpnf'jra rriirru- '2'uor1eu81sepJo eruesqu eq1 rii 'ra,re,tro11 elqalrns eq plno,r,r.'srszarrr,rilpx eql sdeq:a6 perrejord eq lq8rur .serceds,qcns Jo suolllsodslp Jltuouox"l loqlo uaqi'xirleer lqi.r'eeu'e-,iq,txueare ur8t:o b1 se suotlelncodseseql jt 'sercedsleci:olsrq ' 's6uaw$,tDp 'd pu? ale cle \eLq arunssB01 ,(:essaceulou sI 1l 1nB 'sllDwDLlt 'ecnpo.rd J se qJns su.uoJ'uoruod uJalsa,&eql prearol lq8Itu suollBulq -uroce: 'ieere ieqlrnj icuoq,r leuorlnqrJlsrp o^u Jo eJEJJJlulaql 1" pelcadxo eq lq8[u 1eq,trs! ll puBlsueeno,, uJJqtJoN.,^Es ol sI sluJel 'ed,{l., ..,{JotrueJ crqdt:3oa8ur qJrq,$ ttrelrq,r,rx ed,{t,,srl€.lrds,, :uolsseldxe aql ol nullnur -IXoJdde 'd suloJ ed,{l " Jo spuqfq eq ,{puj srrMraqt pre s6uaut41.itpd sE-rlcns I l?ql elelnlsod ,{eru euo uoql os 3q srql Jl JelJ€J€LlJSurlrosse ,{lutopuu se acueitrequt q8norql pa1e.rodro.ri,,oq {zut .rJlJeJEqJe^ooJ3-pu"-o8Pu eql 1"q] pur,t 'lueluuosseeJ 01 eJ" (sJelc"J€qc e8ueleqd,(es ol lalleq) sJel lcelqns 'peldeJJ" -cereqc senads1eq1 ,{lrlqzqord eql 01 uorlruSoceJspJoJe lI sl-serJes eseql ur serJeds ^lrortdrtlnu ? se Suol sB'lool clurouox"l elq"pueoxe Jo 'uorlJuqslp '?rl"rlsnv ^us eq ,{Iuo .(eur rl q8nor{tle lnJesn € sul"uel lI Jo uorner uJeq ou aq] sso.rc?flrnurluoc " sl eJerll esnEcsq sl ll iale:edesI?cldoJl leuollnqtJlstp .lejelduroi lou ar? ser:e! o,r1-eqt;1 uoqcullslp el{l uoddns ?lBP '.os crqderSoe8pue Ieuonnqrjlsrpegl esneceqsr s|t{l pu" lutql ]ou op I Lotn prlsnY Jes pue saTa.irrl5seueg uee,rloq uorlrurlstp eql lentcaJaur eI"tu uaql lou srql seoC 'e^oq? Pel"clpur s€'seues Jo 'seJuonbeso/Al 3ql )lurl ,{lu€sseceuplno,\\ os op o1 i:ted po1ut1e 'spJo^\ se stsLraul:trlopd puu stlouJa\l J ,\\eL\ ol alqtssod sI 1l leqlo uJ ''^I I Dll.Is d SuIqc€oldds '(I 's38u€l"qd osl" snql'earrii t 6) a8u"l€qd r3d slodr"r Jo requno elsrepottlE pue punlo: 'iertj eFutltqd qloorus- raqle-r esl\\l3qlo 1nq utels{s :,r.oo.r3 'stlDw.talJ ipue-e8prr pidolenJp ,{11ei,,nr srq yall^\ ;r ,{q ptre :sllo.tlds d SLtr '!3utier1d -ilcroid'dr red sled.rer riqilinu (e t I I ) q8lq ,{pIeJ " qlrrd se8ueleqd 'Ltlels,(sJo 'l/t,a punloJ 'e8re[ saq lnq e^ooJ3-puE-e8pu SuoJls " scq qcrq,^^ -ut;tttop'4 lq peuol?eJr{l,(1ged sr set:e5or\d} eq}Jo sseue^IlJullslPoqJ €Jne{ 'solerlds -o.llsnv ros ()1pLIoJas eql Jes o1 spuodseJJoseseql Jo lsJUeqJ 'snltnL!1o1 'J-stsuautaquto J-nuLtoLutaDp d-lal!4$ 4-vsuaut.t rttp 4 (7) pu" :stlDut.Iaql d DEau! J stlD.tlds'd snxatuoJ J lI) :seouenbaso,ttl {ttlo ars