New taxa, combinations and typifications in Verticordia (Myrtaceae : Chamelaucieae)

164 Pages·1991·50.1 MB·English

by  A S George

Nuytsia7(3):231-394(1991) 231
New taxa, combinations and typiflcations in
VerHcordia (Myrtaceae: Chamelaucieae)
A.S. George
AustralianBiologicalResourcesStudy,AustralianNationalParks&WildlifeService,
G.P.O.Box1383,Canberra,AustralianCapitalTerritory2601
Abstract
George,A.S. Newtaxa,combinationsandtypificationsinVerticordui(Myrtaceae:Chamelaucieae).Nuytsia7(3):231-394
(1991). ThehistoryofVerticordiaDC.issummarised. Themorphologyisreviewed,inparticulartheanthers,manyofwhich
areillustratedbyscanningelectronmicrographs. ThecytologicalfindingsofRye(1979)arereviewed. Thegenericstatusis
assessedandmaintainedinitsbroadsense. AnewinfragenericclassificationofthegenusVerticordiaisprovided,including
1newsubgenusand18newsections,while3namesaregivennewstatusatsectionallevel. 41newspecies,12newsubspecies
and 22new varieties are described; 3 previously publishednames are reinstated as accepted species. 7 names previously
publishedat specific rank arereduced tovarieties and 1 to subspecies. Darwinia verticordina F. Muell. is transferredto
Verticordia.Thegenusnowcontains3subgcnera,24sections,97speciesand42infraspecifictaxa.Keystoalltaxaareprovided.
Lectotypesandneotypesarechosoiforanumberofearliernames. Distributionmapsareprovidedforallnew species and
infraspecifictaxa.
Contents
Page
Abstract 231
Introduction 232
History 232
Methods 233
Morphology 234
Geographical distribution 251
Cytology 251
Hybrids 253
Furtherresearch 254
Verticordia 254
Synoptickey to infrageneric taxa 255
Key to infragcneric taxaandspecies 257
Infrageneric classification 269
Systematic infrageneric classification 282
The species 283
Acknowledgements 380
References 381
Maps 383
Index to names 391
232 NuytsiaVoL7,No.3(1991)
Introduction
Thepurpose ofthis paperis theconcisepresentation ofareview ofthe genus Verticordia DC.
(Myrtaceae:Chamelaucieae). Itcoversnewtaxaandcombinatifflis,lectotypificationsandneotypifications
and presents an infrageneric classification. Keys toall infrageneric taxa and species areprovided;
infraspecific taxaare keyedoutunderthe relevantspecies.
The papier will be followed by a book on the genus by Elizabeth A. George (formerly
E.A. Bemdt),inwhichalltaxawillbeillustratedincolour,andlateratreatmentbytheauthorinthe
A
“Flora ofAustralia”. Both thesepublications will contain descriptions ofall taxa. Verticordia
Reference Collection has also been compiled by Elizabeth George, with the assistance of many
collectors and observers. This contains representative spiecimens of almost all spiecies and
infraspiecific taxa, and is available for consultation in the Reference Collection of the Western
Australian Herbarium.
AsisthecasewithmanygeneraintheAustralianflora,Verticordiahasnotbeenrevisedsincethe
treatmentbyGeorgeBcntham (1867). Severaltaxahavebeendescribedsincethen, butithasbeen
obvioustoworkersinWesternAustraliathatsomenameshavebeenwronglyappliedandthatmany
newtaxaawaiteddescription. CompilationoftheVerticordiaReferenceCollectionhasaccentuated
theneed both forareviewand its earlypublication. Research on thecultivationofthesecolourful
plants is also increasingrapidly.
History
ThefirstspiecimensofplantslatertobenamedVerticordiawerecollectedatKingGeorgeSound
in October 1791 by Archibald Menzies, on the Vancouver expiedidon. These spiecimens, of
V.plumosa,arenowattheNaturalHistoryMuseum,London. Mostsubsequentcollectorsgathered
specimens ofVerticordia, butitwas notuntil 1826 thatthe first two spiecies were namedby Rene
Louiche Desfontaines,and then in thegenusChamelauciumwhich hehaddescribed in 1819. Two
yearslater, AugustinPyramusdeCandollerecognisedtheseasadistinctgenustowhichhegavethe
name Verticordia. Although he did not explain the etymology, it has generally been taken as a
referencetothe ancientgoddessVenusas the ‘turnerofhearts’ (e.g. Schauer 1840).
