Taxonomic notes on Boronia species of north-western Australia, including a revision of the Boronia lanuginosa group (Boronia section Valvatae : Rutaceae)

46 Pages·1997·13.5 MB·

by  Marco F Duretto

,
Nuytsia 11(3):301-346(1997) 301
TaxonomicnotesonBoroniaspeciesofnorth-westernAustralia,
includingarevisionoftheBoronialanuginosagroup
(Boronia section Valvatae: Rutaceae)
MarcoF. Duretto
School of Botany, The University of Melbourne, Parkville, Victoria 3052, Australia
Address for correspondence: National Herbarium, Royal Botanic Gardens, Melbourne, Birdwood Avenue,
South Yarra 3141, Australia
Abstract
M.F.Duretto.TaxonomicnotesonBoroniaspeciesofnorth-westernAustralia,includingarevision
oftheBoronialanuginosagroup{Boroniasection Valvatae:Rutaceae).Nuytsia1 1(3):301-346(1997).
The revision ofBoronia (Rutaceae) in the Northern Territory, the Kimberley Region of Western
Australia, and north-western Queensland is completed, and a key to all species is provided. The
B. lanuginosaEndl.speciesgroupischaracterizedasthosespecieswithpinnateleaves,acalyxaslarge
asorlargerthanthecorolla,multiangularstellatehairs,andapronouncedridgeonthemicropylarside
oftheseed. ThiscladecomprisestheB. lanuginosaspeciescomplex, B.filicfoliaA.Cunn.exBenth.,
B.paucifloraW. Fitzg. and fivenewlydescribedspecies:viz. B. decumbens Duretto, B. minutipinna
Duretto, B. kalumburuensisDuretto, B.jucundaDurettoand B. toleransDuretto. TheB. lanuginosa
speciescomplexhasfouravailablenamesandwasanalysednumericallyusingpheneticmethods. Two
taxawereidentifiedintheanalysis. Boroniaartemisiifoliavar. wilsonii F. Muell.exBenth. israised
tospecificstatuswhile/},affinisR.Br.exBenth.andB.artemisiifoliaF.Muell.aresynonymizedunder
B. lanuginosa. Additionally,B.rupicolaDurettoisdescribed,andBfilicifolia, B.lanceolataF.Muell.
B. lanuginosaand B. paucifloraare lectotypified.
Introduction
This paperisthesecondtodiscussthesystematicsofBoroniaSm. (Rutaceae) forthe“Top End”
oftheNorthernTerritory(NT)andtheKimberleyRegionofWesternAustralia(WA). Thefirst,Duretto
&
Ladiges(1997),discussedtheB.grandisepalaF. Muell.group:thosespecieswithsimpleleavesand
sepalsaslargeasorlargerthanthepetals. Here,thepinnate-leavedtaxawithsepalsaslargeasorlarger
thanthepetalsareaddressed. Cladisticanalysis(Duretto 1995;Duretto&Ladigesinprep.)showsthat
thesetaxaformamonophyleticgroupcharacterizedbythepresenceofmultiangularstellatehairs,pinnate
leaves(exceptB.pauciflora,which sometimeshasjuvenileleavesthataretrifoliolate,buthassimple
mature leaves),sepalsas largeasorlargerthanthepetals, andshinyseedswith aprominentridgeon
m
the icropylarside. Westonetal. (1984)calledthisclade(lessB.pauciflora)theB. affinisgroup,but
forreasonsoutlinedbelow, it ismoreappropriateforittobeknown astheB. lanuginosagroup. This
cladebelongstoBoroniasection Valvatae(Benth.)Engl,andconsistsofthreespeciescomplexes:the
B. lanuginosaandB.fdicifoliacomplexes, plusanotherundescribed species complex.
;
302 Nuytsia Vol. 11, No. 3 (1997)
The B. lanuginosa species complex is found from King Sound (WA) to Wollogorang (NT)
(Figure 1)andconsistsofan uncertainnumberoftaxa. Morphologicalvariationwithinthecomplexis
analysednumerically,usingpheneticmethods. Leafanatomyandontogeny,andthetaxonomicstatus
ofthefouravailablenameswithinthecomplex(viz. B. afjinis.B. artemisiifolia,B. artemisiifoliavar.
wilsonii, and B. lanuginosa) are discussed.
TheB.JilicifoliaspeciescomplexisendemictotheKimberleyRegionandconsistsoffourspecies
thataremorphologicallyandgeographicallydistinct. Wheeler(1992)recognizedanumberofforms
withinB.filicifoliaincludingtheMitchellRiver(andtype)formandaKalumburuform B kalumburuensis,
seebelow). A third form, fromtheOsmond Plateau, is describedbelowasB. minu(ti.pinna. Boronia
paucifloraisalsoplacedinthisspeciescomplex(Duretto 1995;Duretto& Ladigesinprep.).