A\ou J^\ uoll?nlls Jtlt 3ur,$3llau 's fipn l f 1lDl-I uuoutLuDp s!suaMaqu.tD 'ecuonbes Dltllt -stsltautalrp eq ,'\\ou plnoA\ I{3II{,\\ lsBl el{l rl}l/$ eEeIulJ 'd € urJoJ ,(€ur e,r eJueq .'t/ttlarratlultl ol J"ltturs sI tluuDrytLlop J l€ql luePIAe 'lel3,roH '. ,\\ou sr 1r lluuouutDP snqnttltol surcuat Jrcd E Lluo'(slulaul 'oauenbes puoaes lotpai 4 ruXuou,{i s}t ectlder o1) luaDwwzp d popp" Pus eql ororJ s/sralaf,.i)Luos 'd puB sllou.tard qloq pe^ouel snrll 8ur^?H 'stotutatp nBa1t4-s!lD.ttds 'dr.ota srxa.raoJ ecuenbes aql el€lntsod ,{"ru et\ pu" slpllds eql qllA\,(llp"el 'J'rolct?Jeqc eJour $lurl sryDLuoqt e^ooJ3-pue€8prJ eql Jo slseq eql uO Taxonomy of Western Australian Pandanus
Tentative Key to species and yarieties l. Phalangesslender oblanceoloid, small onoslly 4-5 cm long), usually l-celled or somerimes 2-3-celled. Cephalia when ripe subglobose-ovoid. Staminal phalangeswith racemosely disposedfilaments, these conrparativelyfew and wide-spacedon the colunn. Habitat along streams SectionSEMIKEURA (One species,P. aquaticus) l. PhalaDgesobovoid to subglobose-rotllnd,often large to massive,always of more than 4 cells and often with 7 22 cells (or more). Cephalia when ripe ellipsoid to subglobose. Slaninal phalangeswith iufierous crowded racenosely disposed filaments. Habitats \a ous SectionAUSTROKEURA 2. Phalangeswith severalto many supernunerary ridgesand groovesbesides the normal intercarpellarysulci, in intact cephalia interlocking adjacentphalanges (mortise-tenon effect). Phalangesusually large (7 cm +), equidiametric,rotund, or somewhatlaterally comoressed. t..."nt1l:"' "::* t..: . u:-t ' massive,.:_otunll :1"' .con":l'1:i.i:iffi#,: var. dar$incn\is 3. Phalangesabout 5 x 6 7 cm, convex, the carpel tips ver'ylow-convex to flattened, \ hole phalrnge ronre$har larerall) conrpressed .. 4. P. darwinensis var. latifructus 2. Phalangeswith no, or very 1-ew,supernumeraay ridges and grooves, the facescom- paratively smooth, broken only by lhe intercarpellary sulci. Phalangesmedium to massive,totund and equidiametricor somewhatlaterally compaessed, 4. Phalangesmostly equidiametric,rotund. Carpels rather numeaousto oumcrous (to 20 or more) within the phalange. 5. Carpelswith sharplyacute convex tips; carpelnumber high (15 23) .... l. P. spiralisvar. multimammillatus 5. Carpelstips low convexto nearly flat. 6. Phalange apex slightly concave, the carpel tips flat; phalanges tending toward extremeshallowness. l. P. spiralisvar. flammeua 6. Phalangeapex rotund, low convex,or nearly flat, but not concave;individual carpeltips usuallyslightly convex. 