DiscoveryofnewspeciesbecamerapidaftertheSwanRivercolonywasfoundedin 1829. Many
spieciesweredescribedbyEuropieanbotanists(espieciallyJohnLindley, StephanEndlicher,Johann
Schauer, NicolasTurezaninow and CarlMeissner) from collections sentfrom thecolony. Lindley
published a new generic name, Chrysorhoe, for one spiecies but it was not maintained by other
workers. Onespeciesdescribedduringthisperiod,V.carinataTurez.,wasknownonlyfromthetypie
(Figure 15) until itsrediscovery in March 1990.
In 1859 the first spiecies to be described in AusU'alia were published by Ferdinand Mueller
(V. wilhelmiF. Muell.,published in 1855, is now placed in Homoranthus). By the time Bentham
(1867)treated thegenus in “Flora Australiensis”,37spiecieswereknown. Fournew spxjcieswere
described in thatwork,oneofwhich, V. harveyi (Figure23), has only recently been rediscovered.
Between then and the present papier, however, research has been piecemeal. Later in the 19th
Century,Muellerdescribedfourmorespiecies. SpiencerMooredescribedonein 1898,andLudwig
A.S.George,Verticordia(Myrtaceae:Chamelaucieae) 233
DielsandErnstPritzelseveralin 1904. Subsequently,onlysixfurtherspecieshavebeendescribed,
twobyCharlesGardnerin 1934and1943,onebyGardnerandAlexGeorgein 1963,onebyGeorge
in 1966, and two tropicalspeciesby NormanByrnes in 1977.
Green(1985)listed53speciesforWesternAustralia,allexcepttwoofwhichareheremaintained
atthatrank. TothesemaybeaddedV.decussataByrnesfromtheNorthernTerritory. Inthepresent
paper, 97 species are recognised, together with 42 infraspecific taxa. The large increase reflects
factorscommontoanumberofWesternAustraliangenera: superficialresemblanceofmanyspecies;
theexistenceoflocalisedtaxa,manydiscoveredwithin thepast30yearsasaccessibility improved;
and lackofresources toreview thegenus earlier.
Methods
Thisreview isbasedon astudyofthe gross morphologyofherbarium and, in manytaxa, fresh
material, as well as specimens preserved in formalin-acetic-alcohol. Measurements were taken
wherepossiblefromfresh,preservedordetergent-softenedspecimens. Mosttaxahavebeenstudied
in the field torecord habitand habitatdata, as well as flowercolourand perfume. The last was a
subjectiveassessmentmadeatthe time ofcollection - no nocturnal checks were made.
Becauseandroecialcharactersareusefulindistinguishingmanytaxabuttheanthersareofsmall
size, many wereexaminedandphotographedby meansofa scanningelectron microscope.
Material housed at20 herbaria was examined. Types ofall except twopreviously named taxa
were located; for those two the types are assumed lost and neotypes are nominated below.
PhotographsweretakenofmosttypesheetsandwillbelodgedintheWesternAustralianHerbarium
(PERTH).
Diagnosesare providedforallinfrageneric taxatogetherwith a listofincluded species. In the
treatment of species, only new taxa, new combinations and taxa needing lectotypification are
covered. Foreconomy,notreatmentisgivenofotherpreviouslypublishedtaxa. Thekeytospecies,
however, includes all sf)ecies, whileinfraspecific taxaare keyedout underthe species.
Foreaseofreferencespeciesaretreatedalphabetically. Thesynopsisofinfragenericclassification
(pp 282-283)providesa systematic sequence.
Mapsareprovidedshowingthedistributionofallnewtaxa.Localitieswereplottedbylatitudeand
longitude. These aretakenfromall herbarium collections seen,including those in theVerticordia
Reference Collection atPERTH. Vagueand general localities (e.g. Swan Rivercolony) have not
been mapped. Specific detailsoflocalitiesoftypecollections andrare taxa havebeen omitted, in
ordertoafford these populations someprotection.
