080057-12.002.Pdf
eJotr,t'rollerxs ,e,,ir1c cq-L asuceqleq 1ou.{erusr q11nq u cred not? sulsa,r,ta11aq1ur ux€t er{Uo etuos l€ql peloprsuoJsrlrluouloru0qlle,eldLuexeJog.dno:Fsrq13o,{LuouoxulaqlBurpuelsrepunur,(lqereprsuoc d1eq,{urupacnpo:lutueeqe^pqqcrq^\uxe1esoql,{JrlueprolelqeBureque^g.le,relsaroedseqlapaceld ,e)uenbcsuoc .ele ,{1yolca;sr1eseq louupcr{orq^\ suorltollocJo lcq(unu E € sp sleqtpue uorlnqrJlsrp ur depe,ro sarcedslo.requnu e eleq,lr€rJ€llsnV tuelse/A Jo suer?ulalse^\-quou eq1 uor3 sueurrcadsq1t,.,r fllncgjrp relncrgt:dsr ,selpnts otoql pa^arqcc eq uec uorlpcUrssulJ alqets u teq1,41e1r 1un sr tr pleUqlr,r peurqtuoc'serceds pacnporlur ,{llsoru oseqlJoXBolotq eql3o BurpuelsJopun .rau3q E sr a.roqtIrtull 'luoruuol,ruo fq pacuenUulqcntu puD cqseld (.ra,r sr uorsso;dxe g 'pe1zlosr lucrBoloqdroll ,{lleorqderSoeBflsnor,le.rd are,tr qalqA\ DXD1 uoeA\1€q Bur't.tncco st uotlezrpuqXl .Z 's1sa33ns l uorlergrssulclueserd eql uuqlprleJlsnV ur eraqluesard cxel ssoI.lo aloru s]€ elaql .l :srqluruldxcplnocsctlrlrqrssodJoreqlunuV.sJclcuJ?qcBurqsrnBuqsrpelqurlarpuuolllncUJrp uogo 'scrceds sr lr t€rlt qcns uao,NeqsecueJeJJrp eql epore qcrq,r sueutrcedscru ereql lnq uoxul t?ol peu8rssa eq u€csuatlJrceds Sutlt n4lsot! fl,4s1iqJprrcsqDluaprrro l uea^\taqpu E s.tlDtuaptxJo.Jpue saptolst ) snuDualr1Jla pue.rap?ols7J ,xelduroc J J I uee,{\leqBur qs rn8ugsrp ur pue r.l"i7sa,r,ra77 eq1;o extl eq1SurtrLurlep ,{1uolcu;srles ur ,{llnruJrpppq ueUo spq Joqtns aql tnq srpo,{Jorequnu e ro3snua8 eql ot uelul qJ€orddeleuortrpzjl eql sl eJaqpelueseJd uorlEcrllsselc sq.L s.)rredspelula.r pue rrr,is,{4 's!lDluaPlrro 7 J'sapiotst? J'strlsa al J a.r ltaof,,{urdsqlr,r sarccdsasoqljo ,{lrclncrued.epr,r -pl.ro,!{uorsr aJpelrDlap .1 €Jo paeuur qantu,{le^ sr (e€e3Dll,{qdo?lT) snlnqtt1 Jo ,(ruouoxplcrll
uo4JnpoJlul
'sstoods .r\l Iere^osJoJ peluu8rsep a.re sed,{1ol3al pue petuprle^sr Je{JuU X xa relqcrg Hs/rs.)/agts T xeldtuocv,r;rsa,/./aj J eql ut ex€l eql.Jo uotssncsrp e sreleql puednor8 luql urqlr,r ruorj peqrrrsep,{lleuuoJur elE ExElo^U_ uorsIAOJpuo,\\pJopaauut,{1e:ossrxcldwo)stqs Dt,Iaql Sutpuno.uns .qlueBx? .l sercedsesoqlJo ,{uouoxut eq} tsplluoc ul llenhl.J r?dtnco,rcrrw puu.qlueg ,Je).lpg.W.dsnquoJtlppD ..1,selcads,reu sntnulq J olp3l€leJ,{lasolclsou €Jo uorlrppeeql Luorj 'lnc ''l uude Jeelcsr st4saltal J ol pelplallou ereqctq,n sarcads e,,rrleu eqlgo,{urouoxel .sarcedseseqlgo ,{3o1otqpue fuouoxul ,ur;erlsny ..| eqluo se}ouqlrl\.roq1e8ol urlncco qcrq^\ .\?/ng7rJJo serceqsasorJl ol petuaserdsr,{aI IEg} V (g661)SE-6:(1)3yurs/np.snso^qnssnl(lu.L1o uorleprle,r pue sarceds .treuouo ?urpnlcur'erleJtsnv .Je{rufl ul (e€ecpgftldo? t7) snlnqrtl uo setoupue ,{e11er4 V W.N
lluJfsqv
'.rerral .srl€rlsnv 000Srrlertsnv qlnos oprelapv quoN\uaprDr) rrurlog qtnosJo urnDxqrollolels _/:l
ToIJEBI I U
snsolaqnssnlnqt rJ Jo uorfBprlEApuE sarJadsaau auo Sutpnlrur (equrlsnv ur (ouarBll,{rldoB[7) snlnqltl uo soloupuB fe{ lurrf V
(866t)St-6:(I)Z I ErsllnN Nu)hil Vol 12,No I (laqR) or Iessunanncd coc ci of T.n inutu,vand T. micrococcu.\,bolh considered to benative to Australia,could $,ellrclatetolle,'/eslrisvar.inernisBoiss.orvar.bicornutus(Fisch.&C.A.Mcyel)Hadidiasdepicted "F in thetrcatment ofzygophyllaccac ( El Hadidi I 972)in loralranica" . In contrastto thecomplcxitics within sectionTerrestris, the othernative Australian spccies are usually easily distinguishable.
1'hepurpose ofthis paper is to providcan overvierv of TriDrilasin Austral ia, to h igh light the problem s andto providean interim\\'orking key to taxa.
Gcneric and infragenericconsiderations
All oitlre specicsof Tribulus(and Tribulopis R. Br.) in Australiawere transferred to Kallstroenie 'l'. Scop.by Engler(1890). Onespccies, pl.ttlpterus,prcsumably bccause of itsbushy hab it whereall OthcrSpecies were prostrate,continucd to be refcrrcdto Kdllslloenia cvcn alter the restl'rad been translleffedback to I/rru1?rs.
"the Porter( l97l ) in hisnotes on thc floralglands ofTrlDrilas indicated that principaldiagnostic tloral charactersdistinguishing Tribirlus from theclosely rclatcd genus Kallstroemia Scop., rvith which it is oftenconfused, is thc presencc ofa rvhorloffive intrastaminal glands atthe base ofthe ovary in Z/irrlrri". These intrastaninal glands afe unmistakably present in the speciessurrounding thc I telre.rll/.rcomplex and in theother spiny-fruited 7116a1zs species u'hosc taxonomy is not clearcut as discusscdabovc. l'hese glands are absentin the native Australianspccics, I suberosus, T.pldlypterus, T.forrestii, 7. a.\trocdrPuu,T. hirsttlusand T nacrocarpus, raising the possibility that they miglrt be more appropriatelytrcated as Kal/.ttoenia than Tribulus. Ho\vever, KLtllstrcemiais furthercharacterizcd by a 10-lobedstigma and lruit anda l0-lobeddisc upon rvhich the 5 stamensare attaclted,all charactersnot applicableto thecndentic Austra)ian species. Thus it sccmslikely thatthe nativcAustralian species couJd u'ell form theirorvn section within 7il6ulasbased on thc lackof the intrastaminal glands.
Thisapproach rvould not be supported by El I Iadidi's( 1978)classification where hc proposed three sectionslbr 7i.l6lhrs,based on fru it nrorpho logy. Sectionler/er //ls consistcd ofall ofthe species with in lhc 7'.terrest s cotnplcxand T. aslro.urp s, sectionl/a/a ofthosespecies rvith winged carpels and sectionlirclrrls ofspecies rvith unanncd and unrvingcd fruits. He I urthersuggest edthat T. plqtyPterus might reprcscnta nc\! genusbccause of its uniqueshrubby habit. Basedon his classiflcationthe Australianspecics rvould bc tlisposedas in l'ableI
Table I . Placementofthe A ustralianspec ies of ifr"16rlr.tin thesections rccognized by Irl Hadidi ( 1978).
sectiolt' Terraslris section rr?/rrriJ sectiolt' Aktla
711e|rcslris complcx ?T./itrrestii l . Muell 7 . p|qrypterLts Bentl't. 'l'. 7l /i]r'//r,t R. Br. suberctsus1l.Eicl.tler ex R.M. Barker T. occidentali.sR. Br'. T.mttcrocorpus F. Muell.ex Bcnth. 7. clchlerirarrrl-K.L. Wilson ?7.hirsutus Benth. f. clsIoiLlesi,. 'l a.rlrocnlpasF. Nluell. _-
'aSeurep lnoqlr,nl:rerlxe o1 elqrssodulr eq ol puno.lseq Joqlne srqlqcrq,rr'spcaseqlcJelrn{cJnluureqlSorrcocp.req,(leurc4xseqturqlrMpJSolJuA.Sotlulo{sefupl ,{lunletu otFurdol.Aepl.cocJo raqtunu IenlJueql lnq.lrn{ qlca u.rorjr..oc Setnporcl ol lurtuc}ocleql o,req sercods11y lesredsrproj rrloc.roslcllrn+ olur eiorJossrp i11an1ua,lc op urlerlsnyur sercedssz7ngr,i; eqt '(,'!\olcq Jo llPJo slln4 aqt s s.o)aqns7 rapunaes) ,{re4uol aql ol suorlE^lesqoeuros elrdseq
lrun IEstadsrC
Sur.roqreqlo {q ,{lu^ocql.}o 1ce11u a6etnocsrp snqt puD ,(tl^'cu luu eFuLrrocue0"", ,",r"rr";:tj;rt:l autl ssaccu .uorluurllou_.}les 1uro111sod6ur,no11r leql,{lrlrqrssod u oslu sr eJeql lueAelqpup sueupls 'rlrlltl eql apnlcur{Eru sletad aq13o 3ur11o.r eqt .l urpJ^Jesqo uoeq stq sc,i1e,rr1uule11e ;o uotleurllod Suunsucsnql pue ,p"qsu,1 sucLuuls^llql,(q eLu6rlsaq1 iu rpuno.r.rn"^-Josuucur r eq,{uur1r ,, _,"no11 "q1 tsaq .{rru lcql ltsu8ls ll u,roul lou sl lllsrJalJurpqcslql Jol uospeleql Je,roucql lo erlueJeql tnoqe lcejepuelspup,{llcurpnlrFuol dn s1rlad .r.,{\ol, 11or cql sesolcre,nog oqt ueq,\\ lnq ueclout e1r1_ete1d puru p€erdsapr,\\eJu erle.qsnV ,llp ul seltads.rrl/rg7./,1 lou.]r!solUjo slelacleql teqt alql]ocrlou alrnb sr 11
s1c1ad;o iur;1orluurpnlrEuol
iSolorfl
'snua8 lcullsrpe se .,{pnls.reqlnS pelt;s.tlsr lr rcdBdslrlt3o sasodtnd cql;o.11nq scrrnbaroslr ,sn7nr1r.r1p?pnlcur 'stdolnquJ ur seorr}etuos snua8uur;urlsny oluepuc .s.rclcuJoqJ aqt.+ouorlrsocl cq.1 trn{ pupspuelS luutLuDtsurlul 'sJelJururlJJo -Joef,ucsqD ro oJUescrd's.rolcureqrledes lrqeq Surpnlcul leqLunuu uo peffiqsFurclnol; ..,ErlcrtsnV.'o leuuoJurur pecDldueeq o,req serccds eqt eiolq.,eql.roj luaurleeJl FururoJrluo-] orll ul
'(^\olcq ees)s?{/rDoTapr .1 pcqrrcsep{1.,rreu eqlJolrn{ eqlJo€rnlonrls aql palerd.relur ,{1lca.uorsrq roqlneaL11y sndttcotcnar J qlr,treq ol rueddu plno,rr sdrqsuorttllors1rq8noqtlu 1p1puy1 1E,\qs4sa.ua1uor1:rosurpeluldsunsntltototls,oT.lad:ec qrecaosSut,lr aql uee,,*1eqseurds uerpeu3.Jrrd ,uoutes e surl slr{lol pauirssuos1u.r.,r/.rr ).).t)nw .1, rcuureedde peFur,nEe^rSolloqlo qoce uo ur esdpllocsllrM esaqt teql iur,{.rp uo fluosr ll .petuLlur/(ltcera 3JusllB,!\ eql eJuls.r/tzl/arl uorlcJs.,o ,elcls slrnt-+eql olrl erou qcnurrredde srTrzs,i/ry J Jo slln{ qsc{ Jlsqtul slln{ piSur,r laqlo cql ol ssccordlurre31rp e fq pe^rJepeq ol ulees,olEls ijp aqt ut pegur^\ 'ln ..in ,{lrr]clJelr q^\ trs./i{ J Jo slrn{ eql 7/lgr.r7jo sarcedsalerlsord,{11ensn aroul eql rllr^\ sppo sr 'uortres lu lrqeq ,{qqtrr qs JI aql srqt-}o slr n4 qtl,lr aql llJ^\eer8e qcr qM s1r n4 plBur,u {;.re a1c s^pq r.rsorrgn J pull sn.!aldAlDld J oltrl,f, trlp/'/ uolltts ol pousrssuscrceds uer letlsnV eql utq lt,t\ uorleUc,\osl1? sl eJaql
!a^oqE dnor3lut1l-1osput13 lEuturDlseJlur aql s^IJDItr pelplssu lnq r'/1,lja//.z uorlcesur rprpeH {q paoe ..1,.spuu 1,1 1dse,n s rzrL D)o.tt sD 1.eleut ure lsurlur equo )tul eqt (q I /41./tr1 uorltcs pul?li.//s.r/.1al uorlces ulq1r,,|\serccds 11u uorjta.lirp olrucciduplno.n ir]s-a.lJo.f.Js^tLuJaul uotlcesqceo;ddu puusourds 1noq1r,,n 1so wlrl )Je snnutLu.1,peu s-n)xo.atrurJol perroJo.r.erlurlsnVul st4sdttar ,s4.//r&1 J ro swJo-+aqrJo auos,{Fe lrur5 seurdsuerpau j o red e ssessoclscop lr lDqlldeJXe uorlf,osJoe^rleluasaJder sp(g/61) rprpeg 11,(qpejnltrdrusoIl.H jrasDl.T-1os.1rnr3eq]q1r,n,{1eso1c saat8euTsattof1 setJcdsur rlpltsn V cqtol porldduucq,n uorleJU!sslrlc srql qlr.alsurelqoJd 3Ju er.qJ.
sllnqrr.t. rt) sotou pu! ,(rt trrrr V lilrr8 tr'{)l T2 Nu)t\ia Vol 12,No. | (lq,'8)
Thosespecies which havespines on the fruit clearlybelong to the dispersalsyndrome known as "trample burrs" (van der PijJ 1972). Thus speciesof Trihulussection Terrestris, together with T. astrocerpu,\andpossibly 7l /b//',?s1rlare all prostratcand all havehard fruits lying uponthe ground. Any pressureappJied to thesefluits from abovewill first of all causedissociation of the cocci and, dcpcndingon the lengthofthe spinesand the hardness ofthe objectapplying pressure, may well result 'l in penetrationofthe objectapplying the pressure. he subsequentdispersal wiJl presumablydepcnd on theefl'ect that th c penetratingspin es have on thcobjcct. lfthepenetmtioncauses irritation, it islikely thatthe dispersal will notbc far,but in thecase ofan inanimateobj ect such as atyre or boot,the dispersal couldbe much furthcr. lt isinteresting to note the record of l9 I I (Dill & Bryanl9 l2) wherean expedition "seed" to LaysanIsland in thc I lawaiianGroup recordcd a spiny7,'ir5z1zs entbeddedin thesides ofthe toesofa Laysanalbatross. Th is was identified as L,s//'rt, which is ccrtainly incorrcct. Itismorelikely to havebeen the coastalspecics, Z cistoides.lt is probablybecause ofthis dispersabilitythat there is aproblemwith thc taxonomyof sectionTerrestris.