Thethirdspeciescomplexconsistsofthreerarespecies,B. decumbens(NT),B.jucunda(WA)and
B. lolerans(NT),thatarenewlydescribedbelow.
TocompletetherevisionofBoroniaintheNorthernTerritoryandtheKimberleyRegion,taxaare
lectotypifiedwherenecessary(includingB. lanceolatd)andB. rupicolaisdescribedasnew. Boronia
lanceolataandB. rupicolaaremorecloselyrelatedtoeastcoastmembersofBoroniasection Valvatae
thantootherspeciesinnorth-westernAustralia(Duretto1995;Duretto&Ladigesinprep.). Atthefinal
stagesofmanuscriptpreparationa1995collection(J.R.Clarkson 10473)fromnorth-westQueensland,
neartheNorthern Territoryborder, came to the authors’ attention. The specimen is ofapreviously
unknowntaxonthathasaffinitieswithB. alulataBenth. ofCapeYork(Durettosubmitted;Duretto&
Ladigesinprep.),andsowillbedescribedinaforthcomingpaperthatdealswithQueenslandmembers
ofBoroniasection Valvatae(Durettosubmitted). Asthisspecies issympatricwithB. lanceolata(in
Queensland)itwillbeincludedinthekeyasBoroniaaff.alulata(NWQld,Clarkson10473). AllBoronia
speciesfoundintheNorthernTerritory,theKimberleyRegion,andnorth-westQueenslandareplaced
in section Valvatae.
TheNorthernTerritoiyandKimberleyflorasaretreatedhereasasingleunitforanumberofreasons.
Firstly, apart from B. lanceolata and B. rupicola, cladistic analysis shows that all taxa form a
monophyleticgroupdefinedbythefollowingsynapomorphies:thecalyxbeingaslargeasorlargerthan
thecorolla, the antesepalous antherbeingsignificantly smallerthan theantepetalous anther, andthe
filamentshape(Duretto 1995; Duretto& Ladiges inprep.). Secondly,theboroniasofthisregion are
widelyseparatedfromtheeastcoast, South Australian, andsouth-westWestern Australian members
ofValvatae. Thirdly,andfinally,theregioncoveringthe“TopEnd”oftheNorthernTerritoryandthe
Kimberleyregionisoftenrecognizedasbeingadistinctbiogeographicregionoranumberofclosely
relatedregions;thatis,regionsthatsharearecenthistory(seeSpecht 1958b;Kikkawa&Pearse 1969;
Hnatiuk&Pedley1985;Cowie&Finlayson1986;Cracraft1991;Dunlop&Webb 1991;Crispetal, 1995
Duretto 1995;Duretto&Ladigesinprep.).
The boundaries for these biogeographic units, at least for Boronia roughly correspond to the
,
boundariesofthreegeologicalbasins(asoutlinedbyPlumb&Derrick 1975;Plumbetal. 1980;G.S.W.A.
1990):theMcArthurRiverBasinwhichextendsfromMtIsa(north-westQueensland)totheArnhemLand
Plateauandsurrounds(northernandwesternNT);theVictoriaRiverBasin,especiallythenorth-western
heavilyfaultedarea,whichincludesanareafromthewesternNorthernTerritorytotheOrdRiver(WA);
and,theKimberleyBasinincludingtheneighbouringKingLeopoldandHallsCreekorogensormobile
plates. These basins arecharacterized by Precambrian sandstones.
M. Duretto, Taxonomic notes on Boronia species of north-western Australia 303
Boronialanuginosaspeciescomplex
Endlicher(1837)describedB.lanuginosafrommaterialthatwascollectedbyFerdinandBauerand
labelledKingGeorge’sSound. KingGeorgeSoundisinsouth-westernAustralia,butBauerhadtravelled
withMatthewFlinders,onthe“Investigator”,aroundnorthern Australia(seeSpecht 1958a,b; Wilson
1975). Bentham(1863),whohadnotseenthetypematerialofB.lanuginosa,appliedthisnametoasouth-
westernWesternAustralianspeciesthatisfoundgrowingaroundKingGeorgeSound(Wilson 1975).