7. Phalangesmulticarpellale (carpels l2-24), carpelsbroad. 8. Phalange apex dome-shaped,carpel tips litde proninent. Lateral facesof phalangesmooth; carpelnumber hitl tH '.t',?Xii" ,,^;. "oi."i; 8. Phalangeapex dome-shapedbut the individuai carpel tips somewhat obtruding; lateral laces of phalange with a few subernumerarv ridges and grooves; carpel number noderate (9 11, peihaps more) l. P. spiralisvar. thermalis 7. Phalangeswith mostly5 15carpels -. 1. P. spiralisvar. convexus 4. Phalangesmostly subcompressedto compressediaterally, not rotund. Carpels mostly 5 12 per Phalange. 9. Phalangesabolrt 6 5 7 x 5 cm 2. P. oblatus 9. Phalangesabout 5 5 x 5 cm 3. P. semiarmatus
L SubgenusPandanus, Section Austrokeura(Stone) B. C. Stone, stqra. (l) Series Splrales Stone. l. Pandanusspiralis R. Br. (Fig. l). var. spiralis. Phllangesbroad, rotund, about 6 x 7-8 cm, m ulticarpellate, w.irhas many as 19 23 carpelsper phalange;apical suturesshallow, individual carpel apices very low convex, hardly raised above the general domelike phalange apex; lateral suturesfew, shallow, short; stigmas I 5 2 mm wide. 242 Etz '(dNri^rllalPPsft(I 'S161/8usCSYuroJCl stlDtldt re^stP.ttals sn"p".aJo (slgordpus dol) o8uelEqd Iorn;U Figore 2-Pandanus spiralis vat. convexus. Above: habit and habitat; below; a phalange. (From ASG 12634). 244 siz 'sltdr?3 ZI rllla 38u"Fqd I :sledr?c I I qll^\ seau?leqdt:slodr"c 0I qlr,ia.se8uelerld I :spdier 6 qt!.4\seau€l?qd Z :st€dr"c 8 qlr.& 6e8uzleqd g isledrec 4 qtl,^Aaa8uEiEqd Z:sledrEr 9 q1r,n r8ueleUd I :sllnssl Sul,/tolloJeql s3^13unlle -qdac e$ Jo srs,{lese :s{Eed o,!\t s.{\oqse8u"l"qd Jed :eqrunu 1ad:tc eq1 ',(leu?^ srqt sJelJ€leqc leuo[IPp? eq ,(eur 'urnll?qdeJ Jo ' eql uo euloq se8uzleqdJo requnu llBurspue urJ [I x [I lnoq? urnribqdac eq1 '(pdrureun'1sbui1vs:^Prl tlnp€:ql pu?'^PlJudrpnsn s€ 'se,nee1 llsrus 's3^eeJ eyrue.rnlaql) opr.,'\luc A ? ,{luo stlnp" Jo esoql rePuels oqJ, (Hrxad) 'rlrq^{'esgo89s l|Dauuax !_:aut_:Os X 916I lsn3nv SI :(uoseos)1r^1 SurJnp ^€8oq strelr {lsnP qll^\ 'i>1eo: I'rolleu ol pelclrlser lios i-x"qpa.1 seloq-jele,ll go sor.r:se Suiuriluoc:3ro3 lLetl.ts'Lu :(s;auei€qdei.; gi1qir,n s1'nij'snoJn;18 e8erloJ 'ulrit sLtrsls s ol euld^\ercs .seomip, lol:ecs''S :orloor8 Jo lseo-qlnos(A ,C0 .