The conservation status of all species and infraspecific taxa has been assessed and, where
appropriate, codedaccordingtothecriteria ofLeigh etal. (1981). Theassessmentis basedon the
numberofcollections,localitiesandtoalessextentestimatesofpopulation size. The lastfactoris
approximate,noaccuratecountshavingbeenmade. FullsurveyswillbecarriedoutbytheWestern
Australian DepartmentofConservation andLandManagement.
234 NuylsiaVol.7,No.3(1991)
Two infraspecific ranks are recognised here. Subspecies is used for those taxa that are - 1)
morphologically distinctbut not sufficiently so to be given specific rank, and - 2) geographically
disjunctfromtheothersubspecies. Varietyisusedwherethemorphologicaldistinctionislessclear-
cut (there areoften intermediates) andthe distributionsoverlap.
Latin diagnosesanddescriptions wereprepared using Steam (1966) as theprincipal reference.
Morphology
Habit. Most species are shmbs less than 2 m tall. The tropical species V. cunninghamii and
V. verticillatamaygrowtosmalltreesupto7mtall. Afewsouth-westernspeciessometimesexceed
2m,namely V. cooloomia, V.grandisandV. lepidophylla. Theshmbby specieshaveawiderange
ofform. The most common is a bushy shmb, openly to densely branched, sometimes rounded,
sometimescorymb-like,sometimesirregular. Thecorymb-likehabitmaybeslender,e.g. V. nitens,
V.cooloomia,orbroadandrobust,e.g.V.eriocephala. Severalirregularformshavecrowdedfoliage,
resulting in an arm-like growth, e.g. V. crebra, V. longistylis. Some species are widely spreading
ratherthanerect,e.g. V. huegelii,V.pulchella, V.penicillaris, V.oculata',theextremeisV.oxylepis
which hasamain stem usually less than 5 cm tall andasinglelayerofhorizontal branches.
Mostspecieshavea singlebasalstem andare fire-sensitive,beingkilledby fire. Regeneration
is from seed. Six speciesconsistentlydevelopasmalllignotuberwhichsproutsafterfire; theseare
V. brevifolia, V. coronata, V.grandis, V. oculata, V. ovalifoliaand V.pennigera. Inafurtherseven
species a small lignotuber occurs in some variants; these are V. chrysanthella, V. insignis,
V. densiflora, V. habrantha, V. huegelii, V. oxylepis and V. plumosa. The three tropical species
V. cunninghamii, V. decussata, V. verticillata-sproutfromboththe largerstemsandthebaseafter
fire. The lignotuber has evolved in relatively few taxaof Verticordia compared with some other
woody generain theAustralian flora, e.g. Banksia, Melaleuca. Itis thereforeofminor taxonomic
significancein thegenus.
Fire-sensitiveformsusuallygrowrapidlyfrom seedandareoftenattheirmostfloriferousstage
within five years.
Indumentum. In most species the vegetative parts are glabrous. A few have hispid stems, e.g.
V. staminosaand V. endlicheriana, whileseveral speciesofsect. Sigalantha, sect.Penicillarisand
sect. Verticordella haveciliate orerose leafmargins.
Many species have a pubescent, villous or silky hypanthium, and in some the petals, sepals,
androeciumandstylemaybevariouslyhairy. Thehairsarealwayssimpleexceptthoseofthestyle,
which may be branched. The style hairs are usually arranged in a beard on which the pollen is
depositedandpresented atanthesis. Indumentum terminology follows thatofHewson (1988).
Leaves. Theleavesaresimpleandsmall,usuallylessthan2cmlong,andsessileorshortlypetiolate.
In some taxathere isasmallpulvinusatthebasewhich may persiston thebranchletafterleaffall.
Thelaminarangesfromorbiculartonarrowlylinearandterete. Thearrangementisdecussateexcept
in V. verticillata (in whorlsof3 or4) and a few species in which they arescattered.
A.S.George,Verticordia(Myrtaceae:Chamelaucieae) 235
Mostspeciesofsubg.Chrysomahavetwoleafforms. Thoseoftheseedlingsandthelower,non-
flowering branches are linear and terete to semiterete or triquetrous, and those of the flowering
branches are broader and concave, e.g. V. subulala, or shorter and thicker, e.g. V. grandiflora.
V. cooloomiahasthreeforms, thoseofthemainstembeingintermediatebetweenthebasalandthe
floral leaves (Figure 17).