On thc otherhand thcrc is lessofa problemwith thctaxonom), ofthe wing-fruitedI|.16z1zs species in Australia,and this may relatcto the shortdistances over rvhich thcy arc disperscd.This groupof speciesbelonging to section,.'1l.r1.r produce 5-winged fruits which at maturity, break into five 2-winged cocci. Whileworkers in Tribulus(e.g.El I Iadidi 1978)comment on theunusual(fbr Tr ibulus) upright shrubhabit ofL p/rrtlpterus ard T.suberoszs, dispersal ofthese rvinged cocci fiom a heightwouldseem to bemorc cfllcientthan dispersal from groundlcvcl, and it hasto beasked what lunction a wing could havefor a fiuit rvhichis alreadl,on thc ground. Theonly timewings wou Id sccmto be advantagcous inthissituation would be iftherewere no competing vegetation and wind was a f'actor in the environment. T.macrocarpus is onesuch prostrate species producing winged fruits although the rvings are nuch more thickenedand not aswide asthosc in 7'.pl.tllpterus. It is loundin mulgaareas on redsand and under theseconditions it is quiteeasy to envisagethe circular fiuit rvithits 5 wingsas the primary dispcrsal agcnt.Subsequent dissociation ofthc fiuit intoits 2-wingcd coccigives a secondarymeans ofdispersal.
For a discussionof the particulardispersal characteristics of Z srberoszsan d T.platypterus, see underLsllerosirs below.
Scavengersof sccd
ln themature fruit it isextremcly difficult to removcsccd from the hardened area ofthe coccus which surroundsit. T.eichlerianus isknown to be a lavouredfood source ofthe Red-tailedBlack Cockatoo (D. Albrecht,pcrs. com m. I 996),and it wouldnot bc surprisingif T.forre,stii and T. mquoc.trpus w ere alsosubiect to attackin a similarfashion. lnone specimen ofZ szberoszsat least(-D ichler 23606 p.p.), the hardenedarea about the seedhas bccn ncatly opened and the seedrcmovcd, and it is prcsumcd, becauseof thc strengthnccdcd, that this was alsodonc by a cockatoo.
Littleevidence ofinsect attack was tbund on themature fruits ofI,'irrlrr, butit wouldbe surprising iftherc was not soDrciutelest shorvn bv borersand/or weevils.
Possiblcant attractants
lnT.platrpteru.e,thc area exposed when the fruitlets separatc, padicularly the areacovering the seed, iscoveredwithtiny gland-likeexcrescenccsrvhich are here postulated to bcfood reward forants. l'hese glandsare also to be lbund in 7l "!xrelo.!/Abut do not appearto be asnumerous, and tlrcy havc also, somewhatsurprisingly. been sccn in somenrembcrs of thc 7'.lerreslris complex. T)regland-like cxcrescencesappcar identical u,ith those ibund on SrTaL. (Malvaceae)cocci which aredispersed by " ulsarJoJ'I'9 [1-9srredlegDe.].V/AJoEeJu,{Dg)reqs/uo^J?ul€:l] 'seurds uurpeluJoJlEd E qlr,upuI] popunoJ .pe)luLu ,{lot?lncrleJ 'qtoorus a (llesrop rccoJ:tuecseqnd ,{lasruds ro snorquli 'peqol-9 .snorqelSqns strnlJ 3ur{tol spuelSlpurcue1sur1ul slul?ld / poiur,\\lou slrnr:l 'I 't ...... snqluurutapB [(9)!-_t( e) $ rudlrlJu..t 'lesec uosqlCol q8norqlSutpuelxa,{lqrssod'Eer€ rrouoe'l'uBul,{\eN .u,,lroulun leu8al lunotl,tr]3uo1 ruu g 7-g.I eru8rlspue a1,(1g elDls 'uorlrpuoc qse{ ur prup ur clpprru,\roleq lsnlsaurds o1 luelu,rrnbe seJnlcnrtspaucpJeqqtr,'!\s8ur,r:epr,rruul97-g I,q8rqruru y1-01 strnJC :9 " ..... snlnsrlq.J.C [(g)9-g(g)srred legue-t JN ureqtnosot V,^Au! ueseC,{pueStEeJC puplc^rU lppnU ot qBnolqtuorBej felsroruel I] '3uo1 ,seurds tuurE g etu8qspue elflg elqs qso4ur p€tullur,{rc,r ro 3uruepll]q ,{up Surltul sFur,ulepr,nruugg-67.q3rqruru 3g-g7s1rn,r1 9 's8ur,r saurds3ur1ce1 uao,rloqluocsaqnd,{lasuep slrnrq .sno[lAslupld :g .; .9 ...... sndrecorruru 1/-ssrredtcuee-l 'purs pJJut ,{11rnsn'ygulJquou pu[ JN ul.qlnosol puelu] puu uogpleregol eBuuXode3] sBur,neql ucc,nleqseurds uurpotujo rrede q1r,rlueasaqnd pessajdde .{laueds slrn:1 luacsaJqelSstueld S ,peiur,r rccocpe8ur,,ir-7 a,tg olur Surprlds,{1rrnleru lE sttruC t 'lecr1rutu,{s peduqs-.relslou tnq ro relnia.ll ,pa8ur,n-g.ro,peqol-SslrnrC:t .l ...... snd.ruro.r1se.1 l(s)t( t) srudteliue.I 'plo?t\S puEVS ujeql-lou'JNujeqlnos'Vy'A ut uounqlc/{ ol uo^reuteJ uror; spuulpoo,ue31n6] e^oqp tuo.Upe,ler,r uaq,npadeqs-rets stlnJJ e sqreqolerlsoJd :l . '''' ''''' ..[L-gG) snreld,{1u|d'1 I sllEd]ollee.I.y16 ,uorFaX,{a1sre ure gl 'sFur^r uae,\\teqtuecseqnd,{lesleds,SullDraossrp,(11crnb slrnrg ',{llurxupp snollr^,{lasuapsledeg lruq f1.roc3ur1ce1(llcnsn suels :Z """""""""" ,uor6eu,{e snsoraqns'tr'2 [(S)t-Z(I ) srrDdtel]ea.I V^A lsreule Hl 'sSur,n uee,tleqtuecscqnd ,{lesuap ,Sullercossrp ,{llprul slrug .,{11erxepa .)teq sno1lr,rflrqcled ro snoecrjesXlcsreds sludas ,{IJocqtr,r\ suels Z 'pe8ur^\ fsaurds,{ue 3ur1ce1 slrnJJl seqsnqlq8udn t 'pa?uuqcun e.tusqlSucl e tl€leriraql pueeurl eulosloJ lsts.redflclurre,lur eru8rls pue el,{ls eql ecurstIn.U aql ro Jo,rogaqt ruo4 pe8pnfcq uuceseql lnq luelrodur oslucte sq13ue1eu6rls puua1.,{1s oArl€lo1 strn4Jo sre,lrouJo ecueseld cqt uo flel seop,{eI eqf .uorlnqrJlstpeql qtr,r Buolp ,sercads crll ,sJolauJprlf,JEel Jo qcDoroJ JalcEreq) ,fuDrprsqns Ew pepnllurutaq spq slalJDelJo lequInu eqllnq tsoru ur dEIre o sr a.reqleours ,{lolceJsrlus fre^ aq plno,r s/?/rgr.Uol.{eI e,lrlula8r,re lpql flelrtun sr ll
BrlBllsnv u snlnqltJ ot ,{aI o^!}E}uaJ
',{lrunruLuoJeqlJo sa,trluluaserdarra8rul oql,(qaqote^uqplno,,nsluu,(q1uerue,toLtI,(ue,lellrnJ.]s4.ro.lagr.rJeqlJoezrsaqlua,rrg.Buue;.;oeq ,s1uu tq8lru ,4eqlpre^\eJ eqt olul trlSrsurup JoJpele8rlse,tur uc:q a,lEqol le^au.rEeddespua13 qons lnq
'r.Irr8 lnlnEq |o s.lou pue ,{al lprJ] y N U IT NultsiaVol. 12, No. I (l99lt)
7: Plantsvillous. Intrastaminalglands prcsent. fruits irregularor 5-lobcd, sparselyto dcnselypubesccnt; cocci dorsallyspiny or tuberculate, rarclyrounded, usuall)' $'ith morcthan median pair ofdorsalspines 8 Fruitswith manyspines distribLrted randomiy all ovcr;cocci notbilaterallysymmetrical 9 Fruitswith spinesl0- l7 m long:pctals l5-30 mm long.St)'lc 4-5 mnl long.urruits up to 30 rnnrhigh and 50 mm wide.Redsanddunesof ccntralintcrior ofAustralia. Leafletpairs(a)7-9]...... 9. T. hystrix 9 f ruitsrvith spines less than 8 mm long. Petals6- l7 mm long. Styleless than 2 nm long l0 Florversrvithpctals l0-l7mrnlong. l;ruit l0-20 mm high, 20-30mrn rvide, spincs,l-6(8) mrn Iong. [Coastal areas of WA ftomBroome to Carnarvon. Leatletpairs7-I0]...... 8. T.occidentalis 10:Flowers with petals6-7 mm long.Fruit 7- 10mm h igh,12- 15 mm wide, spines0.5-4 mm long.lSaline soils on edgcsofsalt pans,central Australia.Lcafietpairs6-71.... 10. T. sp.saline flats 8: Ffuitssometimes tubcrculate and often with a pairof medianand basalspincs, but lackingspines distributed all over:cocci bilaterallysynntetrical ll FJowcrsr.vith pctals more than l5 nm long.[Stigma shorterthan stylc] 12 Stvlc4 nrmIong or more: pctals l5-35 mm long.Leaveswith8-l0pairs ofleallets:leaflets less tllan 2.5 mm \!ide. lKununurraand Victoria Rivcrarea.Lcaflet pairs 8- 101.. ll. T. ranunculiflorus l2: Styleless than,1 mn] long;petals l5-18 mm long Leavcswitl'r 5-8pairs of)eaflcts: leallets 3-5 mm rYide l3 Fruit pubescent$'ith relativelydense short hairs overtopped b1,sparser long strigoschairs, median pair of spinesprcsent ol (]ilcs abscnt.ITanami in NT to RudallRiver. andHamerslcy rcgton inWA. Leafletpairs6-8]...... l'1. T. sp long-styled eichlerianus 13:Fruit subglabrous.vcly sparselystrigose; median pair ofspines alwayspresent. ICoastal sands oftropical Australia Leafletpairs5-71.... 12. T. cistoides ll: Florvcrsrvith pctals less than l5 mrnlong 14 l"ruits?-l3rnmhigh,I 0-20 mm w ide,denscly pubescent all ovcr' lntrastaminal glands forning aring. lStigma 1 .J-2.5 mm long.longcr thanst.vlc. Drier arcas ofall statcsexcept Victoria and Tasnania Leafletpairs6-l0l ...... t3 T. eichlerianus 14:Irruits4 7.5rnmhigh. 5-13 rnmwidc,subglabroustomodcrately pubesccntonl,v. Intrastarninal glands 5, distinct[7] le,"t'e.!/r'r.icomplex] 15 Cocciwith distillctdivergent, nedian spincs 3-8 mm long l6 Style0-0.i mm long,shorterthan stigma. lsouthern Australia in wastcplaces, also about Broonle, WA. Lcafletpairs 4-7I ...... l5a. T. tcrrestris (short style) l6: Style0.6- i.4 ormlong, longer than or equalto stignla.[N orthern Auslraliain wasteplaces. Leallet pairs 4-71 ...... 15b.T. tcrrestris (long style) -!
luesatdcJc sta^\ol' o11 siur,naql uee,$laqecJc erll ul puno.}eq ol oJUeJsaqnd esucp a ql ztt'snta1tl,l1o1d 1uro.g 1r qsrniu rlsr p qcrq,r scrceclseq 11o c rlslralJeluqf, reqlout? ,(u1d"^rpeseqlpuuad,(loloqeqluotrresoJdosluoleslrnr-11oraqornuV.(909c2./r7qrr;r.6.0)sseur)iJoc euos dole,\iposlD )^uou .tds srral.it,r/d / lEuorstJJotuql pttou eq plnoqs1r q8noqtlu,sertcds oql 1o sruelsilroc rrlstJrllulprlc.fta^ cql s,(eldsrpuorlrtlloc e,Jl:.ct1l.uttnalpt adi./aL/t ua |luaututd-)
'(t-Z).Jeal ern)puruuo s.rrcd tepcallo Jiqunu l.^\ol cql,{q puu(;uol uturt-z) sel,{tsreuoqs s1t iq'siurrr aqtuc:r^!eq e.uecseqnd csuep qlt,rr 's sttn4 slr,{q ore eql uo sseur ls ).roc3o 1ucrudole,up er11ig sttt,>1dt(1o1tl 1, uro.r.lsJeJ-Jrpr?r.or.)g,/s J trqEq,{qqn.rqs sy Kq snta1d,l1o1tl7 ldacxa sercedss- f(ll.tJ ).tlto Ilp ruol-Jpaqstniurlsrg sa-ruoafELitqstnEutlstg
'?qrp,p):(,wr.)./rq sJpaesnlaldiltlr(tJJoarnr\^u-HJuE.t ^iL++rtfl,:K;ii';:;r:::;:r':; '9861 3nV '.snoqpuo.r 0e encse1oJatrsoddo :VI.IVUJS6V NU.:lJSgl| panLuDxasuauutacl"^
'(A\SN'i,9tz8 t t0 H.l.U:ld:.rsr :Z8S99ZB Ny J :olotl ) 6g9zantt1.t.!tlj l 'LL6l ngw 'erlprtr^nv ,l.Ilrd ,ocnrg I Urotr^e,\\ 1uuor1e51a6uuy ,(a1staure11 lr\ .lo asuqoqt tV ..r.r./fJ
'p'llJ rLtou(9661)l/l louacJKl'1o,tur :/l pjDDtog eprrlcpv t,re)JBg.hJ.Uxi)lJlrlf,r:ll lr?r.r./rg ns snp(ltt.l.
.Z ,1o,r.A'JellEB I/{ U xeJclqor-:l H snsolaqnssnlnqrlJ
Drl'rrsnvur'se^A urre^ru lr'pnl olssorcu uor6e,rr;;;';lx]",1Ji, i,,,".::li:rX:]:u#,;i:;'J snq-1. ,{clraqrur) uteqlnoseqt ur IeerJ sruesuqJ eql tou puu 11111;(o;1o quou tsnl )aolJ sPtulsr.rqJeql sluoserdi.raf,uaJJoold lrqqpU eql uoIeolf spLutslIlJJorllJou selrtu / l ulol.l uortcJlloc pelrJeq.Ls^noeuorc sr (266 1) eJuerMc.l ,{q uorSel ,iaFrqu r) eqtro3 sercecls srqlj o p.rof,a.li ql r.r,r(r^,/
'(l-9)Juel elnlpu Erro slud tageollo iiqLunulaqirq eq1,{c1 ,s8urM puD(3uol Lurug 9-t) sal,{tsr.8uol s1r.,(q iqt ure.,nltqsnorqulF ssel ro rroru sllllr.lslr ,{q ,stuals .7 .1rquq,{qqn.rqs eqtuo sseurlJoJ-,tolucLudole^cp-Jo )oel aqtKq "^n,^otaqns l ruo.rl s-(r.1jrp sz.ra7rlr7u7rl slt .satny:af lq snsahqi^ 7 ldaoxe sercods:^nftlrlJ Je\lo 11uruo.r3 pcqsrn6urlsrq BurL1s1n\u11s167
(966 rilrtlg .( l ) cllssed.{13o uor1u1rc.ro3 :a/,fZ gggt ) OgZ: t lu.rlsnV.ljl.q]ucg sn.rrltl,{1u1rl snlnqrrJ.l
tptpuH sulsaual uorlJas ur papnlJurlou sorrrdsuurlr.rlsny rrru,tpug
srpdteUce,l 'p1f [4-t ryurlerlsnylerlueJlq8rqruuS-tsrrnrg 3uo1ururE1-9 ,f;uo1 s1e1r6cru8rls o1 1rnb. Jo utlql ro8uol ururg1-1.9c1,{15:11 '' sled 1agra.1 'uere [4-9 s^SutJdSoJrlV ot qFnoltpsuou cedspolrn.g rei,rel qlr,n 'erlprtsnv .Euo1 uleqlnos,illsol,{l116rq uuI S /-c sltn.r3 ururl-9.7 ,3uo1 s1u1c.1uurSrls o1 lenbe Jo ueql Jeuoqs tuLu1.9-7 g e1;eg11 3uo1Luru g uuqtssel cserp ro scutdsuerpJru SurlJlll Jeqlre rJtoJ :q I
.rillutl snltlqt.tJ ut:i sitou lur ,{.rj pr.rt V l{ U NuytsiaVol. 12,No L (1998)
Notes.The name Z stberosasH. EichlerexR.M. Barker,publishedbythe author(Barker 1996), was invalid sincethe typewas inadvetentlyomittcd. The nameis validatedhere by thc citationof thetype with referenceto the previouslypublished description and diagnosis (Article 32.3, International Code of BotanicalNomenclaturc, Greuter 1 994). A swell asthe description and Latin diagnosis, all otheldetails ofdistribution,ecology, synonymy and specimens cxamined are to be foundin theearlier publication.