€ZI ,92 "8I) Suuds senSo'I:VI'IVUJSnV NIISJSS,^^ 'slsolsq,ns 'oJqnj-ocequern" oldJeJlrod :snqllsejet srreJ snqllEjrlq sllnlns relljEln-8elIr 'e.tecuoi-osserdSpla^ srnriel srsope,r.redsinlns adope,r|1e;s|p airgjortns'r€ssoldop_oleu€ldtuoc r5l1pcrdt'ie1e1ruc 9.9 S'oe8uoluIJ t t S.t snbuinlold'relerprdrerZI-9 sa8u€leqd '0899 'n'l) uI edflosl qreH u|y 's8ur.rdgsen8o-I t Daauax HIdgiI 16 '3uo1s 'g y :adt1o1o17 (? 'e sernBIC)Aou Je.\ 3 g snaurusu J?^
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'OBO' 'SZ6t (HIdSd) glgt'8t€t I Dauudx.? X lsn8nv {red leuoltEN eleps,{rc :(HJdEd) t€9ZI2troa9 S"X'tl6llsn8nvSZ:poroseq'u^\orq-0EuerodollLnrJ:ll€ltu0leoJl :ilixsqns:,(pues ^Icoi'{eerc Suole SuISuuJ',{llnD ureg'alE.ol etu€S:(HJ{ad) It0Z 'lsn8nv lsoroJ qpau aX'l t .6I tI :?ere stql uI uoruuoc :dor.lno ouolspuesur looo lBnplser :,{elrequr) /A'e^roseU ro^rU luo8ed ecoIJiI :VI-IV{ISnV NU:IISE,{ .suolltallo) $aN '3uo1ruru g-e s€lu8qs:,^AoJ '/t Jo auousornlns :peue]t?Usdrl olleqsse.rn]ns IPJId":38ueteqd lelol?l 'urc ledrec 'punlor 'P?orq Jed slodrec0l g tnoqB qlr^{ L s x 9-9 srbusleqd 'IUg 'J '"ls '0t92 qreH ul N EunS?'IJo s ltu 0z '7tr1 trtra4y'y:ad,(a J'rIV Qg6I)9l tJSJrrEd'uqof lS irSal&/d'u,{S 'V6tZ '.h '(16951)eylarlsny urotselA's8uuds suellrq plDnSzJll /1 :adtJ '01 't8I 'lcS 'ur{of 'lS J (tgOt)St Jrted snxanuo)dr :uL{uoISEg '(7'3rg) 'ruols ,rou'l€ls J q (uqof'15)snxa,ruoc rv,r '(gIUAd) 'o^resed u s I oauuax I X'VL6I lsn8nv 6I .Io-^rU]ue;tu 'aruud :(uortcollo. u,,.toJg;X eq1uI a€.{leslcoldse8uelEqd) (t{Iugd a8.toa)'S V '{r€d )'(r's C/6I lsn8nv IEuort€N ro^Id eleps,{rc :VIIVIIISnV NUaJSE,{I .suolt)allo) aN Flgtjre4 Pandanusspirulix yar..flammeus. Above: habit and habitat;below; cephalium. (From typelocality and holotype).
The stigmas are small and obscure, difficult to make out in the ripe phalanges; most-are 1-2. mm long. Several phalangesexhibit abnormil, peripheral, dwarf or undeveloped, partly free and convex supcrnumerary carpels. 246 L'Z '\HIU '3r't', 'Lu 'lllq ld) 808fI 'S 'gL6l 'asPq pel 'u,{\olq_e8ueroeled lrn{ oel} euo}spuesJo esEq / lsngnv 9I ]e 'lltolLldrersg 8 's,20.9I 'Iled lE EerEe8Edoos're^lu elEPsf,rc uosreJ'a,6t "9zI leuollEN ra^rd oleps,{rc:(gagja) uir raruax, X'vL6t 6nV'e^reseu re^ld luo8ou o.urrd :lsdrooia qrr,'rr-secijp:J1ej :pessaioepqnianbrlqosdrl lrdr?J) (HIUfd) 1r9€l )3.toaU S V '9t6I lsn3nv :puelpoo,{\:puEsut :luacrrdslrlloor lPlrrE'Por llnlj'ur 8 ol eoll:4,6t 'S 0I'Ired ilaN .9ZT ,6t .ti lEuorleN ra^IU oiepsxr(l :VI'IVdJSnV Nt{AlS3,t :suoltDllor 'epr,r ruru g-g. E sEul8lls :selnlns l?lledr?c '.