Later,Mueller(1859)describedB.artemisiifoIia(asB.artemisfolia)frommaterialhecollectedwhileon
Gregory’s Northern Australian Expedition. Bentham (1863) used this later name (written as
B.artemisiaefolia)inhis“FloraAustraliensis”. Subsequently,andupuntil 1975,specimensofBoronia
fromnorthernAustraliathathadadenseindumentum, pinnateleavesandalargecalyxhavebeencalled
B. artemisiifolia, with orthographicvariation. Afterstudyingtypematerial ofS. lanuginosa, Wilson
(1975)realizedBentham’serrorandsynonymizedS. artemisiifoliawithB. lanuginosa. Fieconcluded
thatthetypelocalityof#/cwugmo.sawassomewhereonthecoastofArnhemLandwhereFlindcrs'ship
“TheInvestigator”hadtravelledonitssecondvoyage. RobertBrown,whowason“TheInvestigator”
withBauerandFlinders,collectedplantspecimensfromtheislandsoftheGulfofCarpentariathatare
identicaltothetypematerialofB.lanuginosa{Wilsonpers.comm.),strengtheningWilson’sargument.
The south-western Western Australian species thathadpreviously been calledB. lanuginosais now
knownasB.strictaBartl.(Wilson 1975).
Bentham (1863) described B. affinis, B.filicifolia and B. artemisiifolia var. wilsonii (presently
synonymizedunderB. lanuginosa)fromnorth-westernAustralia. ThetypematerialofB. affinis(“N.
Australia. IslandsoftheGulfofCarpentaria,andmainlandoppositeGrooteEylandf wascollectedby
’)
RobertBrownwhiletravellingwith Matthew Flindersand Ferdinand Baueron“The Investigator” in
1802-03. ThemainfeaturethathasbeenusedtodistinguishB.affinisfromIIlanuginosaisthatB.affinis
isglabrousorhasasparse indumentumwhileB. lanuginosahasadenseindumentum. Thereismuch
confusion surroundingthe application ofthesetwo names.
Judgingfrompresentherbariumrecordsandcollections,B.affinisandB. lanuginosaaresympatric
intheNorthernTerritory. Boroniaaffinisisconsideredtoberare,possiblyendangered,andconfined
totheNorthernTerritory(Briggs*Leigh 1988[notlistedinBriggs&Leigh 1996];Hnatiuk1990)while
B. lanuginosa (as currently circumscribed) is common and widespread from King Sound (WA) to
WollogorangStation(NT)(Figure 1).
Boronia lanuginosa has an ontogenetic sequence in leafdevelopment from glabrous leaves, or
leaveswithasparsesimpleandstellateindumentum,toleaveshavingadensestellateindumentum,as
4hS4aos5m5be,eesnpWeoicbgishmetermnvasend(ei.n1g3o.7tDh4uenr&lmoeCpmrb5ae3v8re0sn;;oDfWusorelecfttetioo&n50VM0aa;lrvGtaeitonare2g(2eD7u)1r3edt5it4so8p;l19lal9yc5nt;hsDihsuarloelntt1t6oo8g&0e;nLeKateidncingeseaeslq1lu9ey9n37c0,e2.i5n,pM3re0ap7n.5)y^.
populationscontain avarietyoffertileformsthatdisplayvaryingdegreesofhirsuteness(e.g. Duretto
477-481;488-491;503-504A;522-524).Otherpopulations,suchasthoseintheMtCahillandMtBasedow
areas(KakaduNationalPark),NorthIsland(GulfofCarpentaria)andthesandplainsnorthoftheArnhem
LandPlateau,consistofsmallplantsthatareglabrousorhaveasparseindumentumandthatareoften
determinedas B. affinis. Interestingly, arangeofhairdensities, which is usuallycorrelated withthe
densityofhairsontheleaves, isalsopresentontheabaxialsurfaceoftheperianth parts. Thishas led
tosometaxonomicconfusion in thepast.
AlongwiththisindumentumvariationB.lanuginosashowssomemorphologicalandfloralvariation
acrossitsrange. PlantscollectedfromthenorthernandeasternareasoftheNorthernTerritory(including
304 Nuytsia Vol. 11, No. 3 (1997)
Figure 1. North-western Australia. Specimens used in analysis ofB. lanuginosa species complex (25, 42-44, 54, 63,
64, 70); B. lanuginosa. Group A (O); B. wilsonii, Group B ().
theislandsoftheGulfofCarpentaria)havelinear,revolutepinnae. ThosecollectedsouthofUDPFalls
(KakaduNationalPark)toNitmilukNationalPark(NT)andwesttotheWeaberRange(WA)aresimilar
butoftenhavelargerflowers. Kimberleyplantshavebroad,recurvedpinnae. Morphologicalvariation
withinthecomplex isanalysednumericallybelow,usingpheneticmethods.
Materialsandmethods
Material
HerbariumspecimensweremadeavailablefromAD,BR1,CANB,CBG,DNA,JCU,MBA,MEL,
MELU,NSW,PERTH,QRS,TCDandWAU,cibachromesandslideswerereceivedfromK,andslides
werereceivedfromBM. HerbariumabbreviationsfollowHolmgrenetal.(1990). Thesespecimenswere
augmentedwithspecimensfromOSS(OfficeoftheSupervisoryScientist,KakaduNationalPark,NT)
andmaterialcollectedinthefieldduringJuneandJulyof1993.