Dispenalof Z suberosu.gand itsclose relative, T. pla4tpterus,has not becndocumented but the followingobservations have been made. Ithasbeen reported that the larger fruit of L sl6erosz.sdoes notbreakaseasilyintoitsfive2-wingedcomponentsasdoesthatofT.platypterus'fhiscommentwas "fruits provided by Mrs P. Glennonwho notedon her collection(C'lennttn 648) of T plaryPle,'r.t,that " separateeasily when ripe unlikeinland wood onesl?7. suberosuslwhichare large and adhere Mrs Glennonfurther credits Jack Paine, the lastgovernment nurseryman lin Karrathal,as noting that the "germ inlandvariety wh ich ispre sumably attrib utable to T.suberosuJ as above inatesbest ifno attempt ismade to separatecarpels". My own field observationsdo notsupport th is While Z suberosasfiuits maybe tardicr to breakup thanthose of7'. pl.ttlPterus,sinae at the green stage they sbow no inclination to sptitapart and this continues even when tlte fiuits are dried, they do breakup quiteeasily (on touch) 'fhe providedthe fiuit isripe and yellorvish in co lour. dispersalunit in both species is thc 2-winged fruitlet.
3. Tribulus hirsutus Benth.,Fl. Austral.l:289(1863). Typecitation: Nich[k]ol Bay, F Gregory. Type.Nickol Bay [WestcmAustralia], -t. dal. I I 8611,P. Wqbolt \'n. (lecto,heredesignated:MEL 79405); Nickol Bay fwestern Australial, Grcgory's Expedition,s.dal , Herb Mueller s n , (iioleclo. K-herb Hooker).
Tribullt.taldtus Ltuct. non Delilc: F. Muell., Trans. Bot Soc Edinburgh7:487(1863)
Distribution. Foundin the Pilbararegion of WesternAustralia through to the RudallRiver area and -ferritory. Great Sandy Desertto southernNorthern
Distinguishingfeat rer. In freshmaterial the fruits aredceply 5-lobed,sofi{extured and without any spines. Other species,such as T.forre,stii, T. nacrocarpus and someofthe 7 lel,'e.t1rl.!group could be describcdas 5lobed but they all havea muchmore leatherytexture and have median spincs. At maturity,in driedspecimens, th e w ingsofthe fruit areusually much larger in I /rrrsrlrr thanany other speciesin Australia,although there are a few specimenswhere the wings are similar in sizeto thoseof T.platypterus and T.subero.sus. However these latter species arc shrubs,while I lr.sll/llsis prostrate
T,pi./ication.The MEL sheetindicates that the collectorof this specieswas PembertonWalcott, a volunteernlembcr ofFrancis Gregory's I 861expedition to the Hamersley region (Birman 1979). The sheet in K docsnot indicatethe collectorbut merelythat it wasthe result ofGregory's Expedition. Mueller "T. hasannotated the M EL sheetas alatus?"and just belowth is Benthamhas annotated it asTribulus ftrisrlrs. Both collectionsarc scrappy but thcreis morematerialon the MEL sheetand so it hasbeen selectedas the lectotype.
"a Bentham'serroneous protologue description ofthe speciesas shruballied to T. platypterus"was presumablybased on the rescmblanceofthe fiuit of 7. /?Lrr.rtr.rsin the dried stateto that ofthe shrubby speciesZ platypterusofthe sameregion. Therearc no noteson habitwith the specimenand it is impossibleto determinethat it isprostrate liom thescrappy fragments which makc up thetype sheets
No1e,r.'lhisisavariablespccieswithrespecttoleafletsizeandpubescence.Thefruitsofthisspecies aremarkedlv different in their lreshstate to thoseencountered on theherbarium shcct or in thedried 'lecrpadaqljoxeduaqlorueqlpucs8eloq,;oqro"ort",lo"q,,J:frtffit"'rH;3;1il,1i#;:::#
.,o lrn.Ueql se)ro,retuur-J a(Ues eql uo pesuqoq olreaddeslrn.rJ aqt,rolaq pelEls sv snqtuo)DppD.J, roj ctlsou8erpsr qcrq,rrcdeqs lrn{ eql sr 1r sercedssl7ngr.i; lsoul rllr,rrsy .satryoalButtlstn4uqst6l
's8ur,,'r oq1urqlr,,rtueppr q ,(;lue.reddee:u sarreds srql ur qtrq,* scurds eql ot ecuere;alul ,ruorllroaurds Jo.I 'DqluozD ,to1rl1, e?JC puEelncsqo lo ueasunloJ IeetC oror3seruoc lcqlrde cgr ccclsey1 -tBolctwtlg
Bur.ra,trogpue,{iolocgereqEuolaqosle,{aulascquosqrgeq,r"r+r"",::t:JJll;::::tjfi""#j""r*? ,Deru lomz?uo16eql uror3euo puEl)llsrp sjouoa.I eql ruo{ euo 1eu8u111unoy1eql uro{ euo .uDul,,|\eN 'su,roC '€rlDllsnV Jo qlnos oollnB tuol-+euo uJelsartAur suorloelloo.rno1 ,{q ilur) u\\ou, uorJnEttste
'(HJugd) uorttpadxqBntuDale .cILue3 lll uasao'a66l deg elFegpesec uosqrD'Ieuoz oLueJJo N 'urnqxA :(l .Z66l.deS.uescC .oBurU {.LUd) 6uotTrpadxgSutwna/e uas-ae uosqrO FunoA:(HJUgd) ,e^rcset 68,tastuI S p uDutdDLt)v'l66l.deg 7,dtuer3o51ru1 g elrstulnq elnluNuesecuosqrC ,eelg ,{pn15 erog a;6e'g :VI'IVuJSnV Nug 1ga,t6 sndtocotcDw 1o1 sarltutffbqlrnpaurur)xa suawrcaclg
(HJuga)stfisra,rur!ft n'I L6t des 'uounqqsv ,uorlEls .L!.s- :(HJUSd) 0n9tpq)tfn V V'BL6I tco € su,\\ocooltn8 :(HJ_UAd) otu17.J y ndnruq g g'9961 'lculsrp .r1?1 lpeoU ,{ln[ r]rouoa.l:(HJUEd) 6/6I / ?lttipDLl)f,,,961 tr6 uotpllrreC uo 1eu8e1,11uno11oro{ rul 0l :VI.lVUlSnV NUE 1gatr4 satn\oa!Joct tl,{| t71tl|paurwora suawnad,g
'rcoolpc8ur,n-7 o,tg olur Surlercossrp (1quqo;d 'seurds 's.tr?rJ ,ra8uol ,JJllorls lDurclxe?urIJ€l lsnqo:crou pults.ueq leuos ,luaq,eldurrsjo crnlxrru 'lecrped e qtr,\\.rc^ollu luecseqnd,{lesuap cql;o xadeeq1 ol ucql puEsrule eql.}o q?eaJo spue aq} ol uotlccfojd xeduruorj ,epr,'!\ ,qFr -g ,uorloe[o.Id IecruocJo sBur,r,(qpaBu r,,n- g rur-uE7-g 1 q ruur 7 1 1 1eJuoo leotdeqlt,,tl pedeqs-.tu1s irz.i-7 3uo1 uru g -S.0€ru8rls !rnJI ur Suol utru €ul8rlspue el,{ls :pOllcJ-9 'luecsaqnd l t Z-g I a1rq,n,{1esuep ,{./l,,1(, u^\ouluns,/a,nn1J.,{11rule1xesnol;r,t1nq,u,{\oulun,{llsoruqr./rs '8uoyruu 7l -9'6Taa rpad Bwlmtg epr,rruurg.t-t,Fuo1u[u11-9g,,{11arxcqu:osuep,,{11urxepesno11r,r lJteurutnap elrq,nfles;eds.ro snorqulS'snotolocuof, + fluoqs ,cnbrlqo-popunor,alu1n1or1ed,{lpoqs 'os,(1,norleu 'c1e,ro saurrleuros ro cqdrllasloU€al :alorled BuoI Lrlu l I -, e^oqEstegual3o srred (9)9_1(g) qlr,\\'olEulellp .Buo1 .g.1r{ fllDnsn3loru'Jo s.rrDd Ienbaun ur sz,rnaT urr 67 1see11u s ulals ele4soJd
(6896st orl :99 g6 :o7o4).u,r,rorq-a8uero .gg BNVJ l 9Ll. 111X96 strnrJ:qJeq 1u.re ueqde elerlsord 99 a8./oaCS.f 't96 f gZ 'ur .pe ., I tnl IeJlsnv urotsey'A atseuoHuotlcts lo,re6uo16 3o qlnosIurl S[] satru ZZ r.r,?d,{J
'su1ll.ta1ur runruurds snqrrud sllnlrlsep snqrttnl lo snqrluecseqndsnqtsuap uelllp pesetuxotcl ctd,tocotct:nl 7 e,rousetceclg
'dorl .htr d.r'rcllug u snqluBJalrpBsnlnqlrl .t
'uorlr:uuo.l lsitrJJealaql ruo.r1pa8ur,,n_9 (1tee1c etu sarccds renlll escql acurs 'uorqspJ .J ,snd)DcotJDa.,L luariJJrpu ut pa,\llep^luruuaolsorulu sr 1r 1nq.vr.ra7d,! olcl l puesnsotagn"^ ur se1c11rn.rj pa8ur,n-o,rt eqt sl lrun lesJadsrpaq1 laprnl3qcueJo s8ur,r o,lrt cqt ruro3o11r;ds ueql pue reqloqcuc uo ur esdellol slle^\ luialEl eqt.Sur (rp ug ulsattof tuln4 eqljoedeqs eql croul iu rlqucseJ 'uorlDrolocpo.rJou,{\oJqqtrMpuppelpUuralrnbclesledtcca,lUoquoslle^\eqt,lnoFur,i.Iparogrg.elelsl
tnl q t uo sotoupLrE .\.1 ltut v r. ug hl )t Nult\ir V{,1.12. Nn | (l()()8)
No/es.Trihulus qlelacr.rirtfur.ris describcdfrom linited nratcrial. l he author \\ ould apprcciatcfurthcr collectionsso tltat its variabilityand relationshipscan be propcrll,investigated and its description completed.
It has sometimesbeen confusedwillt 7'.hir.vutus in the Pastalthough Lawrence (on PERTH " specinrens) suggestcd tlrat it was intermediatc betrveen Tt'lb ri1ls sp. B I Z ?suherotusl andT hirsutus". Z /rlrsirlrshas vcry inflatcdcocci in itsfi eshstatc and the[e is no undcrlying hardened fiamework wilhin the walls ofthcsc ctrcci,as there is in T. odelacanths
llased on the liuit alonc,Z udclacdnlhrcseems to bc closelyrelated to T ndcrocdrpus and T. Lt\ttocdrpr6. I he spines usually present bctween the w ingsof T nacrocurp r.r appcarto havcbecome longerand amalgamatcd with thc wing. A nothcrrva,v ofv iervingthe fru it is asa Z astrocqrpu.t liuitwith wings from thc apexto thc endofcach ofthe fivc spines,these rvings then continuing from the spines to the pedicelapex and base ofthe fiuit.
Thepresence ofspines within the rving is rlot unknown since m aterialof Trihulu.rt:rhl.rlr.! tiom South Africa, secnin AD, hasnunrcrous spines projecting fiom thc endsof itsu ings
T. nacrocorpusrnd T. atleltcanlhttscan be distinguishedby thc subglabrousfruits with only a sparseappressed pu bescencc and external spines betwscn the w ingslbr Z naclocarpascom pared w ith thc pubescentfruits and interna) spines ofthe ncrvspecies. There may alsobe a distinctdiffcrcncc ill the lengthsofthc styleand stignta bclrvcen the two taxa,with thoseof T ntucroctu'pus being longer, but thercis insufllcicntflo\\'crirtg l'uaterial of tl'lenew taxon 1() be sureabout this Thenew taxon may atsohave a nroresoutherly and inlanddistribution than T. ntocrocttrpus
Thc G ibsonDesert collcctions havc the spines more or lessamalgamated with the\\'ing but the fruits arenot asdensely pubesccnt as thosc ofcollections fiom furthersouth Theyalso appear to havcthe shorterstylcs and stignas associatcd rvith the ncw taxon. but thc lengthsalso apply to T mdctocarpu\ rvithrvhich thcse collections also havc afflnities.
5. Tribulus rnacrocarpusF.Muell.ex Benth., |1. Austral. i:289(1863). 71l/e cll.r/lor" N. Australia In the intcrior.trorn Nichol llay- f'. Gregory.7.1,pe:Nickol Bay IWesternAustralia], s ddt , Anon ItJnknorvn.Brought in ll ornirrierior by Partyl(probable httlo: MFL19123)
Tribulus .tp. F. M ucll.,Trans. Bot. Soc. Udinburgh 7: 487( l 863)
Specinensctantine,J WES'l'ERN At-lSTRAt,lA: 125kmSoftsarradalc,Hj Eichler2S6l IB(CANB, I,ERTII);c. 0.5knr N ofNo. 18 Bore,Hamelin Station, /t. Eich1er23646 (CANB);Cape Range, Greall' l0(PERTH). NORTI IERNI IlRRITORY:South Lake Hopkins,1'. K. /.al:7998(CANB,DNA).
Distributioncrntlecolo.{|,. Found in dry arcasof WestcrnAustralia,liom Capc Range to Geraldtonand inlandto southernNorthern Tcn ilory,and north ofOodnadatta in SouthAtlstralia Foundin redsand, usuallyin rlulga rvith,!r'rslir/c,Plctlrtrchtte or fli.)/1.-1,solnetimcs the sandover linrcstone
Phcnolcgr- Florvcringnreclonr in antl)- April throughto Scptcmber,occasionally also otttside tltese onths 'pcuorlsanb eqlqBrru qrrq,r uorlerrlqndgo eceldaqt,{luosrtr os pue lqlrllueJ'loBurrua(auruseqlureueueurusaqlqsrlqndprprcll.nr{.sueprDL) .{\a) ol uorlnqrJlsrpeq1,{q letu ueaqo^ur{ ol uto.splr.to,\\ suotllpuoc eseql os prrr ,.(llulouc8stsruDtoq ol elqrssecf,escrJeJqrl Litr^\suorlnlrlsurI€ltuEloqol.(Ur3lo,a8uuqcxa,c1esqBnorql)JclleurpeluudJo uotlnqt-rlsrp,.,{qpalco.liesruorlucrlqndluq}solEts(t66l7r,,r.telnarD)clntelJuerroNlecrul?loU.locpoJ lsuorleuraluleLlUo6ZelJrUV'Je^at\oll.(g/6lTtrirllrqJlnqJocs),{uururespuu^\e)olluudeltxquusu pelnqulsr p .rc,railuo trlsa.uof1_1o uorldursep cql Surureluoc redud a q1 puu peqsrlqnd rc,reu fltue.rudde .peqsrlqnd se,,i\prolcu ecuorcSuieqlnos Jo €nssttplntruud oq1 lslL' sD,,|\irl.!r./1ol .l eloq,r Jeelc ,{lejrtuttou st ll ir7.o-r.ro/Tsuueqlozru8ocajlouprpeqospuEstrnl.}ppqrsrLllJo.reqlrlutnq.glglut ,{eru3puulse.r:og ,{q suolllcllotrerpNcJo uuo.} eql ut urrLlol.lqelru,lu Ftruamoga^eq prp rellanr{ '(lurluor luuolr;ru eql ol lueLualulssrq elrdsc6 1t.r Jo/ J ad,\olceleq] sepelJeles ua.q stq uosearslLll -sndtootucoa Jo lr .to7 J Jo lrql q1l^\uosuuduoc ur 1rn{ oqlJoedrqs oql uo selouraq.unJ suq ueurcids souofeql lnq sleaqsqloq patploLlutlspq.rcllanhl lucserd slrnrj cJnleululr iluo suqlcuJlf,ru tsJJlol p cql sDereq,rslln{ cJnlDut.JoJequnu spqpur snotdocaloLtl aql sr lcueluu sauol egJ' uouDor/jdiJ
'rJJoc podolo^ep.qt Jo qcueuo scurds;us.rop uurparuJorucle putl (lnurcd ugo 11eqs cql ol.rulruls) ecupns pc1Jeu,(leltulncrlel.qtooLtrs p qtr,r (cjole^ep sla ecIIDlouuall,\\ldorxa)r1?1n3crpuepeqol-q'JEInJrro.JE.r7'.sa.uofJutstt:l.satryDa!Sutqvn'w1s-rg
requaldogpue ludy u:rc,n1eqiuue,nolg iBoJoua4,1
.pulllqnrqs seelupaqlusrp uro{ seurteuIostnq ulqlr^\uror-} ,illensn ,fDIc 'fu1c .pues ,(puusprr puu,(e1cu,,irorq-par s^noe;ulIu. lu04 pcproce;sr11ecre ieg1.ruq5 cqt ur,{1;ulnlyed 'utlDrlsnV pclu ur.lsa/A lo uo^leuluJ eql ol paur+uot st tltsa t)J!1 ,r5o1ocapLlD Lrattn.lj,ltsre
'(lol6L'l:ll\1 u[s) u s 'Zg8 '[cr lt )a.4I | 1er1sny ujcrse16l r.^ rUeuiorsBD :( g9g61 :pelr:uirsepereq'(\)aD.u.,^euo[, 'ggg 'fer1u.rlsny 1E tr1 1 u.re1so16]ra,try cu.,{ocsug :arlr,; paurelqosuaur:rcds Burlrn:1 ,{;uo se .s^cuo1 'J!\,{q 1a,{ lnq eJueq}uo.!.}luas oslc:lsaJJol I uo11'ra,rrg-eu,{ocsugaqlJEoN.r./o.r,/r,r1rali(l(gggl)r/t:rZ 'IqleJlueJ ,,{ucuueg log urlall3n t{ .{qpeqs rlqnd oslu 1nq pue,trey o1 sj oord e8ud se .{1uo pelnqr.rlsrp (ltuarudde'(StUt ,.llent ,\oM (scues,\\.N)l prorc1ecuar.s ueqtnos .J rllsaJroJsnlnqrrJ.g
'(eEeJersnoq)lPtS) DisnoLftrDJS'(euece,r.1a11) uolttn.ly,(e1lex\;tltu]exv) saptouDTtht|n;'1 E atxut ]ueJeltlp lJ[J u r erpro sercedsrrcqUo lse.I aql uro.rgarua8'rc^r p ^\oqsqJrq,r sluuld uoddnsol u^\oul sl c8uegedcS eql tnq EeJueql tuo.Uuor].o11oc euo aqt ,{1uo sr eteq} acurs u,rou)un st ecuecr3ru8rscturouoxlll ,{upJosrslqlJeqleqlA.serJeds.srqlurpunoJ,{JlDnsnsrueqleuirlspuu e1,{1s;e6uolepuu s.rc.nog:a6re1 qrnusuq(ql,rloarg)eiueladu3eqlurorjueurrcodsel6utsy.,rato,l,
eq Pcq leqlrciacql ssnlou prptnq (t9g l lillenl,.{)uoltrpedxa ,(ro6e.rg eq} uo O"rr",,o. ,u" *,.ld".:ltj,l Jo lunorJDup ur sercadssrq1 pauorluaur .le11en11 an301010Jd eql su.ruc,{eurrs aql u1 rucquag r(quees ucaqsuq put: . rellcnt\j/iq c Lucu eqt qtr,'lr pelelouutl sr.llllJele Lu.q1 ,i1.ru.1,,cqt iq urlq8norq sr,u 1r er ur s .(ro8erg Lutqse,n 1l 1uq1{le1rlun sr lr pacpur su,r.rolcalloocql teqlolucrpur ot sloqu;eq1 uo iurqtou sr e:aq1 .1q .uotlt:ttr/idI1 sFur,r,rror'te u elInb qlll\ rJJoJ paturoossrp g 11o ,{1ueu slsrsrroJ !r{ ur cdh eq,1,
'snor,lqos,{u,n1u lou erucseql q8noqtlu'saurds 3urltrrod pre,ru,,vrop uetpeu o^\1fllunsn cre eleql s6ul^\eql uea,\\lcg 'sFut,rr .satryoaf leurpnlrFuol,ror.leu S qlr,\\sllrq lluus elqureselserJcds srql ur s1tnr3 Euttls.rnEutls^t61
..rrvrfl rntnq).|.|.lo sito tur (rt trut V N I Nu\l\ia Vfl. 12.N,, | (loc)R)
7. Tribulus astrocarpusF. Muell., FragmentaPhytographiae Australiae l2: 4 (l 882). TyPecitation; Propeflumen Gascoyne River; J. Forest. Tlpe. MenilyalyaRiver nofth ofshark Bay [WesternAustralia], 1882,J. Forrestrr. n. (1ecto,here designated: M EL 7939 5 ); GascoyneRiver IWestern Australia], I 882, .J.f'orrest ,s.n.(isolecto; K).