{lur?ul ',{1uo -relur ro Sutsuduror 1nq daap ptle psllPtu llJ^\ seln}ns IE.Ielel '^o11uqs 'prdPqs-euop s!:n1ns 1i[ ,{Feeu sdrt [ed]EJ xrdB eSu3l?qd le5rde tlur :sladreJ Il 6 tnoq" qlr.r S.8-8.9 x 9.1 I'punlor'p3orq sonu?lEqd 'tug 's8utrdg 'tttgt qreH ul I N'U selSnoq loH o)elg l S 'd'fi '8tI '€Zt 'uqof I (2961)9l lcs lrr?d 1SstloLu.taLl! 6r ;u{uotseg '^ou lels'JUols J B (trqot ls) sllBurJqlle^
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5l
Figure 6 Profilesand top views olphalanges. At right, upper and.lower: pandanusoblatus (from ASG 14084). At left, upper and lower; Panlanus semiarmatu!(from KFK 4514). 248 6nz '3uo1 tuLu S t-t sstu8tts :l?nboqnspu" €l-II sledr€c:uc 9.1 g x 9.L lnoqz sa8uzpqd :se^ool8 Pu? se8pu qlr.r (seprs seprs o8u"l"rld :snor^qo seJnlns l"l_a]?[ leutpnlfuol ledrec) 'e1oq,ntixa,ruoc :snor,rqo iirnins :ie,ruoi e8ueleqit go xedrl sdtl 1ed:zc iecicie 'drecrde :dJ?JosouIESeq orIS,{JeA IJrql qI^\'punlol'o^ISSauI sobueleqd 'rEA '(8 ', sorn8rC)slsuoult\Jep '(f 'f 'ul^rr"O ' Ug) N Jeeu L76I ur ua11y:arl'{'; 'Wl 'J'LIV 'lcs 'uqol 't (296I) 9I Jlced lS slsuaul'$rBpsnuEpu8d DlnalollsnY seIJes(z) 'vt.torJat d (PelePJ ,tlesolJ ,{lpreq) eq1 slu?ld pele^rllnc ur pecqou uaeq osle s"q pu" aeJl o1 aoll Jo '11e ruo.rJsolr?A lr ppo sr tr q8noqlle Jo3 1a lg8ra,t ou ^JJBJplnoqs puB J3lc€Jeqf, cgrcads e sueeru ou ,{q sr (perurrun uolJo ur8rEru euo-sul8l?Lu JBel eql uo sel{cud Jo acuese.rdI?nboun) ,rnpwtolwas 1at11rdaJqt pa}se88nstlJrtl/rd JelcEJBqt eqJ socsJ qloorus ,{ro^ Jrrrll pue saSue;eqdposslrdruol {pq311sOqt ut 'sseue,ul3ullsrpleJel"l urBuec B s?q lr le^ :dno$ srql uI selpnls lal{ nJ leUE peJJeJ -a:d 'lr oq ,{eu qcrq,r.ruorlrsodsrp a Jo ,{lorr€^ e peJeprsuoJeq IlelA plnoc pu" sllDJlds d ol esolc ,{.re^.sJ ll sdJ?Jopuee^rss"lu aql pu€ soJldeJ€ol peurBun 'lces 'J sll{q u,{{oqs sE'DJn lo.ttsny Jo Jeqtuetu " ,{lJBelJst snlDwtptulas 'uoucolloc u.troul ,{1uooqt,^^ou se,|.l.(ed,{1 ot{l) uorpellos e{?lg 'd lrlun eq1 puzlsuaenf Jo slsuaps.rawos ol serceds srql se]eler uqof 1S 'pu?lJeluIq pue ls?oJ uBrl"Jlsllv rl ou eql Suole uoqnqutsrp Jepr,t\ uo^3 ,{lq"qotd lnq luellrtujelur ul.