Leafanatomy
Thecentralportionoftheleavesofalltaxawassectioned. MaterialwasfixedinMirsky’sfixative
(MAA)or70%ethanol. lffreshmaterialwasnotavailable,herbariumsampleswerere-hydratedbybeing
placedinwaterwithasmallamountofdetergent,broughttothebrinkofboiling,leftsimmeringforone
hourandsoakedovernightbeforefixinginMAA. AIIfixedmaterialwasplacedin70%ethanolovernight,
dehydratedthroughagradedethanolseriesupto 100%ethanol,infiltratedwith100%LR-White(London
Resin)througharesin/ethanolseries,andpolymerizedat60°C. Sections,2 pm inthickness,werecut
onaReichertUltracutultra-microtome,stainedwith0.05%toluidinebluesolution(pH4.4)andobserved
andphotographedusinganOlympusBHScompoundmicroscope. Anatomicalfeaturesaredescribed
inthetaxonomicdescriptions.
1
M. Duretto, Taxonomic notes on Boronia species of north-western Australia 305
Scanningelectronmicroscopy
Trichomes(ofleavesandstems)andseedsurfacesofalltaxa(wherematerialwasavailable)were
surveyedusingaScanningElectron Microscope. Dryleaves,stemsandseedsweremountedon stubs
usingdouble-sidedorcarbon tapewith conductivecarbon paint,coatedwith gold usingan Edwards
SputterCoaterSI50Bandexaminedandphotographedat5KVusingaJEOL840 ScanningElectron
Microscopeequippedwithalanthanumhexaboridefilament. Allphotographsofseedsweretakenof
centralareasonalateralside,exceptwhereotherwisestated. Trichomeandseedcharactersaredescribed
inthetaxonomicdescriptions.
Pheneticanalyses
Characters
Seventeencharacters(Table1)werescoredfor105herbariumspecimens(Tables2,3)thatcoverthe
entiregeographicrangeofthe B. lanuginosaspeciescomplex. Somecollectionswere includedmore
thanoncebecausetheyconsistedofanumberofplantswithvaryingdegreesofhirsuteness:viz. Craven
6705(specimens32and104),Wightman1337(specimens71and72),andRussell-Smith286 (specimens
1
81-83). Scoresareanaverageoffivemeasurements(wherefiveorganswereavailable)andratiosarethe
Table 1. Morphological charactersused inscoringherbarium specimens.
Binarycharacters
1. Style glabrous/hirsute,0/1
2. Leavesrecurved/revolute,0/
Numericcharacters
3. Lengthofpetiole(mm)
4. Maximumleafletnumber
5. Lengthofmostproximalrhachissegment(mm)
6. Lengthofmostdistantrhachissegment(mm)
7. Lengthofterminalleaflet(TLL)(mm)
8. Widthofterminalleaflet(TLW)(mm)
TLWATLL
9.
10. Lengthoflateralleafletprecedingtheterminalleaflet(LLL)(mm)
11. LLLvTLL
12. Lengthofpeduncle(mm)
13. Lengthofanthopodium(mm)
14. Lengthofsepal (SL)(mm)
15. Lengthofpetal(PL)(mm)
16. PL/SL
17. Lengthofsepalonflowerwithmaturefruit(mm)
1 1 1 s 1 8
306
Nuytsia Vol. 11, No. 3 (1997)
Table2. Datausedin analysisofB. lanuginosaspeciescomplex. Principalcollectorgivenonly. For
quantitativecharactersmeanvaluesaregiven (seeTable 1).
Specimen Collector Herbarium Character
21 WCirla&svoennnum13b27e68r9(ordate) DD&NNAAs54h5e9e19t041n5umber 001 211 00..368 77..400 31..150 34..600 89..758 21..807 00..12849 681..002 00..871331 011..250 321..300 571..400 351..500 00..716160 71.07?
543 THKinhniozgmhs1t1o01n4520474 ODDNNNAAA3433349426465934 000 111 001...553 1197...000 421...907 333...625 11019...20 212...112 000...211092 677...666 000...776347 001...055 234...000 566...000 444...000 000...866077 78..00?