ReDresentatiye.\pecimensexamined. WESTERN AUSTRALIA: GreatNorthemHwy, 45.6km S of Kumarina,2TAug. 1995,R. M. Bqrker1067 (AD);No I Well,Canning Stock Route, L.A. Crqven 5041 (CANB, PERTII)t WarbunonCon'rmunity, between shop and river bcd, F.,1.Zlcfr 76 (CANB, NSW, PERTH,PRts), NORTHERNTERRITORY:PlentyHighrvay,22kmNEofstuartHighway, Kl. l/l/'mn4641(DNA,NSW). SOUTII AUSTRALIA:-l'imberCampBore, Hamilton Station, 100 km N ofOodnadalta,F. Badnan309 (AD, BRI,C, CANB, MEL, NSW). QUEENSLAND:72 km S of Winton,Mar.191l,|'. Knowless.r. (BRI).
Distribution atld ecoloS'. Occursin WesternAustnlia from Carnarvonto Warburton,cxtcnding into southernNofthern ferritory,south-west Queensland and nodhem South Australia, invariably within mulgawoodland. Soils vary from redsand to pebblybrown clay.
Phenology.Flowering has been recordcd in mostmonths ofthe year but predominantlyMarch to May and Augustto September.
Distinguish ing fealirre.r. The star-shapcdffttits ofthis spcciesare very distinctive
"Tribulus T)piJicqtion. Both shcetshave bccn annotatedby Mueller as astrocarpus",although the "asterocarpus" "c" lectotypeis annotatedas with the crossedout. The MEL sheetconsists ofthree fiagmentsand contains florvers and young and mature fruits. In contrast,the sheet in K hason ly a single fragmentrvith one matutefiuit present. Both specimenswere seenas photographsin the Eichler nanuscnpts.
Arotes.Tl] ere secm to betrvo flower sizes amongst the m aterial studied. l'he f'ew specimcns with larger florvers,i.c. petals up to l0 mrnlong (e.g. Kt owless. n., Crcnen5044), were more robustplants and tended to be floweringin the carliermonths, but theredocs not appearto be anypattern to theirdistribution sincethey came from all statesexcept South Australia. These larger flowers tend to haveat lcast2 or 3 stamenswith longcranthers; thcse are possibly staminodal. Also associatedwith the largerflower size,as might bc expected,was a longerstyle.
Section 7'erlestfr:rHadidi, Taeckholmia9: 60 (1978)
Thetaxononly ofthis wholcgroup is very unsatisfactory and much in needof world-widercvision Many specimcnsexist rvhich possess characteristics of morethan one spcciesand tllere is an urgent nccd,particularty in viervofefforts to find a biologicalcontrolfor T. lerrestris,ofan understanding of thebiology ofthe wholegroup before a satisfactoryclassification can be achieved. It is likclythat some ofthesespecies are much norc closelyrelatcd than the present taxonomy suggests, and it is alsolikely thatbarriers to crossesbct\I'een spccies are not wclldeveloped. Even some bas ic eXperimenta] wOrk to establishwhcther such ctosses are possible and thc characteristicsofany progenymay helpto place thctaxonomy on a firmerfootittg. '(tUA stlDtuapt)).)J u! u s tJuruDg) puglsw eno ur ?urlueluurC eqt urol.J,rEs eq leql r:/,//rif{J -lo ueluroedsfguo eqt 11eco1 FursooqrXq rcqlrnl uJ^e uorl€ntrs eql pesnluoo,{lqrssod(668 t) {c|ug (g991rellenl,{) r1is,q J sp u,rorg puDDoclpy' fq pa1ce11oc 'tuEeul s1uu1dpeleerl ,{lurupeJ eq acurs le11en11luqt uorlrpcdxg ,{lo33JO eqlJo lunocce eqtueeq e^l?Ll 'JeAo,r\oH '(6 ,{pur}r ereqlpiuouueul lou stsllouapl)JoJ ?xuls 9gl lellent!) rrorlcelloce3€qquO eqUo 'srql tunof,f,esrql ur tou su,\\tr ucql pelseSSnslsrlJ oqM JellenrlJ ss,\\ tt JI uorlrpedxle;Dqqu0 aql.lo s1ue1deq11o lunoccp srq ur uolsrcepslLll Surleur roj alqrsuodse.tse,n orl,n aq se,r\ll leql pcrurulo(96g l alEJ d, rallint{ ur)Jollenhl udorg u.qo1 ,(quees suerurceds eqt atucol o1clqeun se,m eq ecurs,(1uo srsouBErp oqt uo poscqf llue.rudderrls,{r7 7;o ru,{uou,{se sur i 14uap no J p:4el.Jl ( t9g I ) tueWueg
'setcads srqlur 01,(1seql ueqlre8uol eq ol rur34seq1ro3 lunsn sr 1r stlaJeq,,r'(urut I ,) eLu8rlseql uuql -ra8uolsl (ruur9 l r) a1,{1seq1 lnq'elD/uap)Jt) J.+o slrn{ Surdola,roppue (3uo1 ruru Z l r slI]lsd)ezr s Ja,roUeqt seqetlells nV ujetse/[ ur (9tg Sl./orrf) elppeJJegulo;; ueurrcedsy
'saplolslc 'peqluplq 'seurds;o J Jepunsalou aas stlDiuapt))oJ ul cq,{eurqcrq,rJo eruos 'l€seq 'seurdsjo uorlnqr4srprc^o llEeql qlr^\ peJpdluoc euopuE uurpeLu auo srred7 (1uosuq saprotsrr ,; 'luesard 'puDlur oreslrnr3 ueqr\\ qsrn8urtsrp01'elqtssodtrlr touJ I llncUJrpele o,,!\lesaqt slr n4lnoqtr,\\ puu 'septotstJ :aqung 8ur:rnrco oslu q8noqlle J o1,(1r1eco1 lelscoc Jt?lrrurs u setdnx.ostlDJuapno J
'xt"tlsftl J Japur se]oueql ur passnrsrppuu e^oqu petsrl eta xitlsfi.l 7 qceotdde qat\M sllDJudptJJoT 3o sueurredse ruoS raqlo qoeJtuo.t3 lcurlslp etu (eql srTrluapt Jco J utseclaq.r Sutrelelduocururoj olrcaddenttst7 1ur spue18,(;e1ceueq1 qt6ue1re1urrsjo eru8rls puu e1,{1s eq1fq (q puesJe,ttoL+ pup slrn{ Jell€usaqt rlrtr',{r1J ruo;3etquqsrn8urlsrp ,{11e nsn sr s.rlDruaptno J '^}toN
'5uo1ruur 3-1puu qt6lral ut l?nbJsselJo aJou elE eru8rls pue e1.{1s cq11nq (5uo1 uur 1 1-gI s1e1ed)e8;u 1 eru s-re,to1g (ri.i7s,{ry 7 u13uo1turu11 o1dnqlr,n 'l peredruocSuol lulu 8 ol dn)rris',{ry urpunoJ asoql ueql;egoqs eru seuldg 1rnr1eq1rero11usaurdsjo acuesard eq1,{q peqsrnBurlsr p fpsee srsr loqapl no J'tueseJdelu s1rn,r3ueq16 sa,tryrnlSut4smButlstg
'Jequetdes ot Iudv 3uua,no1g ttBolouaq4
'uo,\IlureJ ol euoor8 ulo{ EtleJlstlv uelse/AJo lseoclse,r €Lllol peuuuoJ trotlt(lltJslo
'(sleeqs Z'CIV)9t t tuDeLtD^ H ',tof 'SNVJ 'CV) '.uortuls suoJo S Lru{Z0ll setrLut9 :(HfUEd tr7yguowtg ? O 8uruur16jo 1,,1 [ru{ 9g]salrrrt St :VI'IVUJSnV NUAJSA16 xu1sI11c4 Lltoo,lddD awosBulaaqs pautwt:>xa suautrcadg '(sn'ata'Hrusa'A\sN'vNC 'gNVJ) SSI {rlz J'ql!urs}lod'eerE 6urduer q1ruggo4urorg ur>1 1uoo8elle eereSurvudrceu peor eprseg ., :('lAht'gNVJ) .Sl,rro{S S dr'f^\H lplspoJy'ANeqtFuolu uo^IuurEJjoAS ur)il :(,\\SN) ao.rlrtl .1 'ppou 'olppel.lpBJo 8 9V89sqorDl i^'8861eunl 8 uo^Jsur€J S rul g l I :(HJUgd) llgl taup.tDCy '1761ru1197'rrosdlueslurod:(H.LUld'1!\SN'SNVJ)Z09e7n1qag ) tg,(nor.tulsI€rrSolorg ouuEt{) 'pqor f,IulCrerdw?C le'rerdLupC :(palnqulsrp oq ol ydnp'qy) *IIra\Dg lll U oltxeu.q.eeq tuo{ I??q 'qceog eunp pupslsJg uo eqturrc) :VIIVUISnV NX:l ISA16 pautaDxasuawtcads a,vy4uasa-tday
(pZ66l) uosir/Acos sercads strll.]o uoqecgrd(1 eqljo uorssnrslpu ' .g rcg.adt1 ftyg1) 69 ddy 7 erlurlsnylerluca olur uorlr pedxg ntsulJg U srlntueprJJosnlnqrrl
sntn.fuJ uo s.tou pur I.I lerlr V rotrr8 ht lt Nu)tsiaVol. 12.No. I (1998)
9. Tribulus hystrix R. Br. in Sturt,Expedition into CentralAustfalia 2 App.69 (Jan.-Feb.1849). 'llpecitation:incollinisarenosis.Lat26',D Sturt 71'pc:Sand Il ills in Lat about26'' s dat.,(:qPtain Sturl109 (holo; BM tr.t., seenas photogrilph in Iiichlermanuscripts)
Re]lre.ectltqtivespecitnens exctnined. NORTI IERN TIIRRITORY: Camel Lake,Sirnpson Desert, D.F.Glbson 6l (DNA). SOLJTHAUSTRAI-lA: Priscilla Crcck. 30 km NW of FinnissCreek on C)odnadatta roitd, K.L. ll/ilson4601 & R.M. Btxker (AD. CANU, NSW); 5kmNof StuartCreek Ilomestead. F.1 Badnon1581(AD.HO, MEL). (BRI) QTJEENSLANt): c. 27 km SW of Bctoota, K P Ni c o l s on 29 l & P.E Nor'e/il
Distrihutiott.Found in northcrnSouth Australia, south eastern Northcrn I erritoryand southlvest past(Hnatiuk 1990)and is in the Queenslancl.It hasbeen recorded from westernAustralia in thc WAHERB Ccnsus,but ntost ofthe !natcrialidentified as such is probably 7i occilerlalis (btltsee notcs bclow).
DistinguishingJr:unrres.lfliuits arepresent, this species cirnllot bc confusedwith anyother 7i'lbrl?irt spcciesin Australia.fhe ir size (up to 30 rnmh ighand 5 0 mrn u'ide) far excccds that ofan1, other species, andthe length ofthc narrorvspincs ( l0- 17mm) isalso diagnostic. In theabsence offiuits, the large s ize ofthc flowersand Iength ofthc style(4-5 rnm), coupled rvith a knowndistribution, should be sufficient '1 to identify . hystt ix.
Typilicdtiol.Theholotype shcctconsists ofasinglebranch with aflowerandasinglefruit. Ithasbcen "-l'l annotatcdby Brown as ibulushystrix Br'"
"vaf. N.rter.M ueller annotirtcd a numbero f specimcnsof z l?lrl,'r.r as grandiflorus"irrc luding that of McDouallStuart's from Attack CIeek (MEL 794121)rvh ich rvas seen by Bentham,but tlle name has ncver beenpublished.