|s€q sertrSdssrql lqnop ou pu€ e^rsseJxalou sI ecu"lslp eql lnq '(urMJ"O Jo g lsnf qefurdlooy) ,{JolueJ uJequoN eLltur uortnqlJlsrplLlJol snor,\eJdrreq] ruo{ elou u"rlerlsnv uJelsey' er{l 01 uoqrpp? u? selcedsslql e{elu sp.rooeJeseqJ peceld ,{lerns ssel sr 'lrn{ ernlEultur ,(re,t e go Swaq'(76761 'sntDlu.totwas atloat) uorurceds pelrJ ls?l eql qSnoql d Jo uoll"Jlsnllr pu" 'Jeqlo slsouS?rperl] qtr,r pu" qc"a qtr^a lle^\ ploJJe sueurceds eseql llv '(HJd3d) dS.nat '9L61 'uaar8 'qslnlqf,llqElls eAeIloJ 'llet 'purJpoo^\Z0Z 'S S V lsnSnv€'slecol lrnrJ rLl9 6oJl urpues ur ,91"SI':] ,rt .9ZI eLu€S:(HIUgd) tl9t ,1//r/ {r) 'S/61 pue oipoBuole uolLrluoa'S,tt 'a,ts 'tJ tsn8nv'al€.ol0Z'slood lueuuer,{q )aercJo '9161 'Iearc tL "9Zl ells etU"S:(HIIad) 8I€t 4/Darrlry tsnFnvfl oprseqS /6t otl 'A slrs'etecol elues :(HIUAd) I Dauuax'Sl,6t lsn8nv 'sloodlrPuluar ,Sg,97,1'Zg 'poq)eorJ,{rp 'S n]f 'g 'ol€.ol OI punor€,{luouru-Iotr Joeprs ,l.t "tI ,90,tZl aues:(HIdEd) 8€lt ,(In,)uuax '91.61 'arolllluocxre^ '{oer) sosuqot luEuuor e8peuo 'S,lT !'a ). tsninv 6 ')red Jo lood Jo .Sf ,Zt ,Lat'tS etrs lEuorleNJa^rU aleps,{rc :Vt-lVdISnY NUEISS,AA 'a 'qefurdloo; '0dg) 51 uro{ y6961 a>1o19:aclt1 '(s9 'uqof 3rnirl) ,tt J'lZV Q96I\ 9l rcs Jrc"d 15 snf€uuElruassnuBpuBd t
'rrlr/9o tllll^ 'd d l"rrluapr s" (62 J'019 (1961) SI rrs Jr3ed) uqol tS srsi/alr.s./z4os reprsuocI 'roqlouE Jo setredsouo jo ldJJuoJoql Jo JepJoq euerlxe eql le sr qJrq^\ur.IoJ ",{lsnol^qo si ll ll ol pelElsl.{lssolJ Jo'snxatuo) 'JE^ '.ta1r"Ilds sllotlds d Jo rrrJoJ elrrednrdep ,{ro,r e flqrssod sl lI les uJ pecrld Sureq s" JeJosursarJeds srql qltm puodse;roc o1 :eedde plno,,it lI ssJeJ '(3uol eSueFqd qloours eql Jo asn?ceqq8noqll? Luc I uaql ssel) se8uslsqd rplncolrcned pue ll€u.rsfre^ eql Jo esns3oq ,u/r,/?ds J Jo ldecuoc (pauop€oJq ,rAou)eqt urq}r,r popnlJureq ,(lrp"ej louu"c l1 a8elsslqt te uorsrcepe e)"tu o1 olnudoldd? psuoep sr 'dnol8 pu€ e,tttrdecepslr{l uI setu€u lou 1r llncurp 'seruBu serceds (,{Lu,{uou,{spuB) ,{cu"punper tuepl^e eqt pue peqstlqnd 3o 'serJeds fpeerle 3o relle,r eql ur tnq A\eu€ eq ,{"trl ll ueese^?q I qalq,{\ sueul Figr1rc7 Ponrlanusdalplrer?srJ. Phalange,profile and top view. (From BCS 10650\. 'o resou re^ru lue8eu e.ulrd'II'ZI 5SV 'runrlpqder pu€ so^eol 'qouerq 3ul^\oqs ']JqeH 's$uautluDp s aDpaDd-8 afi3\4
'gVI 'J'OZV 'uqof '75 (2961)9I lrs JIt"d snDn.4ltotJ:lu,4uolsog '^ou ]els'3uols'J'g (sqof lS) snpnlJllsl J"^ 'pools.l0pun Ile.^d.lou ',{lrpa.reg 'aJu?Jgru8ls 'Jetc€r"qc sr slr s" II?,\{ w s}I le,{ snol^qo ,{JeA3 sI }I 'pelc0J3 'urnrpqdoJ dllenbe sr e8urleqd f:a,te 1ou pue " ulqll{\ l"q,\\etuos ,(J?A ,{eur -(sdeqrad a,rqcege,{1par lou) SurlcoFetul uouel pue asllJotuJo pul) ? spJogB qcrq.