6 Craven5963 CANB313896 0 1 o.s 9.0 1.5 2.3 8.B 1.2 0.15 7.6 0.37 0.0 2.0 6.0 4.0 0.67 8.0
7 Russell-Smitfi5220 DNA47609 0 1 0.5 9.0 1.5 3.4 9.2 1.0 0.11 5.0 0.55 0.0 2.0 6.0 4.0 0.67 8.0
3 Wigntman4283 DNA50903 0 1 0.5 13.0 3.2 2.8 11.3 1.6 0.14 9.4 0.81 0.0 3.0 8.0 5.0 0.63 7
9 Lazarides7928 DNA52723 0 1 0.7 13.0 1.7 3.0 10.8 1.1 0.11 7.6 0.71 0.0 2.0 5.0 4.0 0.80 8.0
10 Cleminson261 DNA79417 0 1 0.5 9.0 2.0 3.0 10.6 1.5 0.14 9.2 0.86 0.0 2.5 6.0 4.0 0.67 8.0
1 Hinz196 0NA43177 0 1 0.5 7.0 2.2 2.8 10.4 1.0 0.10 7.6 0.73 0.0 2.0 6.0 5.0 0.83 ?
12 Wigntman 1374 CANB3S25I6 0 1 1.0 13.0 1.0 3.6 7.4 1.0 0.14 4.2 0,57 0.0 5.0 7.0 5.0 0.71 9.0
13 Craven2299 CANB271721 0 1 0.6 7.0 1.7 3.0 10.2 1.0 0.10 8.4 0.35 0.0 5.0 7.0 5.0 0.71 12.0
14 Lazarides9042 DNA19608 0 1 0.9 9.0 1.4 2.7 11.0 1.1 0.10 6.2 0,56 0,0 4.0 7.0 5.0 0.71 9.0
15 Hinz26 DNA32735 0 1 1.3 17.0 2.4 3.2 8.2 1.1 0.19 4.0 0.66 0.0 1.0 5.0 4.0 0.80 6.0
16 Clark1344 DNA34728 0 1 0.9 11.0 3.4 6.0 8.2 1.7 0.21 6.4 0,79 0.0 2.0 7.0 4.0 0.57 9.0
1137 HPeanrskhearl9l061680 DDNNAA5141397084 00 11 01..83 1151..00 22..42 23..02 88..04 11..20 00..1162 55..26 00..6666 01..50 32..00 6s..0o 44..00 00..6677 88..00