7. ,l,stro.can bc djstinguishedfi.ot.t'r T. occidentolisbythe muchlongcr style than stigma, by the largerflowers lnd tiuits and possiblyby thc intrastaminalg)and being a completering latherthan 5 dlstinctlobes. Distributior also oftcn helps in distinguishingthe trvo species, wilh L r.i,slrrxcoming 'l'. from central Australia and occidenl4/r.r from thc west coastof WesternAustralia bctweenBroomc andCarnarvon. HOlvever, (lrere irrC some spccimens from the wcst coast Of Western Australia and the interveningarea which appearto breakdorvn the distinctionsby approaching7. l7],r'//r-{in some charactersrather th an T. occidentalis,raising the possibilitythat the two speciesarc mereiyends of a clinc. Ihe interveningspccimens lcnd to havelarger florvers tlran is usuallor 7'.occidentalis (petals up to 2g mm long) andthe stylcsare longer than the stigmas,but not aslong as the 4-5 mm found in T. h).stri.\. Spincson the symon collcctionare lessthan 8 mm long, which is charactclisticof '1. occiden/c1lis.Thc cliscmay be a continuousring ratherthan distinct glands but this characteris not alrvaysrcliablcoreasy to intcrpret.Bentham treated z occidentLllkas avarietyof z l?*,\//'iland his conceptmay evcntuallyprove to reflecttlte relationshipbctter than the maintcnanceof thc two as spccies.However, complcte specinens irfc needed ofboth spcciesas we llas morcspecimens from the areabetrveen lheir prcsent knorvn distributions, beforc a decision can be made on their taxonomic status
10. Tribulus sP.salinc nats (P( L.rt.4530)
'lTribttlusocc rclenrulls (smallcr florvers and fruits) K.L. Wilson.Ielopea 5( 1): 28( 1992) -
'elnunun) ,pr.r ,.rntlncr.r6ylo .Luru_r :(Hl1gd'UNVJ) IOlgu1dy H :l.J,BL6l rrl J I I nunun) ldoq lulucruuadxg.,{a1;eqLLl;jo epi:rlrsocldo pmr ururu:Vl.1VU-LSn V NUgJSg,lA fautLtnrasu)utj)a(JS
.1J.iclra.r,\\D.l' - (266I ) l0 l a I' I 89 uorBrl,(alrequrr; y clss n 1 nq t.t.1
\>l:u:tt) u sDlldnl.\' :l '9931 'ftorrrre.l. 'laerJ lrrdy ur.rllroN s rucrt)trr1\:(ZLi6l .y-g141.uis1.u s .ra11arr71g .9Eg ilunuel ',ltotr.ualuoquoN ',re,t1e 1 uolcrrlraddq .rrl,U (068I caq)gg:(t)t nrt.'uezucgd.leN !tuer,1ry'1iugur.16ug - .u8erg,.Ilentr1 \'lpn!^ J)Dta tl ruturntDt uao.ttt^lltiY (S SS t ) St: t .1:(998 I ) eag:g Iunllorsrl,{ uepru C 'll1o, .lle .ll ,\\e) ?t ,iuulo8J o lautnols,lJlooH ur( pr!.r ,{lqrssod) n!\ C snrourlnrunuBJsnlnqrrl
'o.lcq pazlluLruoJueeq lou seqtr irl,,r\uor^ee.r leqloult sr puu,rro\D]stqlJo iltprlu^ aql uo etuosslsec '3u ,uortnq lqnop tlf,elloJelDnbeppur lceget ,{1arou ,(uurtr qBnoqt ue^a r.qstptcunls rp,{ 1opr,u e qJns e)cllles€ ol luJcElp€DelE pues pel u uol-lcpcu suMlr qinoqtlu.potsrlsuotlf,e llo. lerltocrlt ulol-] pe^oucJ,{lq€reprsuoc oslu sr Erle.rtsnV uJ.tseA\.Jo lseoJ tse.r eql uo.Uuollcellor Jcllug aqli1,ru1rur5 'llrqpues u uro;16urtuoo se peproooJsr lr puDsuollJellor aqlJo lsJ.reql orol-lpcloLLrej llcpraprsuo) sr uorlnqrJlsrps1r 1eq1 lclecxo slccdsar 11t Lrr letJeluru slql qlr,r saerip uotloellof,zsueupl\l eql
'(uoxrt tuLll.rapun selou ces) ^Dptolsro .ds J, to snuDtt;>1Lptapa1I1s--Buo1 J Jo sel,{lsra8uol puu s.ra,'!\ogrr8re l qtnor eq} qtr,r puno.} ucaqe,\€qsarf,JdssrrpJocsoqlollulrurrsqrnr1Surjucq(1q17 d()lu O-l e)suewnodsleuorsrcc6 s;:rlt.r,y
'sltntlxrrliri/JJJntD .ltruirltlJ.loute.rrslnt.l.lorlllnq.snLfil)aHJD J .sd.Itpaf Jo stll3uclttuSrts pue ellls pu? ozrste,{\oll Il€rLls erll scssassod uoxrl srql ;iutrls t'Lltlsj(l
'suleJ ol esuodsalur .,{1qlltunsl.rd,(r1r,1 o1 ,(rtnueJ 6ut)en\oll .t1opu)tld
'(89 .euolscrLrrl LZzsual.tnN)]ntqpues u uro.r1 .{lcraur Jo le^o ,(puusuto.r.1 trjo spJoc.JJeqlo etu .JJLll 'sued lros lnq tles-iosa8pa aqt uo lo slrosiurlcs ,{puusur punoJBoraq sp peplotel ,(llcnsfl scl!y'Aqlnos 'Pr ,\\cNu r lerllno uurllr ^\ lpllsnv ulclse^\puu ,{lolr e.l.uloql-loNulsJnf,JO ,iJolooa puo urtTnclttls.tg
'(BNVJ) .af,uonelS g9 LZ.:r.r1rl.rrl su^\oc,{rnqsrleS :SA,lVA\I IlnOS,^ASN '(HJU'Jd'VNC)0t9t.tD7 X,l .17'aloq{rou luljl lrqqeU s [ruM] selrurt I :(VNCI'UNya) 7111ytttuag ,\\olgzu0no y'NlrLr)lZ e l!^elrtu Z :(Hlugd 'VNC 'g 'CIV) .uorttts NVJ tZ I Zro Zl_| Z:/,r? ) df ui^Erl^\eNrAUOltUUg.t. NUAHIUON '(HrUSa'gNVC)OS g y 't961 1ya8.oag ,{lnf1'aiue1 q6auu,re3:(pelrlqutsrp oq ot.ldnp,CV)60Zl .tayDtJN U.9661 dcg 9,,(.,ug IstsDoJy'A N oro.Upcol uqqonf luro.lSuop ruI t'Z :VI.lVXlSnV NUg.l.S3A|pautwDx),^u) !t)a.lS
'JJUlln\ iuol trrrrry-c '1unbe lr'.JoPl)\ol;r:s.'ruds 6 ssalro erourqlt,r'rccor ipoo,m'luorsrqnd ileluriporu'luculetuuri(se S-! olurArrrterJossrp 'epr^\ -Z 'q8rq '3uo1 urlll SI I wrt 0l-L tin.t:l urur6.9 eLu8rls:te,\\ou ur iuol urur lr1,(1s:pellac-g 'luaosaqnd 1-g.9 ellq^\,{lesuaptJrr,lo Suolunu Z l sleq}up:Auol utru Z , slueuull}:Dur;rls ol lunbe,(lun}DLrr '3uo1 lu'OI suawus^ 9 spuD18lautwDlsorlr4slrd.s uErllJe8uo 1 uur1-9.r.snolqu[6,e]E^oqo..rollei sTnTa.TurFrcru eurlu,{q uo ldocxafllarxuqp snollr^,il.suip pue eruacsoqnd ldocxc,(11crxupu '3uo1 lucrdu.rol snorqulS urLuy r sTrrla53uo1 uru91-61larrpad8uqrm1,tq8r.rdn,iuo1ruurg-gptpatlButtatolg 'epr,\\ '.euol .,{l1urxepu .snoJoloJsrp rurugZ-E I ruruc 9-t',{llerxuqusno11r,r ,(lesuep orour snollr^ ,ctncl] '3nbr 'etDInlorled 'or1dr Iqoiseq ilyoqs 11a,{l^\o.ueusleUeil :alorled iuol Luurg- e^oclrsli)UeJIJo sJmcl 'cluurellu 1 /-9 qll,lr ,{leJurarou's.rrud 1r:nbcut-tut ta )e.I ?rroILurgEotdnsuels,g./rrysnolllA,elurlso.rd
.0 'Z \r,,r.ir,/'ru Jli I'i'. i:t t'Ur \ r.)t t\ ll r.\u)lsiavol. 12,No. I (1998)
29 Fcb.1972,D. Bedbroolcl0B (PERTH);Kununurra arca, I F. B1ac,tl0 (PERTll);bitumenroad verge, blackclays ncar OPRS, Jan. 1981, C. G-lover I -7 (PERTH);opposite Kuuununa airport, E.C. Glot er 56 (PERTH);KimbcrleyResearch Station, Ord SandyLoam, E Zarg/ield58 (PERTH);?9kmN ofKununurra atMocks Dairy on E sideof lvanhoeCrossing road adjacentto milking parlour,6 May 1993.A.A.Mitchell 3085(BROOME,NSW n.r,.,PERTH);Airportentmncc, KununurIa, RII/. Pttrdie3303(DNA);Kunununa, streetedges near swimm ing arca, 4 June1975, D.6. Slnon 10331(AD, CANB, K, PERTH).
Specimenexamined with o;//initiesto T. ranunculiflorus. WESTERN AUSTRALIA: Mackay Creek , 31 Joly1917 , Anon.s. r. (PERTH03776085).
Distributiotl.Apart from the typc,known only from the Kununurraregion of WesternAustralia.
DistinguishingJbahrres. This specieshas the large flowers of T. hystrix, but does not develop the extremelylarge spiny fruit ofthatspccicst instead it hasliu itsmore like those of I /e/re.r1,"lr,with only amedianandbasalpairofspines. Mature leaveshave 8- I 0 pairsof Ieaflets,with thcsclcaflets being narrowlye lliptic and no morethan 2.5 mm wide. Thestyle is 4-7 mm long,considerably longer than the stigma,and longer than any otherofthe sect.Dlbrlr,r speciesin Australia,apart from Ll,il,"rr (rvhich hasa stylelength of4-5 mm).
T))pificatian.Both syntypecollections have been labe lled as Tribulttsranunculiforus by Muellcr.but bothhave petals c. I 5 mm longand styles 4-5 mm long i.e. at the lower end ofthe range ofm easurements for thistaxon. LIowever,in the crucial measurementsof p€taland style Iength, the protologue gives a lengthof"7-l0lin."Il5-22.5mm]forthcpctalsand"3-4lineas"l6.75-9mmlforthestyle. Thesepetal measurementsplaca thc typeswith thetaxon rccognized here, but the stylelength is longerthan that recordedfor anyspecimens (see Table2). Bothtypeshave onlybccn seen as photographs in theEichler collectionofprotologues and need to bc checkedfurther befbre a decisionas to the applicationofthe name T. ranunculiflor?.rto this taxon is made.
Noles. Sincethe typc collectionof T.ranunculrflorus has petals c. l5 mm longand a stylelength of c. 4.5 mm it was initially thoughtthat T. rantutcul(lorttsmight provc to bc synonymouswith T. ci,\/oides.This was reinforccd by a furtherthree collections from Kununurra(Langfield,Purdie and Syaor, ref-erredto assmall-flowcred in Table2/ apparentlysharing these attributes. However, other collections (lp1i,, Black, Glover and MitchelD frorf. Kununurra have flowers with longer petals and styles,makingthem vcry distinct1'rom any other species in Australia.Acompa sonofthese entities isgiven in Table2 andfiom thatit canbe seen that all ofthe specimensseparated as T. rantrnculiflorus, despitetheir variation, can bedistinguished from l ci,rlolde-rbytheirstyle length,their leafletnumbcr, their feafletwidth andtheir fru it width. Fruitsof Z cr-r/olde.rare similar to thoseo f T.terrestris in shape but muchmore robust fruits of T.rununcuhflontsbcar aremarkable resemblance to thoseofI lerre.r/,'rs exceptfor the presenceofthe very long style. T. tetestris and T. ranunculiJlorusdo apparentlyoccur togethersince tlre c ollectionsMitchell3085 and 3086 are both given the samelocality details.
El Hadidihas annotated a duplicatco fthe Symoncollection in K asZ .zeyierlSond., another spccies which is only doubtfullydistinct fiom 7. crlolde,r,but this is clearlyerroneous since T. zeyherilacks an elongatedstyle (see under Z clrl.)L/eJnotes below).
Differencesin flower sizeprobably relate to soil type sincethe larger-floweredspecimcns of T.ranunculll orus areallrecorded as com ing from blackclay arcaswh ilc thc smaller-floweredspecimens arefrom sandand loams. -l
pe)ro^\oq^\'rauod.{qpauuguocsr.rrp.rolr.r, J.}o uor}r?culr".n,"ltNjf"';;[:$J:::r:Xr:lil; 'puelsueenO ,0// ul€lsee-qUoutu04 ,{lqpunsard I urset f,eds s rqt pelo:rlloo lepuElos pue sIUEB .raloN
'arn1aa3 SurqsrnFurlsrpln1asn p oslesr stelrqEq ol uorlclrls.llsll .roFlDI,(lqercprsuoc 'st4sa'ual lelscoo 1nq J,loasoqloladeqsut.tultruts'sltn.r3snorqel6qnspueaur34sueql.rc6uol(6uo1Lurug-7:r)e1,{1s '(3uo1 ruurg 1-g 1 s1e1ed)sra,rog aSrel slr(qpeztu8oceJ,(lrsEelsour,(llensn.satnloafButqsrnBu4stg
',{lnl .t8o1ouat1,1 ot Iudv,{UEIncrgud1nq rce,{ aq1;o sqtuoru lsoru sle,!\og t1
'rs'oru'rrurrsnv urersay'A eqrJJospuursr uro.,,,uo',.",,o"lLll,.i,li.ii'.ln:::ll;::;i:]1,:T,::jJ:ii uIDrlpJlsnV .IEolon ujeqt'louUr sonrl€f,ol lelsuoc ol pauryuoo,(lutlllllsr.rrplols.7, .1. puo uo4nqt4stg
.(CV) .g ,1961 .Burllrns r08 f I aoar,{5 {e118 I odel rVgNtnC,^\_JNVndVd '(B !.1.r.rrtJrl '096 .lec .allr^su,\\oJ .uorltts NVJ) t 1 g l lrpuejsilpd s).rcclIltuorleN:sldulsnuEtu J€lq3rE ut qderSoloqdsu ue as uetrlrrads - (t{ ,,{tllroo B) , s rapuDl..)<-tr,eluDg,0r, I tnoqt!,\\:C NV.l SNgEn '(vNo 'BNVJ) O uDtutLlSttll 'dnorg,trel1e4prearpE 'puPISl I8s I 9 lg qlnos :(VNC) &rr-3? .CV) lse,^A 98tZ I.'pua1s1qeddul6 o1aetr :,{eg pn1,1en1g :(t.t t,VNtC tglZ lq)alqlv g '9661 rdy 'pualscurog pr'olf,rA o 3 I su^\oo13^ru EN nr) s.e :AUOJIUUAJ.NUEI.IIUON (Hlugd,gNVl,EI^JOOUS) Ie ! ral.i/,JIU,pUBl -JerdLUp(l 'drrlsuMol N lulod rlJV euO :VI-lVUJSn V NUEJ.S!.ly\ .:paunrnxa sudtu1)a.ls a^jJDlursarcla{
'(t0Z;!S6lspueuli/Aap1/'otn1)9g7 1,9g2:869luugtrtralj.rrl,.iJ.arorprlul .1 oJIreI'rlVurtulrq?H.uottDtitadilJ(gStt,(utr,t l) tgg unruluelds.rccd5 saprolsrcsnlnqrll .Zl
suolflln)unuDt J Jo-jpeploJel trql rBr,r sppole eru (tuLrlg-t) tllpl,t\puu (8-4) roqtunu leUeel eql pu€ (ullurlsny ur slrceds srTzgr;7 ,{ut.roj pep.tocarun sre1,{tsjo q18ue1 srqt)uorct aqlroJ eBueJ lDnsn eql uDql (turu g re8uolqrnu s!el{ls aqlqFnoqllu ,ereq sBuoleq flqrssod 'u^\otDlun 1) srErlDllsnVulolsay'A ur uorlecol esoq^\,(gg09l/ E0 HIUEd {aerJ,{EilcDr\,1uor3 uerurceclsy
91 ol dn 0l oldn (uu) qlpr,rtrn{ e) t-l (tutu) qlpr,r laueal (8)r_-9 0 t-8 0 t-8 0r-8 requnu leuPel s l-t 0 90 r.0-s 0 (Luu)q13ue1 uu8rls t-8 r , 9' ) L-t (tuu) qrBuale1,{1s 9t-sr OZ.SI SI ' st-Sz(!-r) (uu) qlSuclIEtad
sueurrceds suaurrccds 11c pera.rot+-llurus suerurradsed,{1 suautrceds11u
salt!ors!J'J suq Iln)unuDt snln(t!tJ SJJIJEIEqJ
.,1 'snto1/)lntunun_r saplotstJ J pulzstuoLlllln)unu 1e 7;o suarurceds €Jrnunun) .7 peJaMoUllatus\l|^\ stuollliln)unuD.l/ .'o sueurceds:rd,(t eq13o uosl.ruduro3 e1qa1
.r.Irufl sz snlnquJ lo silou lur irl lsrit y l^JI 26 Nu!t\ir V,,l ll, No. I (lr)()8)
A rcccntcollecfion fiom theVictoria River Downs I Ionlestead(l). L. Alhrccht1434)agrees entirely with 7:cls/ol /er specimens tiom coastalareasand probably indicates thatthe sPecies is capablc ofbcing transportedinland and sustainingitself.