In speciesofsect. Chrysorhoe, subg. Verticordiaand subg. Eperephesthere islittledifference
(exceptsometimesinsize)betweenthelowervegetativeandupperfloralleaves(e.g.Figures21,23,
25).
A few species can be recognised from the gross leaf form and arrangement, e.g. V. serrata,
V. crebra, V. oculata, but usuallyfloralcharactersare neededfordetermination.
Inflorescence. Despite the impression ofcorymbs, racemes and spikes, all species ofVerticordia
&
have solitary, pedunculate axillary flowers. In the terminology of Briggs Johnson (1979),
Verticordiahasauniflorescenceconsistingofapedunculatemonadwith2prophyllsbutlackingan
anthopodium.Greatvariationinthelengthofthepeduncleandinthelengthoftheflower-bearingpart
ofthe branchletgivesrise to the appearanceoflargeinflorescences.
Inthosespeciesthatarecorymb-like,thefloralintemodesareshort,thelowestflowershavelong
peduncles and those above are progressively shorter (Figure 17). The branchlets may also be
similarlyarranged,givingthewholeplantacorymb-likeappearancewithallflowersofeachseason
at the same level, e.g. V. nitens, V. eriocephala, V. roei and V. endlicheriana (especially var.
compacta). Inothersthebranchingmaybelessregular,givingrisetoscatteredcorymb-likegroups,
e.g. V. plumosa, V. vicinella.
Thepedunclesmayalsobeofmoreevenlengthandtheintemodesshort,inwhichcasetheflower
groups arerounded, e.g. V. brachypoda.
Where the flowers have peduncles of even length the groups are raceme-like or spike-like
dependingonpedunclelength(e.g. Figure25). This isthecasein muchofsubg.Eperephesandin
V. mitchelliana, V. humilisand V. staminosa.
Theflowersmaybeerect,asinmostcorymb-likeforms,turnedsideways,asinmostraceme-and
spike-likeforms,andsometimesspreading(F.staminosa)orpendulous(F.mitchelliana,V. humilis).
In allspecies theprophyllsareoppositeandsessile, immediatelybelow the hypanthium. They
may be deciduous before or at anthesis or persistent. In several species they are united, e.g.
F. grandiflora, V.nobilis, V.rutilastra,F.chrysantha.Theyaresimilartothecheiridiumasdefined
in Calytrixby Craven (1987) butforconvenienceare termed bracteoles in thetreatmentbelow.
Flowers. TheVerticordiaflowerappearsinuicate,butmuchoftheintricacyderivesfromhavingthe
sepals deeply divided in various ways. Thepetals are usually lobed, fimbriateordentate, thereare
ten fertilestamens(fivein F.pictaand F. rennieana) alternating withten staminodes,a simplebut
often hairy style, and the hypanthium is often hairy and/or sculptured and, in subg. Eperephes,
bearing appendages. In all parts there is a large number of characters and states useful for
distinguishing taxa.
236 NuytsiaVol.7,No.3(1991)
Hypanthium. FollowingJohnson&Briggs(1984),thistermisusedinpreferencetoperigyniumand
thelessprecisefloraltube. The hypanthium is variously turbinateorhemisphericalandisoften 5-
or 10-ribbed or -angled. It is sometimes constricted at the apex (e.g. V. grandiflora) or below
(V. eriocephala, V. brownii). The surfacemay be smooth, rugose orhoneycombed.
In subg. Eperephes the hypanthium bears five reflexed appendages around its upper margin,
alternatewith thesepals(Figures9G & D, KKH). Theyaregreenand usuallyarecloselyappressed
tothehypanthium. AlthoughtermedherbaceousappendagesbyBenthamandlaterworkers,theyare
succulentorfirmin textureandareherereferredtosimplyasreflexedappendages. Thefunctionof
these appendages is unknown. Theirsizeand form areuseful in assisting todistinguish some taxa,
especiallyin sect. Verticordella.
Sepals. Thesepals(inallexceptonespecies,V.verticordina)aredividedintoanumberoflobesthat
are themselves plumose, fimbriate, erose ordivaricately divided. In many species they also have
recurvedorreflexedbasalauriclesorcilia(Figures9G, lOE),andinmostofsect.Pennuligerathere
arereflexedbasallobes. Therearefiveofthesereflexedlobes,butonesepalhastwo,threeeachhave
one,andthefifthhasnone. Theselobesareusuallyclawedandhavealaminathatispeltate,scale-
likeandciliatein somespecies,deeplyfimbriateinothers(andthedivisionsmaybeupturnedatthe
baseofthehypanthium).