19 Wigntman 1615 DNA23559 0 1 2.0 11.0 4.0 4.0 13.2 1.7 0.14 9.0 0.68 1.5 2.0 6.0 5.0 0.83 7.0
20 Maconochie860 DNA85534 0 1 1.3 9.0 2.3 3.2 9.8 1.8 0.17 a.o 0.83 0.0 1.5 5.0 4.0 0.30 ?
21 Pullen 9233 DNA315385 0 1 0.8 9.0 2.4 3.0 7.4 1.5 0.22 4.8 0.66 0.0 3.0 5.0 4.0 0.80 7.0
22 JacoDs 1587 NSW244407 0 1 1.0 11.0 3.4 3.6 9.3 1.6 0.18 8.4 0.71 1.0 3.0 6.0 5.0 0.83 8.5
23 Halford34114 DNA76573 0 1 1.1 15.0 2.8 3.5 6.8 1.3 0.20 5.2 0.76 1.0 1.5 5.0 4.0 0.80 ?
24 Maconochie2096 DNA48271 0 1 0.8 11.0 1.2 3.7 72 1.3 0.18 4.4 0.61 0.0 4.0 5.0 4.0 0.80 7.0
25 Reichenbach 3.vii.1955DNA40515 0 1 0.8 13.0 1.8 3.0 8.0 1.4 0.18 5.4 0.68 0.0 3.0 6.0 5.0 0.83 7.0
26 Gallon53 DNA28469 0 1 0.5 13.0 2.3 3.4 10.0 2.1 0.23 7.0 0.72 1.0 1.0 7.0 S.O 0.71 7.0
27 Hartley 13828 0NA44421 0 1 0.7 11.0 1.8 3.6 9.2 1.8 0.20 6.0 0.66 1.0 2.0 5.0 4.0 0.80 ?
28 Chappill30.vi.1993 MEL 0 1 1.3 9.0 1.0 3.1 9.5 1.0 0.IQ 6.0 0.63 0.5 3.3 6.0 4.0 0.67 8.5
29 Leach 2757 BRI AQ4S26I3 0 1 0.7 9.0 1.6 1.8 9.8 1.1 0.11 5.8 0.59 0.0 2.0 5.0 4.0 o.ao 7.0
30 Menkhorst357 MEL1582546 0 1 1.5 9.0 3.4 3.4 7.5 1.2 0.18 4.4 0.58 0.5 1.5 6.0 5.0 0.83 7.0
31 Cowie 1165 MEL1582645 0 1 0.5 9.0 2.1 5.0 12.3 0.3 0.06 8.6 0.70 0.5 2.0 6.0 4.0 0.67 7.0
3332 KFriynxgel2l948222 ADDN9A829227500 00 11 00..55 1111..00 01..90 22..36 109..23 11..00 00..110 87.04 00..8669 00..00 44..00 1100..00 38..00 00..3300 1121..00
3354 CFrryaxveleln48970075 MDNEAL357276223339 00 11 00..53 1113..00 01..19 33..64 167..08 01..80 00..1017 13501 00..6862 00..05 42..00 150..00 46..00 00..8600 1110..00
333678 SKKiiynnmgge4s22.V8I8.1982 DDDNNNAAA12203432444587 000 111 000...644 111753...000 011...672 324...404 11271...024 111...000 000...101084 775..433 000...866168 000...000 222...000 667...050 455...000 000...776177 111300...000
39 Barlow558 ONA20750 0 1 0.3 9.0 0.8 2.3 8.0 1.0 0.13 6.2 0.79 0,0 2.0 7.0 5.0 0.71 11.0
40 Dunlop 8086 DNA42635 0 1 0.5 11.0 1.4 1.4 7,4 1.0 0.14 6.6 0.90 0.0 4.0 70 3.0 0.86 12.0
41 Lazarides8426 DNA56830 ? 1 0.5 15.0 0.5 1.9 5.2 0.8 0.15 4.3 0.94 1.0 2.0 7.0 6.0 0.86 8.0
555554455444444263459805346721 lCrFFDKKWGCLDFFihrrrrlaeaeoueulyyeyyaznnarnnolxxxxsrannibclleereetksreeellnloloegiaallllspr4p3edllf213444e4lli.263580s6671yye0v3929080l3.5689118d52905197063649921888949245 MCDBMCDCCBCDDDDRRNAAEAAENNANNNIIANNNNALNAALAA38B2BB3284111A1A330331722556Q4776Q0367753734S47456677544421195062064S2844893S933441266207614548 001110001110011 000000000000011 222222213013011...............120554600980522 1111111111111553593333913951...............000000000000000 322235355443211...............802880444006062 324443462431231...............486822322009828 1111111823458091569815...,...........838044888440248 433542343322111...............510060812880900 000000000000000..,............434522324323411160387204635083 6337686a6354556.......,..-....00436666040602 000000000000000.....,.........775865548764668105521078006326 00000000000000o...............0000000000000s0 243433535334244..,.,.........,005000000000000 575775796966595...............000000000000000 366454467444648............,..000000000000000 000000000001000...............685888768660868067606807077970 11168778817907718..............00000000000000?
averageoftheindividualratiosofthefiveorgansmeasured. Therearesomeproblemsassociatedwith
theuseofratiosinpheneticanalyses(seeDuretto&Ladiges1997andreferencestherein fordiscussion):
hereratiosareusedasameansofquantifyingandstandardizingleafshape(characters9and 1 1), and
measuringthe relative lengthsofthesepalsandpetals(character 16).
1
M. Duretto, Taxonomic notes on Boronia species of north-western Australia 307
Table3. DatausedinthepheneticanalysisoftheB. lanuginosaspeciescomplex, analysis3,juvenile
leavedplantsonly. Principal collectorgivenonly. Forquantitativecharactersmeanvaluesaregiven
(seeTable 1).
Specimen Collector Herbarium Character
&number(ordate) &sheetnumber 14 15 16 17