'l'. ci.rtoidesof south-easternUnited Stalcs(Porter l97l) and tropicalAfiica (Launct 1963)is dcscribedas having thc intrastarninalglandsunitcd into a 5-lobedring Squires ( | 969)also suggested thatZ cr.r/oilerhas the glands in a ring.This is not in agrcen]entwithspccimcns in Australiaherereferred to 1l catoidcrisincc intrastaminal glands havc been found to bc quiteseparate Howevcr,Poncr also notedthat th is character brcaks down in theGalapagos Islands. F-rom his mcasurements, st)'leand stigma lengthof 7. ci.\toides\.t soutlt-easternUnited Statesalso appearto be much shorterthan thosein Australiirnspecinreris. 7. ci.ttoides is predom inantly m aritimc in itslocations in thc Amcricas and Africa, in agreemcntrvith wlrat has bccn documcntcd Ilere
Distinctionbctween thc two specicsI cisloides and T. zey-hel"/apparcntly rests on the relative length ofthe stylcand stignra, with Z cAloi./e.rhaving a d istinctstylc. whereas the stylein T.zelheri is morc or lcssabsent (Launerl 1963). Schrvcickerdt( I 937) rccognized thc two speciesas d istin ct butreferrcd to thcdifficulty in distinguishingthern in Afi.icawhilc Porter( I 97I ), aftcran c\amination ofspecinrens fionr both America and subtropicalsouthern Africa, consideredT. zeyherito be a synonymof 'l'. ci,\toides.Ill Iladidj (1985) separatcdthe two on thc styleand stigmalcngths and aiso on the pubescenceofthe plantsand thc lengthofthc floralpedunclc u'ith respectto the subtcndingleaf; hd/ commcntedon thc difliculty in allocatingsone inlandfornts. Fromnotes in Eichler'slnirnuscripts, El Hadidi's annotationon spccimers (seeunder 7l rununculiflortts above), andanlcntion by Wilson ( 1992a),there has been solue thought that f. :el'l?e/imight occur in Austualia, but there secms to bc no nraterialrvhich rvould suppoft this hypothesis.
7i cruloklesntay rvcll introgrcss with othcrspecies in Australiaand Aliica andthc followingnotes dOcumentitS rclationship to otherrucmbers ofthc Z /e|resll/.!group in Australia.Ho\\'ever, variatioll in charactcr.isticsdocqmented in overseasmaterial (see abovc) indicates that it is in needof review throughoutits fange.
T. cistoiles antl T. sp. hng-stlle eichlerianus
Specinlcnsr,ir. linel WESI EIIN AIJSl l{AI-lA: PicrCreek, Warralong Station, N l Butbidge826 (PflRTII,2sheets): 5kmNEof DeCreyRivcrcrossingonroadtoBroome,DE Symon 10100 (AD, PERTH),
,\bten.Sonr e collcctionstl1' Z sp. long styledeichleianus,c.gBttrbitlge 826 and.Sl'rioir 10100,have fr-uitsmore like thoscof I clstoirl'srvith respcct to theirshapc and distributioD and numbcr ofspines, but they arc invariablymore pubcscentthan coastal populations. It may be that 7l sp long-styled eichleridrut:;should be thouglttof asan iniandlbrm of t .lsto'1cs,but this nccdsto be investigated fufthcr.
T. cisloides and T. occidennlis
SpecittenscxLlninc.l. WESTEItN AUSTRALIA: North Wcst Cape-lighthousehill, 3l Aug. 1960, ,4.5.Georgc lr391(PERTH). .Ellujlsnv urlxcpue r rleJ lsnv utilsoy'ALr I slntoo oslu s^ct?ods erlJ qtnos puu,itol rJ.re.l_ ulo qltoN .silulA qlnoS,\\aN'puelsuJtnf .srn7lral4rla uro.rj,,^ueurrcads pclcelas pcts^rl(e266 t) uoslr/A Z Aurqucsepul
'(9661(luroc sJ3d 'lqctJqlv c) soolu)ooJlculg pellul-pouorluo pooJ elu no^EJe xJD stluot ta14 rtz JJo slrnr-]oql srlo,!/
reBuorEruBrrs^ oqlpue s.urds l''srop uurprru !orrs rrlr \\ qrur pr," q. q"rq, r1"l;l,t"ll;::: l;:ljlill,iTl 'ila,l 'esucp ecuccseqnd le,rlsoullc oqsaql .{q pezr uSoccr sr rru Jalqt..,l..s.atnloaJBut4s.tnEutlst6l
'(eZ66 l) uoslry'eas :SglVlS)t:IHJO '(HJUgd) tt6l ar,LrXO U' Lt6t Ktt\l9|,r€^rU.{o.rC .C reddn')carJ ,,{e11e u,\\DIleJ:(HJUg(l,BNVJ) 0ZS!-I ?)i.).rlf S ,,r,spuDIelquJlsiqtno5 ,1uopeaJglo eprs ,o6og,lcoppud ,.(915114) S:(l i.l.U:ld)tzo0l )auptDC V)'L-c61 rdy 71 es:o11 us nuptDg.y ;1 'lt6| .rdV6l,re^rUpmuuc.l,drr.rDCloodc?rlod lco'eurnoqaouieou:(HlUgd)tStg pp!/uD.Dl A,gg6l 'pusls :(HJ.U:ld)9901./1a1lrrfl U'0861 lro l .,!\olllrB:VI.IVUJSnVNU! ISAM pautLuDxa sL!]un )a.lS
'(lN 'BNVJ :os/:/A SN. oloq)talDg ll ',y 'tg6 ,,{.lottrre.l. ,J.}o_urnl i9t9luasll/14 ? I ludv L I uroquoN Drdolll Jo tsc^\_qtnosu1 9.,(u,$q6r;1 .'1.) ie^opulrs:rp.roN ll?Jtue) :adIJ. (a661)lZt(l)geodoleJ,uosltlA snuuuJlrlJrrsnlnqul .rl
'ariq,{ldde 11e,lr,{uuruorluucldxe eurus.r ueql.Erlerlsnyur slrraclsrlloq Butllsrl eq ol punoJeq .llele)coJ uDcroleurllodeJl iapnlsii J ur sl€^\ol}Jclleuts -+o uotlcnpo.rd lq1 o1 iLt rpuol puu .rr1.iin./rp 'acq . ..L odoJollx raluadler crruapue eql iq palelrlrcel l],t\uHul .lncJoso op st 4 s.nr.!.t puesa? o/st. .J uaa.,'\lequorsserSoJlur ,Ja^a,rol ltql palpcrpur( tL6l) leuod lpqtpelou iq osll?plnoq\^ :ll I ttn{ eql uo Sutra,nocluq ollccdsetqtl,l,t elqullr^ sr slql e.urs ( ,i /r)I /rtsr-r./r/ JJo uoxelpelils ro8uolerp se poteorl eq Jfe,{\lqBrtLI,{eql strtsatlat,l,ursee;esaurdseqlpupr^ctoedslsrl}utsr luerseqnd flesuep se lou sirnl-1 ,(e,loqr: eql'pul6rlscql uuql.rlSuolsr lr1,(1seq1 :rcurs srzr.rDrDlLpta J lou a;u,{eqtpue cos)uruirls aq1 uut{t .sucrurleds .11ou Jeiuol ,{lquteprsuocse1,{1s e^eq iaql ecurs aqlJo euo uo pasodordsu r.ra4,ta, ers,{orJJ_ ra^o e.uaf,seqnd Ilu uorlsu Lllll\lnq t-i.Osr-/./rl7.1o ltlcrdil ores1n4 eqtolrq,\\ sj.lts Dt J, 'scrceds roJleql uEqlJa8uol lnq saPo$i. J rol lJorlsoot sr q13ua1r1,{is eq1 eq1ucc,rleq elrrpourralur lcq,lteurossr ezrsra,\\ollraqj serf,adslerpreqtr,triuriuolcqilluerlouelrq^\r:)p/ots-t)..I.pulr.tJlsltJ)l.J Llloq scrlslrolJurcqJ .J",l.)N Jo ssassoderlerlsnv utequou prn?luluro4 suaurrccds-+o raclunLr V '(g 'e.rog Nlvc'tug) tgl,1,Trrrls. 7/ N'uorlulsiJaiunllr11 lsrnqpues rc NV-IsNggnO lSNVJ 'CV) uotyrDl' 'a:U|l3 .ir.,{^\H ttOl S N E qle)Jo N ur) 0l lrrnls eprse8:AUOttUU.JJ NUSH.I_ION '(aNv)) 'E.|Jnunun)'uorluls zrtatna7 , qcrDeseu,{euaqurr) OUISJ :VI'lVUISnV NUg.l.SA/\\ pautaDratLtaut))dS
lDl s strlsau4.J pug saflots!) -J
xolduroc slqsa.uatJ oql.Ioserteds sr?1/lgr_rJ ur eduqs 1rn.r1 oqljo lueudole,lep:lql ut ,trer;su1deql el'llsnllr ol scAJesuaur Jeds :t qJ stlDJuaptnoJ. to slr nU e qUo J qs uatJp.reqc ero u seurds Bur dole,lep eq ol sreedclr uoncellorsrql uo lllt{ 8uno,{laqlo eu6 sno.rqulBqnsaJE,^dpjolstJ l, -Jo esoql sueleq^\ lucJsaqnd oJe 'le,\ea\oH '(trun!nlq slrn{ esaql d .1,peq s-ttlsat)atJ,.}o s[LroJeuros pur) srf.foll.iJ j ut es^oql otedEqsJulrutsu-]osyr nl3jo ,{1r.roli:ur eql seq (t6E la.t.rrll) aduJlsa16qyop LuoJ-luortcelloJV LaloN
.r.IIrB uttrqtrJ uo sitou pur iel IrtI V N ll NuylsiaVol. I2. No. I (1998) specinlensare Iisted here. Across its range the fruits are not alrvaysas large as recorded by Wilson,nor arethcy alwaysas densely pubcscent. Thcre is a greatdeal ofvariability in thespecies with respectto thesizeofthemcdian pairofspines, rangingliom al ostabsent to almostas lalgeasthose in Z cr.tloides. A collection froln thc rocky coastsofBarrorv Island(R. Bactle1,7066)agrees with T.eichleriqnus in a1l respcctsexcept that thc coeci are only sparselypubcscent, while a collectionfrom Southeskl ablelancis (George15520, CANB duplicate)has thc styleon oneofthe fruitslonger than usuai for this species.
14. Tribulus sp. long-styledeichlcrianus (,1 S Gettrge10666)
lTribulushyslrix(shot1-spined,smallerftuits)KL.Wilson,Telopea5(I):28(1992).
prostrate.villous rel,r, stemsupto,10 cm long. 1-e.7yesin unequalpairs, more ralely alternate, with 6-8(l0) pairsofleaflets above 4- I 0 mm longpctiole; leaflets narrowly elliptic, shortly petiolulate, base - -4.5 oblique,acute, discolorous, v illous adaxially, rrore dcnse ly v illousabaxially, 6 5 13 mm long'3 mm 7- 10mm long, widc. Floweringltedlcel 9-28 mm long,u pright, fru iting pcdice l l4-35mm long Sepals glabrousadaxially exccpt for apicalpubcscence, dcnsely villous abaxially except on hyalincmargin. Petalsyc llorv, obovate, glabrous, l5- l8 mm long,longelthansepals. I nltastaminal glands 5, indistinct. St4ner.t l 0,at maturity hcld.iust below stignla; fi laments3.7-4 mm long;anthers l .5- l 8 mm long ')t'r.v denselyrvh ite pubesc€nt, 5-celled; style 2-4 mm longin flower;stigma 0 6- I 7mm long F,rllll8-I5mnl high,I I -20mn widc,dissociating into (3)-5 symmetrical, moderately pubescent, woody cocci, with short -4 tuberclcsall ovcr,somctimes also lv ith a longerpair of medianspines I 5 mm long.
Specimensexdtnined. WESTERN AUS I'RALIA: PierCreek,Warralong Station,N T BurbidgeS26 (PIRTH,2 shcets);19 May ]971,RudaltRiver,'4 S Ceorge10666 (PERTI l);Bloodwood Bore, nearBalgo, J,rly1g72,C.1!. Giuins2452 (BRl); Fanny',sPeak, 85 km SofGiles Meteolological station, 29 Aug. 1973, B. Lay861(AD); MinderooStation. Site 28A LandsatStudyl , May 1980,AA. MirchellT04(PERTH): AbydosStation, S ofPort Ilcdland.Sep. 1968, F Richardsttnl9 (PER']'H);!7May 1947,5milesN christmascreek. Rabbit ProofFcnce, n. D. Rolce1741 (PIiRTII);5 km NE ofDe CreyR ivercrossing on roadto Broome,D. r. syaror 10100(AD, PERTH);Rudall Rivcr area,7 Aug. l9'71,P.(i. lltilson1029'7 (cANB-2 sheets,PER'l H). NORTI IERNTERRITORY; On l'anami rrack, c. 472 kln NW ofAlice Springs,140 km SofRabbit Flat, P.A. Fryxel1,L.A. CrtwentlJ. McD. Stett'ttrl4505(CANB); RabbitFlat,42 km SE ofTanami,D E Synon 6904(AD). souTIl AUSTRALIA: plainbetween Tomkinson and Mann Ranges, on westthca of inselberg,c. l5 kmbyroadNNliofturnolTonPipatyatjara-Putaputaroad,c%km E of Waltjatjataroad, 6 Scp l9T8, N.N. Donner 6582(AD, CANB).
specimen.texunined of intennediates.NoRTHERN TERRI'fORY:46 miles[c. 74 km] E MongrelDowns, 20Apr.lg7l.C.R.Duntop2l07(DNA);AlongsideTanamiCreek'306miles[c.490km]NWofAlice Springs,20Junc 1976, K. I/rut 76138 (NSW)
Dist butionL1n(lecologr,. Occurs in thcl'anamiDesert arcaofNorthcm l'erritory acrossto the Rudall Riverand Hamersley areas and south as far as Gilcs in WestcmAustralia. Little is known oftheecology oftlris poorly collectedtaxon. It hasbecn recordcd fiom red clay loam \''/itl1Meldleucd glomerutd, At'istiddand Sl'lobajiurn-fron stonyloam and f'r'om hunmock grasslandin sandstonchills
Phenology.Flowering is rccordedfiom Aprilto August. -l
'uorlsenb olur ,{urouoxelluosord cql Surllec.lotccJ leqloua 'suoudocxe sr qcrq,lr ,{uuure.tu a.taql pue elerutxorddu,Oe,l sr peluclpuruorlnqrllsrp aq1 e,toqepelonb seuosurol i T,llof,s Jo llo,r eqlol taJorsdnor8 eufzosr puerequnuauosoruoJqJ sdtqsuorleler;req1 puu a:aq pczru8ocerxeldtuo. s?.r,1sz.//at snlnE,U eqlJo exa!rnoJ crll Jo uollecrpurue se,rrBg e1qe1
'8tJo Jeqtunu eutosoruoJqc€ eluq osluqctq/$ suorlcelloo tllul\n)/uerpul qlr,{\ uoxel srql ec€ld osle plno,n srs{1uue '(e966 au,{zosrJeqleqM oas ol 8ur1se;e1ureq plno,r lr 1) DocS?uosrJlol^trotSurproJle(gt, =uZ) 'ErlpqsnV proldolcouu sr sneeo Jo,rna J e)uts pecnpo4ursr (e1,(1s ljoqs) s?.r;ixa./,/a;iJ alr q^\ ol cArlEueJE 'snr)o)or2tLu (e1,{1s3uo1) sr.i7sa.r.rar J pup.srynulw J J l€ql slsoSSnslJo,r s.uostJlot{ ? uocs lnq '(ereqpazruSocer 'u€rl esoqlo1se1nqul1e ut f,1ue Itrurs elquUe uo.re ^\oqs qJrq,,$ ro.J(Z/61) lplpuH tg {q pazruaocetv4satta,i J Jo serlerJE^eql eldurexero3 ees) e;eq,nesle uo{ sarradseqUo sulloJlu3JeJJrpJo .e1,(1s suorlcnporluro8e 3uo1 tueserdar ,{eu ex?lJnoJ IID tuql elqrssodsr 11 eq13osq13ue1 e,lr1ule: uo ' 'suorsn pessq s sl /4srrla/ 7 urqlr,n craq pezruEocel uxul IEuuoJuro,rl eql ol uoddnspuet ,{utrt lnq IJuoc Surlcrguocleq,neuros ol pcl s€q(q966 I llo.S Z' uosuJol,!)seru,(zosr puu (s9661ltols ? uosulot{) stunoceuosotuorqc'(u966l uosrlot{ T,llocg) .{3o1oqfuotu:.rnquo oar,{ruunurlardt:xel secslc^o qlr.{\ drqsuorlulorrreql pue erl€rlsnV ur uxe] cql uae,rlaqsdrqsuorlelol puelsJopun ol qsl,ne,,rr jr ,4pn1s,{3o1orq r€lnJelotuJoJel€prpuuc poo8 fue,\e sr 11 epl,r pllo^\ puEerlEJlsnV ur qloq uotsr^etJo peeu ul qrnu-lf.la^ st 'Jleq dnor8eq1 paullsnfsr snlelssrrll tpqt InJlqnopstuees tr q8noqtuc^c uorlecgrsselc1eq1 e8ueqc ol pesodordlou sl ir pueerlurlsnv urqlr,r Is^el serceds1u pazruSocar ueeq fpee;1u e^eq'snnuta J '€x€l pue snJJocot.twJ' ,tlss st4satJalJ eseqlJoaelql ,{8o1oqd.rouur epulSlelurflqtldariedLul pue uorlnqlllstpur depe,roqcrq,r ExstJotequnu EJo slslsuoc€rlurlsnv ut'tDl s st;4s tat J
xalduot '1 syl.lia at snlnqlJJ
'uoxpllcLlt ur rJllELus qJnlrrJre qliuel el,{ts '( puesle,\loU eqt lnq ^ b) snfio)oJzttuJ ol pou;tssusueturcads uer;erlsny 1e;1uec pelrnl.; .re3ju1 eq1 Jo oruosol e3uelqueser?uo4s e recqosle slrnrg sorcadsleql ur se3uo1 su 1ou sr e1f1s eq1 1uq1ldacxc '4.4s,f? '€ul8us J ul punoJsr s€ ?qlJo lpql urqlja8uo1 qcnu ,411ensnsr q13ue1 e1{1s a ql all q,{\ t?rli.,{q1J pu€ stlDtuapnxoJJo sazrsle,rou eql erou SurlqrueseJ'snunlqzra 73o teql ueqlre8rel qcnru sr ozrsre,rolC snuDlolq)ld J ur punoJlpql wql e1f1sta8uol qcnrue ,,{qpeddol puu tuetsaqnd ,{lasuJp .ds ss3l ere ^aql lnq sruDlplqqa I Jo esoqtelquesar snuDual4napal!s-*uo/ J Jo strn{ eqJ
'Eet€3ql ul 'exel ,lue] ?ut1oa1locrood lceger lsn[,(eru,{aqt -ro cqtlo flrpr1u,r eq} uo lqnop lsec snql putl eq,{e ur pxt;l esaqlJoqloq JoJ uortnqrrlsrp ur suorlcunfsrpe8rel eq1 uxelaqlgo sdrqsuorlsleJ eqlJo SurpuutsJ.pun .Io)cel eql ol solnqrJtuocos11? prle{snVJo uor?er lsa^\ quou pu€lur eql uor-}suolgcallocjo ,(lrcned eq.1 '€xel 'sBurssorc l€sreJl] oseql Jeqleq,4\ pouleuecsE eq up, 1reJoJeq'Jncco op feqlJI qcns.;oseuroclno ,€ru6rls eql puu a1eur11od-sso::ot srynqi)J Jo sar?cdssnouul eqtJo {trltqe eqt puu a1,{1s3o sq13ue1 e IlelaJpuE ezls te,roU ur uorleue^ pueJo aruzrgruBrs lctrSolotq JqtJo pJInba.r sr Surpuuls;epun repeq 'J .ds V saptoJslr ro snuDual4)tapaltls-Buo1 7 go esoq1,{pee1cete sqlSueluu8rls pue o1,{1spue ezrs '(slrn re,\\og eql lnq 4 ed(1xr,t1stt1J lleLus e't) stDL[au11rs ds J Jo asoqlot .cuelqruesejJ€elc e ^\oqs slrngoql suorlaalloc eseqtul tlesecrtuEuelaqtluo{ qloq'g€ lgLunHpue LOIZdoT,z6re1r1 suaullrods 'stolfaLulos .ds elElpeuuaturf€uorsDcooJo ecuelsrxa oql sr ds Jp\n snuol,talq)Dpa1t1s-tuo1 sn1nEt1, seureueserqdeql,{qpel€eurlepoJeqexelo,nleqlSurqucsep,{llDturo-+louJoJSuoseeJeql.}oeuO.la,roN
'ralrefl snl q J uo s:rtou pro iit lelr V I,\ )l NuytsiaVol. 12,No. I (1998)
Table3. Comparisonofthc taxarecognized within Australia in the Z /er'len1,"$conlplex.