\\ relJarsqc eql Jo aruaprJur aql tBql enll osl€ sI 1I uo4"lnJsos laeqs sr srql esJnoc :puof,es eql ruorj ura1s,{se,too:3-pue-e8pu aql 'ls.lu oq1 ruog Jo 'laltlI 'd 'J soSuEl"qd eqt Jo ,{trpunloJ pu? azrs e^rsserueqt pre slp,tlds LU1JJ sJetJ"r"rlJ Jo uorwulquoteJ E lJ"d uI eq ^ew slsuaulltt,lopdr'peepul sa^Il?lel slt puB raJtLlAg,{llercadse 'lsee eql ol D.tn p.ttsfiV reg ut setcedspal?lel qll^\ 'setceds uorl€zrpuq,{q tuo{ pa^uep sr ralJ?J€t{csn t"ql etu 01 slBedde 11 eq1 ur rnJco u?c seSueleqdoql stuets,{se.toorS-puu-e8ptr Jo Jo 's.rynld.t l€uIpnlfuol IBJaIBI Jalc"r"qJ aql l"ql o8pal,a\oulJeI d ol slsuaw(A.toPd 3u4€lel uI
'099012ao$ ')'g 'Ia6I 'ur.{1reoreau 'IeerJ poo^\qsec:.\uorIuusl{R l99OI ldes t 'S '9161 -HltON (HJ,Ugd)Lgrtt'9tttl aSnat U lsnSnv'Ir€d leuotleN$^!Uepps,{r( :(Hj,Xgd) us ,tpaauax aures:(HIXfd) a8loaD 'tl6l 'a^iosouf t't16I rsn8nv€'ole.ol lltzt S Y tsn8nv 6t re IU lue8aderulrd :VI'MfSnV NXEIS1,^A:suoltralloT I/.aN Type: H. St.John 24225,22 mi. S. of Darwin, N.T. in BISH. Phalangesbroad, rotund. about 5 x 6 7 cm, with 12 20 carpels per phalange, apex of phalange dome-shaped, carpel tips very low convex to flattenedbut apical suturesdeep and rather wide; lateral suturesdeep, nurn- erous; stigmasI 2 mm. Very sinrifar to vat. aullrcmensis,also to var. clervinensis. Ne$jcollections: WESTERN AUSTRALIA; West of CapeLondonderry, 5 August 1975, A. S. Geotse11375 (PERTH).
II. PandanusSubg. PandanusSect. SemikeuraB. C. Stone, Contrib. Herb. Austral. No. 5- (1974\42.- This small, characteristicsection is composedof two subsections;ofthese only one so far is known from Western Australia. The other. subsect.Elaphro- carpusB. C. Stone, is monotypic, and its single species,P. basedowiiC. H- Wright, is known to date only from Arnhem Land, Northern Territory.
Subsect. Serrikeara Type species:P. delestangii Martelli, a synonym of P. aquaticusF. MuelI. This subsection,which I consider monotypic, rangesright across north Australia from Queenslandto WesternAustralia. Although five specieshave beendescribed in this group, thereare no usefuldistinctive characters. althoush one form, namedP. kintberleyanusSt. John, has fruits of which the drupesare rather short for what seemsto be the average. With Blake (1954, ll0, pl. 7, f- 3) I considerP. aquaticusthe correct name of the species,and although slightly variablethere seemsno justificationto recognizeadditional species. Somenew collectionshave recentlybeen made, cited here.