57 Craven3480 DNA55102 6.0 5.0 0.83 7
58 Latz 10096 DNA 7.0 6.0 0.86 9.0
59 Craven5796 DNA19609 6.0 5.0 0.83 7
60 Tidemann 13 DNA51300 6.0 4.0 0.67 7
61 Russell-Smith3072 DNA43330 4.0 3.0 0.75 6.0
62 Must1041 DNA4995 4.0 3.0 0.75 6.0
63 Grinns3 DNA82012 6.0 4.0 0.67 8.0
64 Craven631 DNAS19618 6.0 5.0 0.83 6.0
65 Wightman2229 DNA26585 6.0 4.0 0.67 7
66 Clark1363 DNA49016 7 7 7 6.0
67 Russell-Smith2766 DNA30067 6.0 5.0 0.84 8.0
68 Smith579 DNA39890 6.0 5.0 0.84 7.0
69 Clark1178 DNA34488 6.0 5.0 0.84 7.0
70 Cowie26.xl.1985 DNA48810 5.0 4.0 0.80 7
71 Wightman 1337 NSW244394 4.0 3.0 0.75 5.0
72 Wightman 1337 NSW244394 4.0 3.0 0.75 5.0
73 Wightman647 DNA21610 5.0 4.0 0.80 5.0
74 Clark1529 DNA34617 5.0 4.0 0.80 6.0
75 Symon 7736 DNA67794 5.0 3.0 0.60 6.0
76 Maconochie 1482 DNA35810 6.0 4.0 0.67 8.0
77 Must 1018 DNA4890 5.0 4.O'0.80 6.5
78 Wightman 1981 DNA26336 6.0 5.0 0.83 9.0
79 Bowman308 DNA26280 7.0 4.0 0.57 7.0
80 Wightman 1084 DNA22555 5.5 4.0 0.73 8.0
81 Russell-Smith2861 DNA29879 6.0 5.0 0.84 7
82 Russell-Smith2861 DNA29879 5.0 4.0 0.80 7.0
83 Russell-Smith2861 DNA29879 5.0 4.0 0.80 7.0
84 Wightman 1130 DNA22554 5.0 4.0 0.80 7
85 Hartley7.vi.1974 DNA8233 5.0 5.0 1.00 5.0
86 Wightman3798 DNA30364 5.0 5.0 1.00 5.0
87 Latz3462 DNA36986 5.0 3.0 0.60 8.0
88 Menkhorst337 DNA43605 5.0 4.0 0.80 6.0
89 Craven6063 CANB313897 6.0 4.0 0.67 7
90 Muir5976 AD98904071 6.0 4.0 0.67 7
91 Muir6057 AD9890623 5.0 4.0 0.80 5.5
92 MartenszAE583 BRI [AQ151157] 10.0 5.0 0.50 12.0
93 Cowie272 DNA27078 10.0 7.5 0.75 7
94 Gittens2612 NSW244427 12.0 9.0 0.75 13.0
95 Henry896 DNA49279 12.0 8.0 0.67 7
96 Bowman383 DNA37156 6.0 5.0 0.83 7.0
97 Ollerenshaw 1594A DNA50113 6.0 4.0 0.67 7.0
98 Lazarides8944 DNA19605 6.0 4.0 0.67 9.0
99 Brown9.vii.1985 DNA26772 5.0 4.0 0.80 7
100 Leach2829 DNA47633 7.0 5.0 0.71 8.0
101 King 16.vl.1981 DNA18842 7.0 5.0 0.71 7
102 Pullen 10602 CANB264130 8.0 5.0 0.63 9.0
103 Brooker3149 CANB259815 11.0 8.0 0.73 15.0
104 Craven6705 DNA20963 8.0 6.0 0.75 7
105 King45 DNA20464 9.0 6.0 0.67 10.0
Fornumericalanalyses(cladisticorphenetic)homologousfeaturesonlyshouldbecompared. For
example, comparing leafmeasurements at different stages ofdevelopmentwould be erroneous and
resultswouldbemisleading. Also,asthereisanontogeneticsequencefromglabroustohirsuteplants
in the B. lanuginosa species complex, characters such as hair density are unusable. The phenetic
analysis,outlinedbelow,usesmorphologicaldatafrommatureplants(i.e.withadenseindumentum)only
and,floraldatafromspecimenswithmatureand/orjuvenilefoliage.
308 Nuytsia Vol. 11, No. 3 (1997)
Dataanalysis
AlldatasetswereanalysedusingPATN(Belbin 1987)followingthemethodologyoutlinedinDuretto
(1995)andDuretto& Ladiges(1997). Datawererange-standardizedbeforeManhattandissimilarity
measureswerecalculated Forclusteranalysis,bothflexibleUPGMA(unweightedpairgrouparithmetic
W.PGMA
averages) and flexible (weighted pair group arithmetic averages) were utilized as fusion
strategies. Datawereordinatedinthreedimensionsusingthemultidimensional scaling,MDS, KYSP
algorithm(Kruskaletal. 1973). TheHybridoptionofFaithetal. (1987)waschosen. Twentydifferent
randomstartingpointswereusedforeachanalysisandtherunwith the loweststressvalue isshown.
Charactercorrelations withtheordinationvectorswerecalculated usingthe PCC function ofPATN.
Minimumspanningtrees, MST,werealsocalculated. Threeanalyseswerecompleted.
Taxondescriptions
Descriptiveterminology follows Theobald etal. (1979) and Hewson (1988) forhairs; Briggs &
Johnson(1979)andWeston(1990)forinflorescencestructure;andMurley(1951),Powell&Armstrong
&
(1980),Barthlott(1984)andDuretto Ladiges(1997)forseedsurfaces. Conservationcodesfollowthe
format of Briggs & Leigh (1996) for all taxa, and that ofthe Western Australian Department of
ConservationandLandManagementforWesternAustraliantaxa(Nuytsia10,p.471, 1996). Authority
abbreviationsareasgiveninBrummitt&Powell(1992).