approximatc chromosome isoz-vme style cocct coccl dislril)ulion number grouP length(mm) spines
T. tcrrestris northcrn 24 C 0.6-1.3 not dorsall), longmedian (long stylc) Australla rounded spines
T. terrcstris sout h ern 24t36 0-0.3 not dorsall)' Iongnredian (shortstylc) Australia rounded spines
T. micrococcus centlal 48 0.7-1.1 dorsally medianspines Australia rounded lackingor short roQId
T. minutus southern 0.2-0;7 dorsally medianspincs Australla rounded lackingor short
'1 . rnicroc occus and T. rr iiirlu.!,w ithsnlallcr fru its in which the 5 dorsallyrounded cocci are usuall,v tubcrculateallover and either lacking or rvithmuch reduced mcdian spines, havc been recognizcd in thc pastonly fionr lhe eastcrnstlrtes ofAustralia. A ccording to Wilson( 1992b)th cy canbe distingu ishcd frotnT. tertetjttis by stigntacharacters. but thesediffercnces are oftcn not clearcut. Instead,within spccimenswhich havethe vcry longmedian spitlcs oftrue Z 1elrs.t/rl.e,thcre seem to be northernand southernfbrms bascd on st1,lclerrgth (the longcr styles usually found u'ithin the northem populations)' Sirnilarlywithin the smaller-and shortcr-spined spccimens T. micrococcu.rlcnds to cometiom mole northerlyareas than L?l/?r/1i./sand also has the longer style. Again,this is only a tcndencyand there arespccimens (e.g. -D unstun(AD9624707 3) frorn Adclaide has thc longerstyle ofZ tnicrococcu.s)which do not agrce.Thcse snallct- andshorter-spincd species are not confincdto the eastemstates but are to be found throughoutAustralia. although thc fruitsare not aNvaysas small as in the eastcrnstates. Thcycan usually be distinguishcd from T.e ic hler ienus because thc fruits are smallcr and not so dense ly pubesccnt.but tllcy resemblethat species in therounded dorsal surfacc ofthe coccusitnd any revision ofthe cornplexshould also illclucle that specics.
Thcreare some specimens which have thc attfibutes ofscVeraltaxa. For cXam ple, somc spccinr en s iclentificclas L n icroc(tccus,have fruits approacling T. eichlerianus in pubcscence,but the lcngthof spineson thefiuit aretrrorc rem in isccnt of 7l /crrestr'ls,rvh ilc thestyle an d stigmalengths are simi lar to thoseof I micrococcus(.e.g. P. Cunt.v2186,Nullarbor Plairr (AD), M Cole42 Fr.uitsagrccing rvith one ofthc smaller-fiuited species can sometimes also be fbund on thc same plant astiui{s rvh ich arccloser to 7l /?/'/d.t1/r.r,e g. felrl s ti , Doodlakine(CANB). '1'. acanthocpccus ntayrvcll coinciclcrvitl\ T.minulus or T. micro(accus (sccbclor.v) but at this stagc only a photographofthe t1,pchas been seen in thc Eichlerrranuscripts. From Mucller's description, thestigma is only hatfaslong as the st1 le, suggesting that it wouldbe placed either with trucf. /elre'!1/h -l '\cv ) s6afl t)l|.unH J Y '6S-61 JdV L'suurlueSrV'uuninuulule oqurn:'urnb1e6cpllpou lp nr)tt J:VflU'J!\,lVH"l,nOS '(6y) g9gI t.i;77t27,r5O ,l'L96t txoj.. 'clrozuv 'uosrnj, 'suxe.1_'oJ '.klv) :(CV) //l ItnA v'9-61 8ny 1 tu0Jrel S6 uD aj)]4 e '7961 'rruroJrleJ',{}unoJ 'sr^rC 'leloH '6t6 ietr1 9Z oloA ,(lrsra^run:(CV) t0lt srTd',/ t ,(lnf 8l 'uuDrsrno'I 'eoruo!r{ etrqcrno ur 15rltg.lo 3 pur?iS prrseo lo qlnosprc,{ puorlruu oUroed l.rrossrt\i '816 'uutrsrno'l'uosrpD!\ :(AV) Ztt WtV O i 61889rDtuoLl.l. alD0 U i eunl Et :VJl)lgl,,{V I IIUON L^ASN'BNVJ'CV)0080 1 p./o//tJ U7'rrequuJ 'urDo reue,\uoS:(A\SN'BNVJ) g6t t4rsdaj f {T'8uot?otjostrr)istno ureqt-roN :SE.IVA\ I IJIIOS ,{\:l N '(g1y'.1.1) '0t 'urlteqrupJ 'BNVJ) 9;g;a,ro,l I 6 t ,teN :0 IlU jd trr1g6l11o;uu,ty.4 y 'curoorg 'UNVJ) :(H.LU:ld /96t ra,1D./Jy7 ylpxltg f ,1't86t,(u119'iurssorrre.rr)llcocpVlu 'BNVJ) 'q!.d 'pDoU puou foruelD uo uornrlsosnol I tN :(l IJUgd 1,7711pp(un4)/ U qlnos rll?rrrf 'eJnllncuSV ldoe clrsoddo'1co1qlueceA :VI'IV)llSnV NUgJSgA\ paLlturDras-uautna.ls papalas (alils lroqs) slrlsarral snlnqrrl sSI -c.roq pczruSor:r.r(11,{1s yoqs) ri;7r'a,r,ra17 'adit qlt,lrploeld ,(llcerrocsr eureu eqt to edit aqllt;qt pounssc sr lr eqtsu ueurf,eds rulncryud srql ctrc ',.luesqD 'uoqs,,3ureq qloqif,urs pue ro uoqs,Ce^,, sr lprprH MuD ..paJnpa:qcnur sulueulr?e.rl srq ur sr.4sar,rai7 jo al,{lse qt scquJsJptprc)jcro,rLJcS ro^J,\\oH taatls:^nr'outEnL!n l J crl} uoteqt uEq}.rellcLrrs^ 'lrutep ([quraprsuoJs! lr lErlt,{cs o] tdeJxe lno eltlu ol elqrssodurrsrlr pur lueserdtrnr; elFurs u,{1uo seq t1 (!-S6I)IplpuH lq (q put (LE6I) lpieloro,nqrgiq sorocdscLIrJo rd.(r cLIr se potrc ucaq suq'(7'1yc 11511'1) s1.ttsd.ltatJ sDpall.qpl sr qcrq,nueurrcads eq.t (/ 861 e1u,(uuussrqeldurexJ JoJ cJS) st.lt,saat JJn Lu(uou,{sDsp pcleiJl,(11ensn,nousr snsoutBrtLltl .1,:urlurlsnV ur s7rlr'r.1./a7 17_.1o sueu rceds,,poJnpoJlrr,, eql .JosuluJ Jqt ol .lzlrrurs.Oe,r sjreddu puu'seurds losrop ul?rpetrl 3uo1 q1r,\\ slrnl.] lsnqor ,{ra^ surl (e1uu1tr5) uo1,{e3tuou snsoLBnuDl J.Jo Z'ltS NNII lciqs cql'unr.ruqr.qrmeuuur'1eq1ur luaserd sr qclq,,rrleuetuu oql-J6 serledslqt 3ur,{1rd,(tolcclurtunocru olur uelct eqol spacuqJr q^\ IurJcle u.rorllo aq Ile,\,(utuereql puu sdrqsuorlular t:ur8rls puu e1,(1s su slrelep qrns euruualap 01 elenbepeur sr 6y 1u 'slrclds sed,{1uucuuurllo eqJuojcrttreql snoeuoraJq plnoxsrq] tnq cqt3ood{l eq1q1r,$ seprruroc 'DlluJlsnV ol peJnporluruarq Sur.ruqsu p:rzru8occrsr qotq,\\uoxlll pel,(ls-lloq\JLlt lrrlt liLLrn\\r ueeqsuq ll \utsa,tratJ Jo uorlecgrd,{loql sl pesserppuueeq }o!r suq qllq^\ l)edseJeq'unJ ouo 'urlErlsnvot eArlsu,{lruleq ot pareplsuoJeq uur uxDlqf,rq,\l -+ouolletrpur ue c,lr8furu 1ruaql arnlnl arll ur apuur cq uuosuostreduloo Js:tqlJl Exp]uerlc.llsnv crll rltr^\aprcuroJ,{lnlt oseqtre{taq^\ oprocp ol clqrssodlou srlros puuexDl aqlJo suorldr;csep rprprg 13ur palucrunoop,{eqle-rD roN s.rloue^oseqtjo sqderSotoqd eqluol..l sqlSualuurFrts puu el,{ls eqtuJecsrp ol olqtssod 1ou st lt ,{1e19un}oJuf} uEJIJoJ(Z/61)rprpl?HIlfqpezluSojar'st)t|a,ua/ J-Josetteuu,\eqlol alulo.llle fuu .'rroleqpezruSorcl exel rno3 eql l€ql illltqlssod u sleleql'e,\oqL' pJuorlutur iprup sy 'pcztuFocer .{11ua.unrsruuql eaJe JeprA\ e ralo tlxtlleseq}.lo aLrros.lo o.ucJiniJo Irin}uu aqt elclrpur ,{lqlssodplnolr lr JoscosJelo uoI ulll?rlsrlVolul s^oltcdsJues aqlJosrulo-IJo suoqcnpoJlur lueJoJJrpJo ,{lrlrqrssodcq}sesreJsrql'ouoJuaqtlourqtjoc1{tsraiuoleqlscqqJrq,\\utpulutor.,tuorlcelloce16ursusr 'DJu-]V ereqt'rJ^o,,!\oH r?.//.t4././a,?J .lo urJoJuer lcllsnv uloqlnos e ql qll^\ sapn rlloJ Eu t rlJ puDedol n:l 'spJIJeuV eqt uo{ luuctErulpql selcrrpulCV lo pleqsl/r'a./.1rt J -joIeUatEu seas.le^oJo {pnls V 'porJrd,{tolcelpue porpnlseq ot llrlsspq 19y1 ured,{1eq1 (696 1 ) scrrnb5 ,{q pelurr pur s.006 t i1.rueeql ueLlt.rer 1.rua qcn ur sD,\^rrleJlsnVursutsalJat J?e.\JoeJueJrnr30eqluarltreur:o1lqlqlr,nsiuolJqIIJ| .r/i.r.),,r,,/.)/rr:/ salupatdsncooco1lurz J ueql relel eql qlr^\ Suolcq o1e,rotd plnoqs 1r-11 str2)oJo,tJiut.l qtl,,!\Jo NuytsiaVol. 12.No. I (1998) EUROPE;Italia, I I July I 969,L Chqrpinet dl. s. n. (AD 97030 122);Yugoslavia, MacedoniaL Dolina Vardarja,24Oct. l9 64,E. Mayers.n. (AD 98135009);Corsica, l2 Aug. 1932,P. Aellen 128 (AD); East Bulgaria,Varua, 7 Scp.1953, Ar. Vyhodcevski s.n. (AD 98580847). ASIA: without focafity,s.ddl.,R. Schomhrrgi.!. /?. (AD 981461l9); Betling, China, T 9ep.1982,B. B. ll/an 82181(AD);neartheSummerPalace,Beiiing,China,23Julyl986,B.M. ltang053 (AD). AFRICA: Farm Erfrus, MessinaDistrict, Transvaal,2l May 1982,E. van Hoepen 1779(AD). AFGHANISTAN: Kalne,27May 1969,J.E. Carter323(AD). IRAQ: BaghdadAirport, 17Oct.1956,K.H. Rechinger, A.K. Khudairi&RIl Hainess.n. (AD). Noler. This taxon representsthe introducedcomponent of Tribulus tefte.tlris. It is distinguishedby thestigma being longer than the practically non-existent (0-0.3 mm long)style and has apredominantly southemdistribution except tbr somespecimens from aboutthe Broome-Derbyarea. Many of thc collectionsare subglabrousbut somearc villous. Z lerrestris(short style) appears to grade into T. minulus. Specimenswere seen iiom a numberofother countries as listed above. These resembled th is taxon closely,particularly those fiom Nofih America.Specimens from Afghanistanand Baghdad were not sowell developedwilh respectto thc dorsalspines on thc fruit andtended to be morepubescent, thus showingsome approach to I'. minulus. l5b. Tribulus terrestris (long style) Selectedspecimens examined. WESTERN AUSTRALIA: BungleBungle National Park, Ord Riverat B lueHoles. r(. ,llenlrol.rl625 (DNA); 9 km N of Kununurraat MocksDairy on E sideof IvanhoeCrossing roadadiacentto milking parlour,6 Mny 1993,A.A. Mitcnel/3086 (PERTH); Wittcnoom Gorge, c. 4 km from Wittenoom,Hj. Eichler22549(CANB, PERTH). NORTHERNTERRITORY: 9 miles[ 4 km] SEKatherine, T May 1960,G.Chippendale s.n.(NT 6712, PERTH);Mountain Valley Station,Gove Roadat BarmguerikbaCreek, T. Henshall3903 (DNA); FjrstSt, Katherine, 19 June 1964, D.,1 Nelsor 1229 (NT, PERTH). SOUTHAUSTRALIA: OlympicDam minearea,A. SmithperT.J. Fatchen39l I 164(AD). QUEENSLAND: BruceI lwy, c. 5 km W of Ayr,J.H. .Brome583 (CANB). INDIA: University Campus, Delhi, Mar. 1959,P. Masand s.n (AD 96033083). NoleJ.This taxonpresumably represents the componentof Tribulusteftestris which is usuallycited asnative, although, as mentioned abovc, there has been some Indian material seen which resembles this taxonin itsstyle charactcristics. lt is charactcrized by the stigma being shoftcr than or equal to thed istinct (0.6-1.3rnm long) style and has a predominantlynohem distribution. Thesing)e Indian specimen scen has a style1 .? mm longand stigma L2 mm Iong,making the style somewhatlonger than is recordedfor thistaxon in Australia. 16. Tribulus minutusLeichh. ex Benth., Flora Australiensis l:291(30May1863). - KQllstroemiq nlinuta(.Leichh. ex Benth.)Engl. in Engl.& Prantl,Nat. Pflanzenfam. 