5. Pandanusaquaticus F. Muell. in Hook. J. Bot- Kew Gard. Misc. 8(1856) 329, nomen provisorium; Fragm. Phyto. Aust. 5 (1865)40 (validation);Srone, Contrib. Herb. Austral. no. 5 (1974)42. Syn.P. kimberleyaLusSt. John, Pacif. Sci. 15 (1961)180, f. 9. Type: Fitzgerold 2395, Fitzroy River, Kimberley, W.A. (NSW). WESTERNAUSTRALIA: Drysdale Narional Park, 15 02' S, 126 55 E. commonon riverbanks, trees5 6mtall, ratherslender. aerial rootlers present, fruit fallen,6 August 1975, A. S. Georse13486 (PERTH): Same locale, C_arsg.n !ye1, 14'49'S, 126'49'E, 1t Augusi 1915,A. S.Georye 13705 (PERTH); Same locale, Site C2, l5' 03' S,126" 44' E,very common, l7 August1975, K. F. Kenne.illy4388 (PERTH\. Of these specimensthe drupes of 13705 are like those of the form called P. kimberlel,anus, but not quite so bluntl while in 4J88 the drupes are more slender. as in P. delestqnsii. Addendum There is in the W.A. Herbarium (PERTH) a collection madrc "Pandanus bv E. M. Bennett,no. 1815.labelled lectorius" and consistingof sorne phalanges of what is clearly a member of the Austokeura grouf; however locality data are lacking (collected 20/5/1967) and I refrain from describins- it- It is almost certainly an undescribed new variety of Panclanusspiralis.
Acknowledgements I am indebted to the Curator and staff of the Western Australian Herb- arium, Perth, -for loans of specimens and photographs, and especially to A. S. George and K. F. Kenneally for their interest and cordial asiistance in the 252 '6'Id'OL-Lba stl ot.at snuDpltDilur 'seprrqeH I€ Ilng,{\ex uoqelre^ Jll!€dserlul uo ^esseus qlr^\ ^\eN eql Jo oeoJeuepoed6qI 9L6l- '0ts .qrr{ef.tog 6st: t6 ls^s leeeoeuBpu€d)sn npaDAul uollecglssel. tueueS€{ut pa^ordtul ue pre^\ol qtl6l-- 'tt-lt'. oN rlsnv qraH rluoJ (eEao€ I 'etL6l-- -\tepued) D.rnbllwasuo]l'aes snuopuDA Jo sorcadsuerterlsnv eql Jo aer^er v 'L9Z IEZ'. ZZ erode8ursIIng supD I'eP.ieuepued uelsolet\lJo solpnls q/96I- 'tt6 'sn 6t6: t9r.uols .lo{ I rauv DPuDdur uo!.rollrf,cluouoxel € se requnu ledrq) e/96f J g ''t t-68 : €z t s, Jt ed DuattsotrtN uorlrosaql ol ^el e pup'soraedsuErlErlsnvJo slunoDv raqunl:€t tred plql 696I- 'tzn -ZIt: ZZ r.S JrcBd 896I fq poqsrlqndsor.ods uerleJlsnv oqI:82lred plql 896t-- .0€9 €29 : IZ TJSJTJed srr./rpral Jo s)rJadsuer lErlsnv .rql : [Z UEd plql tg6l- '8zt-60n | 'rrs ',ftoluJal 9t Jrc€d €rlerlsnv uroquoN e\| ur snuDpuDd:€I trEd prql .2961-- '9lt 16Z:9I IcS JI.ed rrlt 'rlrd' puelsueeno: ZI lred plql qZ96I- '18-2. : 'puelsueono 9I tcs lrcEJ €rlertsnv tuo{ sercadssrrrprrJ ^.\eNeorqf :6lred plql EZ96I-- 'sag*t9s: sLI.s JrrBd erlertsnv puelsuaano luo4 sanads snuopuoJ l.t'aN :9 lred prql qI96I- '98I-08t: r.S ',,{oi11roJ -prqlSI Jl]ed uortras eq} uo saloN pue erler}snv urolse^\ ur rrrrprrd :Z U€d EI96I 'Itz I rcs 'suorlces nzz tI lrced eql ol ,{e):Iu€d urlalls srrrplDA snuaS aqt Jo uorsl^eU 0961 H'uqof tS uopuo_I 'ueurorc '0lgl u€^ eelnsul lo oerpuelloH oe^oN oeroll snluorpord Ueqo{ 'u^\orE '0tI : stueJd uerl€Isnv urequou sno.u€llessruruo serpnls ',ta^rns66 Z log f lsnv Il'e{elg leuor8ell €rlErlsnv urrqtroN eqt Jo suoltnqutuo) lerruelog tS6I I S saf,uaraJau
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