Specimencitation
In order to accord with the policy of the journal, specimen citations have been deliberately
abbreviatedtoachieve lessprecision inordertoprotecttaxawithconservationcodesofE,VorK. A
complete listofspecimensseen isavailablefromtheauthoronrequest.
Resultsofpheneticanalysis
Analysis (specimens 1-56,characters 1-14)
1
Thefirstanalysisincludesplantswithmature(havingadenseindumentum)foliageonly,specimens
1-56,andcharacters1-14. Twogroups,AandB,arerecognizableintheUPGMA(Figure2)andWPGMA
(notshown)classifications,ordination(Figure3),andMST(Figure4). GroupAincludesallspecimens
fromtheWcaberRange(WA)to“Wollogorang”(NT)(specimens1-43)exceptspecimen54(Figure 1).
GroupBincludesallKimberleyspecimenswestoftheOrdRiver(specimens44-53,55,56)andspecimen
54fromthelowerVictoriaRiverarea(NT). IntheMST(Figure4),GroupsAandBarelinkedbyspecimens
25(and 17, 18,21 etc)and44(and45,56etc),whicharefromVanderlinIsland(GulfofCarpentaria)and
BougainvillePeninsula(KimberleyRegion)respectively(Figure1).
Charactersthatarehighlycorrelatedwiththevectors intheordination(Figure3)are:characters2,
3,8, 1 1 forvectorl; l,4,7and 10forvector2;and, 12forvector3(notshown). GroupAischaracterized
bysmallpetioles(character3),fewerpinnae(character4),andpinnaethatareshort(character7),narrow
(characters8and9)andrevolute(character2). GroupBischaracterizedbylongerpetioles,andlonger,
broaderandrecurvedpinnae(Table4).
M. Duretto, Taxonomic notes on Boronia species of north-western Australia 309
Figure 2. UPGMA classification, analysis 1, specimens 1-56.
II
O
Figure 3. Ordination (KYSP), vectors 1 and 2, analysis 1, specimens 1-56. Boronia lanuginosa, Group A (<>);
B. wilsonii, Group B (^). Specimens referred to in text are numbered.
90 08
310 Nuytsia Vol. 11, No. 3 (1997)
Table4. CharacterrangesforGroupsAandB. Meansinbrackets.
Character GroupA GroupB
1 Style glabrous/hirsute 0 0(38%)-1(62%)
2 Leaves recurved/revolute 0
1
3 Lengthofpetiole(mm) 0.2-2.0 1.2-3.
(0.74) (2.2)
4 Maximumleafletnumber 7-17 9-19
(11.1) (13)
5 Lengthofmostproximal 0.5-4. 1.6-5.4
rhachissegment(mm) (1.92) (3.5)
6 Length ofmostdistant 1.4-6. 1.8-6.2
rhachissegment(mm) 3.2 3.6
7 Lengthofterminalleaflet 5.2-16.0 4.0-18.8
(TLL)(mm) (9.3) (10.6)
8 Widthofterminalleaflet 0.8-2.2 1.9-5.
(TLW)(mm) (1.3) (3.4)
9 TLW/TLL 0.06-0.24 0.2-0.53
(0.15) (0.35)
10 Lengthoflateralleaflet 4.0-13.0 3.2-8.8
precedingtheterminal (6.7) (6.2)
leaflet(LLL)(mm)
11 LLI/TLL 0.55-0.94 0.46-0.85
(0.73) (0.61)
12 Lengthofpeduncle(mm) 0.0-1.5 0.0
(0.33)
13 Lengthofanthopodium(mm) 1.0-5.0 2.0-5.0
(2.65) (3.6)
14 Lengthofsepal(SL)(mm) 5.0-10.0 5.0-9.0
(6.36) (6.46)
Analysis2(specimens1-43,characters3,5-7, 14)
Twogroups,AandB,wererecognizedinAnalysis 1. GroupAwasre-analysed,thatisspecimens
1-43,andcharacters3,5-7and14toseewhethermoregroupswerediscernible. Uniformcharacterswere
deletedpriortore-analysis.
Intheordinationplotofaxes 1 and2(Figure5)specimensfromKatherinearea,westernNorthern
TerritoryandtheWeaberRangelooselyclustertogetherbutarecontinuouswiththeotherspecimens.
NodistinctgroupsarediscernibleineitheroftheUPGMAorWPGMAclassifications(notshown)or
MST(notshown). Charactersthatarehighlycorrelatedwiththevectors(Figure5)are:3and5forvector
1;6and 16forvector2;and,7forvector3 (notshown).
Analysis3(specimens 1-43,57-105,characters 14-17)
The third analysis involvedGroup A andsympatric specimenswithjuvenile foliage, specimens
57-105,andfloralcharacters(14-17)only. Twogroups,A1andA2,arerecognizableinboththeUPGMA