3(4): 88 ( 1890).Type citqtion: Queensland(?),Leichhardt's Expedition. Iype: IQueensland, Leichhardt D istrict],before Canal Ipastoral run] andafterwards , s.d.lt., L. Leichhqrdt s. z. (.syr:MEL s.r., p.p.,excluding lorver LH S specimen); without locality, Leichhardt's Expedilion,s.dut., Herb. Mueller (syrr K-Herb. Hooker, type seenas pholographsin the Eichlermanuscripts). 'urlpllsnV 1z.r1ucoolur Surpuolxe,puplsue.nO pup seley'A qlnos /|icN .uotUE)tsle 'zsser,tua^o).y,elt6urs o]peor uo eriunlcqlo 165 tur,, ,r."r,$:il"Hil;i'"lfJJi^tri .ppor ft\SN.gNVJ) g6l,uosl!ll 7 N uujleulrhtr_ql]o,rsllrd!uo{ peorurnq,{a13uo1u .a rul g:(SNVC,IUS) rr{rl8 J S,p^Jeurr{:CINVISNEgnO p anruorns.uarr,,rads '81,61 u! Ud u! urlscylacnrg ,{q ueID} sqd€l8oloqd :stduf,snueur .lelqcrg erll ur sqderBoloqd sBueos (Xd r.,1ll.r)y6yg uiwo11.0161 ,[puelsueeno] .cldo IIV rpl tr or{cuef pndr :(Ud) Z6V9u!Mo(j '016I rul 'lpuplsueeno] ,01 .re ,Lpuulsueenol tr sra^{oJ.s.lcue qJ :(XA ,trs)t6l9 ulutoe 6 I ,\l DpuoN 'ddopndusuaroqiurlloX.arl{-r'fpwlsu.on|ltallerqrl,rlqo,upun16l6f.rur\_ qel r.i/a1.)O)€puoN req su,toq .puEItuad .sJa^\oJ 3ur11o11;epucqcDL]ssetD reqlepl€,\\uauu€ADS sJouDqJteq ua11e1ssu.rg cBrg.rnp :uc.tr1t:4rtadil, 'lco) DJluDtoB .ururoc ./ GZO\ atz 4g lJi]H D)aqp!lcl!B sDJroJorJrrusnlnqrJr l .,{poo,r ssM1ua1d aql jo espqeql eJuts uuercd let u se,,nseocds e ql reqlaq,u.( 1 1 g LI apg )uo\)a h srquo pauorlsanbelDl8 .l .suortcallol (UnrlI]qJeq s luol.+luesqs ^lq]lrie^ur ele slJolsloolacurs sancds eqllo uotlnqr.4stpjoeBuul iql ssolae ple$eqluluorle8llse^utteqUnJJopeauursrJJlf,uuqJSrqf sr,r7r:.ij..7l j o lrquq 1enuuu X1.rue1c eq] q1r,,n parcduocs1ro1s1oor,{poo^{ a rsstlu eql3o esnucaq leruuctadKlsnorAqo aJo,{s.]JllC p?uwoJJsnnulLu J suorlelndod .1i17.ururor Jo usuolcrA lprplJeJ eql uopelrrru3J (966 | deg17 lii srad)suosru.f.,1.1 ;q ,{lesuapsselqcnLu c.rclnq s,4,?.,,,0",, , r""Tll',"fi:ii:":::ililil;jtTli,fli""';.t:;p;1i3 peluerlueeq oA€rl os pug aztsutsntnutM J ruo.r1,{re,l o1 ,{1uo urces,{eql ept,'!\ruru r l ol dn ?uraq.re6;u1 3rs,{eqllnq J'nrroro.r1tMJpLre r^Nn\otu IJoesoq}oledeqsr?lru SEJoslrn4a^lttl(.u.salDpua(ldtq) puu ztol' Lts uoBLoll .3.0)uoxpt .r.a,roN I0lZ O srq]ol alelalrllqM sueuncadsuurlEJlsnV lpllueJ ruersrxe-uou,{llpJrrcurdeqruutp:aBuole*r,o"u,rnu"tiorltJ,l,f;'ffi:,J[*:llH:;:i"1,::ffitit red uerp3ru .XloltJteJ eql puDpapunor ,{11us;op ere rccot aql u;equoNoLItul punoJels .lceosalJeqlo ,tue,teur earBeqcr q,n 'sue ,xa1dwoJ urc€dspeltn.U-Je8.ru1 qBnoqglt: s:t.rJr-tat J eql.+olseJ eql ur puno-'t osoql ueqlrcllDurs eJu uoxelsrquo$rn{,aLupueql LuoJJ pellodxe aq 1q6rur sy satnloaf1urtls.tn8utT stq 'ueqtuaB ,{q pclElouupuaeq o^Eq sedf}u{s qlo8 uotlDct/itlt1 '(sNVJ'ruB) 'tS6I 9t I Lltto^)luttlg y I?ll0I.gpunpFgJoSIrulZZ]silrrutI:(y1-99yy1g).u.suD|tol,]/O,uortDts lu.^\reo:(6tltl y .u.s VNe) lOVaztDT rr,ee.msa8uul uueuueled:(9 E/ I VNC) alopuadcl14lO,su.{oc uollrureH'uarv SurzeJD .J ,Lr6l ,su^\oc uase(I :(lUU) I I gaI ay)ft1 S {ul,{ 9l elaunrg puuuoo?D-l ,{uoqluyuee,rrlaq,{a^\JlpqlnoqV:IuoJluualNUggllgg.sTirz.r/.rai'rrl4/t^pamutoxasudat)ads c1dur,(.,1eq1puuq,e)eprspEor eqr uae,r,"o.o",I+1;;);?,ffi:'jfiJfl,f,i,XTl1iT:ii#li '(sNvo) 'qtno.IJo^AS sosg aloory g ll ) url 19.e1u8poo.turl uro4ur1g.1 ,,,ud1u,{ur::rnJ,,:Sg-IVA\ '(IdB) HJ1OS 11i\gN OOZZuosuLlol I U.pEoJoqLuEJ_-ul1rq1ainy uoulloLpe8ny ruorj u111 :CNV.lSNlEnd 'rur'c reaul s.ruaD,Furu,{ErJ:vr.rvuJsnv NuEJS.rrA\ ,,};i::};t;::'l:::I:3;yj}",lrT NuylsiaVol. 12.No. I (1998) and usually lacking any Distingtlishfugfedhrres. fhe iruits of Z tfiicrocaccusare roundcddorsally dorsalspines. lJnlike T. mhutus,towhich it is closelyrelated, ?i micrococcusttslally has a globose stigma,rvh ich is considerablyshorter than the shod (0.7- I .4 mm long),but distinct,slyle' Typific.ttiotl.Tl,pification ofZ n14ococctisis impossiblefrom thcphotographs sincethe dctail ofthe siyleand stigma is not disccrnible.It appearsfrom a 5ou ng fruit presenton thecollcction from Jericho thatthe styleis distinct,thus agreeing with thistaxon. Ilowevel,spines prcsent on the fluit of Donlfl 5493are I mm long,much longcr tltat thc usual2.5 ntm longspines ofthis taxonand so it may be that rhcsyntl pes repre'ettt a ntircd collcction Notcs.There is a l-cichhardtspecimen in MEL (Leichh.vdt26,MEL s.r. ) whichhas been annotated by ,,Tribulus h im as parviflorusm ihi" ft om BokkaraF lats on 3rd January1 847. It almostcertainly bclongs hcre eventhough only flo\\,ersare prcsent. "Flora The longcrstylc with a shortglobosc stigma as pictulcd in of Nerv south walcs" (wilson 1992b)ceftainly occurs in somespccimens ofthis taxon, butlhere remain a numberofspecimens which secmio bclonghere rvhich have thc non-globosestigma as long asthc style lt scemsunlikcly that T.nicrococcus andT. rl ltrlr.is canbe maintainedaS diStinct Species, although Z mlclococclJ doestend to havelarger flowcrs. Specimensfrom SouthAustralia, Northem 1'erritory and WestcmAustralia usuallyha; largerand sometimes, spinier fruit thanthose tiom eastemAustralia' and approach those ol7'. iichleriqnus and L sp. long-styledeichleriar's, differing from the former in the lessdensely pubcscentfruits and tltc lattcrin thc smallcrflowers and shol1er stylcs Acknolvledgemcnts Much ofthc initialpart ofthis work wascarried out under a grantfrom ABRSto providean accoullt for.,FloraofAustra!ia", but thc timetaken in tlle productionofthis papcrhas becn mostly unfunded pafis My thanksto l-erenaLal1y for thcilitatingthe compilation ofthc Eichlermanuscripts in theeally ofihis pro.iectand fbr herco'ments on f ibrrlusgenerally, siuce she had donc some initialwork rvithin group. the genus. l'hanksalso to JohnHosking for his earlydiscussions aboutthe T. terreslris The Eiclilcrcollcction ofprotologucs and photographs oftype specimensenabled the typification ofmost speciesand it is verymuch rcgrettecl that thc descriptionin anearlier paper of I .tr1relosr.sfrom these manuscriptswas invaliclated by ontissionofthc type. Karen wjlson's cornmentson the nlanuscliptwere much appreciatedand shealso pointed out a lectotypificalion of T. cistoides,which I had not seen. M), thanksto the statc Herbariumof south Australia,both for the useof facilitiesand for thc opportunityto seea t]umbcrof thcsespccies in the fleld in WesternAustralia in 1995,and to Bruce Maslin,Stephcn van Leeurvcn, I'hilip Shorl and Bill Barker.who all contributedin differentwaystothis fieldtrip. References "QLrccnsland Bailcy. INl. (1899) Iri6r1,A /": l:lora" Vol I p 172 generic ol Zllqaphylfum ^ld a ne\\ B rker. ll M (1996) Nc\\ ld\a. ncw combinalions. kcls and connnents on concepts j. Adclr;de l}ot. Gard. t7: l6l-172. spccresol:7iiaul,s {ztgophlltaccac)in thc narruscriplsofrhc lale Eichler..t "lilora Vol l pp 2S6-294 BcDtham.(;. ii8(,3). llgoph)il.rccac l,r Australicnsis al-ZI d.l f toA r^.t'rr\\rllio;-,\riN-Jo nrot,t.. (pr) .f).u.pnrtI .r/ snyqtU.lq166') .t.\ .uostr^\ 6Z-t7 :(lJSD.dotil_ (erl).n {qdoiiz) sr/r97{ uulrllsnV ur uoDr.rrrd{tor r tur s.r..rts,\\.u V 0ra66t)-1;1 r,,,.1,,11 ( .spUrui.r,!\ |urrplio)t :rur.)l!fl V V) ..surtirrltuo:) ,rl] .to .iurlo{t .rt.t...(t86t) O c (urtr:rn r.;urd() up. puz..snrrtdr lSrrtur psir.l\rp.1osildr.uud.. (ztait) I.tId ref ur^ rrlrrtsnV tt-8Zt 9ll tnN lirA rt tDl su)stutta.uat s lrvr.U.loursrqdro[rilod lup Uortuqrr]sr(J(6961) )l A,scllnbS ,,n{r r (e0irru,irrdoir'1)st.tte.kat sntt tu.r ro suorrurndod,,",llf];f;t,u;;,;'l,o\.1 .H::,vil:lXli\:,,". 8Lt'ZLl :! rrllrqlog uLll \i -tfr)rl.r.rltlr( I |uno.Lrrl,rlrl 1o srrrids ur.r.r]vqtnos ,rlt.to 1 no..r uv (1t6t) l,\\oll :8i p{rD 'to8 Unossrht .ritrod t-l uuv (ir..eltirtdotiz) snl9r.2 ur spLrrliluolt eqt uo si)oN (l16t) hl (I 3u,,ordx'rprr(srux,iorcrnr ,",,,,."^n,t"t:,"t"T,,Ll",ijiiliX,r"'l))t;),'"'ffi"1,;)l(;:;I':]'l];rf;"ll"l:;1""^ L8t :/ rl.ernqurlg i)o\ t{)g surx.l_rrl0ltsriV ts).\\-rltroNolul uortrp.dxt n-urroldrl '1981 s.iroBirrJ rl,\i iu nl rr..i.ql ur 1rr\orfl purltn,t\lrl1lJ puu jq .lrllanr\ Iorlu,!\ uo!.q[rid rt\ p.]o.llor s]lrrldJo pror.r V (f9BI) jl (curnoqt.t\ jotUrrd l^oD) I oN rrdrd i_rr:prun:rpr:,1ScSI ur rrtrrlsnV rltnos_.lo _ror.lurlsir\ rluou lt otur uorlrp.dxl s..trqqr rr\ SuDnl pil.illo. slurld irltJoUodi(6rgl) j r.ll.nl,\J '(ose.,rr,{rrdoitl) rruiTosrr s,js.i.u)tr,/,err./.lo sL,o,rrrndod ,,,,,,,,,,,.,"X1i,11:,,,ti;l',".,,i,"f;"1^.1'l]:l;.,"^ eruosof('qrz (.r..er,(qdot{7)ri.,,)t1;t snt,tqt.t.! la suo,rrndod u,,,",,,,,o,X|'flrll,i'i{,.1,"r,:.'i';".^:l;1lll),", ( LI!.d rt{ uortrd j,) .i.uir^\r.l IVJ) .. {.tr.qtur\ .rtl jo rjol.l.. l, )l.t !iti IA\ q s lnqt)J.(za)61) rt N -!at 0 :(t)Z crrr\.qurtZrrot:t ,/ iI..DItirldo.e,(Z(tqOI) I t.r:runc.t (r,IiqUt.):s(lDV)'IloNs.|J.\.runP:I'l]loI:Ie|ltl]snV..'slut|dlt|n.slrAUrr|ltl]slrVJosnstl:)J.'(66])}I.ln|IuFI ( ^utLtuiC:z1t:ro)) .lt1 ...po"l o.{)of rrnltt.uiLUoNIu.rflrNol] Jo irfo:l Iruortrrrilrl.. |t66i t;, ,,\\).]nrrD uirtllur-ttszurLId .ret;uJ 88:(t)t orqrrlrnleNer(I :tV),1]urr(l ii' VI) jq (068t) Vr) rittujt ( r\e) rsu.lrDD i)luelog jr.{ lD,\oI)!t-l ..n.l{tvtsrlp.rdor.tJorrot:t.,(pe),\U,llllo,t ar..r .illdo8(Z(S8a,l) Nl,\lplpoU l:l 'I 99-6S :6 rrtntoqlrinfrltntoqlrinf -I s)ttuqt.tJs)ttuqtlJ uortriLtrssctJ 99-6S lo uortriLtrssctJ.rlr ot uo|]rnpol]utuo|]rnpol]UtuV (8/6t) NhJNn tplppurl,rt)rU Ij (Drrlsnv:7uD) ecrurrl (p.) Ul-it:8a). erot-t lt\.r.;urqrilt ,/ ri.rr {Ltdir8,i7 (Z/61) Ni,\.!p!peU trl ll :at ou url.llnB ^.^rns rrntln.uivJo ( . It.rSolortl ldl)c it I I()l ur purtsturs,(E.t ol uorlrt:rd)i.uu-Io lrodil (Zl6t) v.\\ urrrg ]' U II ral rxol.ror pisr,\.x.. ..,oliiJ-Io tooqpurl (pi) ct\.rryicuussrc rrl e'..r1.i!rdoi-,(/ fr*n,l n ,ltl.-',l"',1",,J-*o 0ZI-t :(t)t ru. .nht (9681-SZ8l)r.lt:)nt\ puxurpr.:t.to . Ilf stro\\p.rtsrtqnd.r{.t. (tL6t) nC rro)urS,} O.t !rnr\'I I(1 rq3.rnrt:) .ocrUItl ( qur(t :s\ir(l V,,\\il).. Ll!o.t\urn}I Jo .iro.q.r,)..(6/6t) ,\\ eq,,qi.tJ la slou puu irl tlrrJl V r.)jllr8 I^j ll