Verticordiaverticordina,previouslyplacedinthegenusDarwinia,hasundividederosetoshortly
lacerate sepals butinmostmorphologicalfeatures is similarto the species ofsect.Infuscata.
In thespecies treatments, the lengthofthesepalismeasuredfrom thebaseofthe lamina to the
apexofthelongestfringeorlobesegment.Incountinglobesonlythewell-definedsegmentsaretaken
into account, notthe small lateralcilia presentin many species.
Petals. These may befreeor variously unitedwith theandroecium. Thelaminamay bedigitately
divided,fimbriate,dentateorerose,orrarelyentire(F.picta,V.interioris,V.habrantha,V. Integra).
In sect. Pennuligera and in V. rennieana eachpetal has twobasal auricles.
Theorientationofthepetalsvariesbetween species. They maybespreadhorizontallyorerect,
andinmanyspeciestheyspreadwidelyatanthesisbutlatercloseoverthecentreoftheflower. This
openingandclosingoccursovernightin some species,e.g. V. mitchellianaand V. capillaris,butin
others occursoverseveraldays,e.g. V. endlicheriana.
Thepetals aremeasured from thebase to theapex ofthe longest teeth, lobesorsegments.
Androecium. Allspecieshave10stamensand10staminodes,exceptV.pictaandV.rennieanawhich
have5fertilestamensand 15 staminodes. Asintheotherfloralparts therearemanycharactershere
useful for distinguishing taxa - form, size, orientation of filaments, anthers and staminodes. The
stamens and staminodes may be free or variously united (into prominently tubular form in
V. staminosa and V. monadelphd).
A.S.George,Verticordia(Myrtaceae:Chamelaucicae) 237
Figure 1. ScanningelectronmicrographsofVerticordiaanthers.A,B.-V.citrella(A.S.George16830).C-V.endlicheriarta
var. endlicheriana,withextrudedpollenpiled infrontofappendage(AS.George 16883). D.E- F.subulate (A.S.George
16506). F-H.- V.brevifoliasubsp.stirlingensis,Hafterextrudingpollen(A.S.George 16519). I-V.amphigia(A.S.George
16318). Scalebar= 100p(0.1 mm).
238 NuytsiaVol.7,No.3(1991)
Figure2. ScanningelectronmicrographsofVerticordiaanthers. A-V. Integra(AS.George 16468). B-F.serratavar.
serrata after extruding pollen (A.S. George 16461). C - V. rutilastra afterextruding pollen (A.S. George 16315). D,E-
V.coo/oomiu,Cstamenandstaminode(A.S'.George16843).F,G-V.aurea(A.S.George16359).Scalebar=100p(0.1mm).
AS.George,Verticordia(Myrtaceae:Chamelaucieae) 239
Figure3. ScanningelectronmicrographsofVerticordiaanthersandstaminodes. A,B-F.starrdnosavar.cylindraceaafter
extmdingpollen(A.S.George 16465).C,D-F.plumosavar.plumosa,Cstamenandstaminode(AS.George 16576).E,F-
V.plumosa var. grandiflora afterextmdingpollen (A.S. George 16498). G, H - V.stenopetala,G stamen and staminodes
(A.S.George 16437).I-V.vicinella(cultivatedCBG,31 March 1985,A.S.George).Scalebar=100p(0.1mm).
240 NuytsiaVol.7,No.3(1991)
Figure4. ScanningelectronmicrographsofVerlicordiaanthers.A-C-V,densifloravar.roseostella,Cafterextrudingpollen
(AS.George16835). D-F-V.eriocephala,Fafterextrudingpollen(AS.George16562). G-V.fastigiata(MonjebupRd,near
BoxwoodHill,D.Hutchinson,7April 1985).H,I-V.dasystylissubsp.kalbarriensisextrudingpollen(type,D.&B.Bellairs,
18October1987).J,K-V.oxylepis(A.S.George 16484).Scalebar= 100p(0.1 mm).