WESTERN AUSTMLIAN HERBARIUM RESEARCH NOTES

N o . 5 , 1 9 8 1 : 2 9 - 4 8

THE PHYSICAL ENVIRONMENT. FLORISTICS AND PHENOLOGY OF ABANKSIA WOODLAND NEAR PERTH, WESTERN AUSTRALIA

By A.V, Mi lewski and Christ ine Davidge

Environmental & Life Sciences, Murdoch University, Ifurdoch, 6153,Western Austral i -a.

ABSTRACT

Ihe phgsicaL enui.r'orlnent and flot"Lstic cornposition of 10 haof mature, tu.tdLs tutbed Banksia uoodlmtd on deep" Leached sand axedes cr"Lbed. The s l;udg a.z:ea cotseyed part of a siLiceous &,Lne undera medLterranean climate on the s outhern outskirts of Perth"Westetw Austz,alia. The 6 m high tree s tratum consisted rainly ofBanksia attenuata (Proteaceae). The L n high shrub stt:atumcornpr"Lsed g0 perenni.al speaLes in 25 fantlies. SLight uctt'iationsin the conposition of both stztata uere found anong lour s tud4sites in assoctatton uith small dLfferences in strbsty,ate, TheperLod of mat:imwn flrn:erLng uas October" aLthough a feu speciesfLoaered i.n uinter on in sunrnez,. The times of crppearance of anaddLtiornl 25 species of annuaLs and herbs, apparent onLy inuinter and sprf,lry, one z:e coz,d.e d.

INTRODUCTION

This paper presents data on the physical environnent, floriqtics and ^phenology of an area of Banksia woodland at Jandakot Ai rpor t (32005'S, 1 l5o53rE) on the souther :n outsk i r ts of the Per th netropol i tan area, l l lestemAustralia. The nain aim of the study was to provide environnental andfloristic data as a counterpart of an intensj-ve study on the reptile, frogand b i rd comrnur i t ies of th is area (Davidge L979a, 1,979b, Mi lewski unpubl ished) ,These data arso provide a detailed description of an AustraLi-an nediterraneanarea, It is intended that this description will permit future comparisonsof plant and aninal commr.mities in Australia with those of other continentswi th a s in i lar nedi tenanean c l inate,

DATA COLLECTI ON

An area was needed which was large enough to include all plant andanimal menbers of a natural comnunity but which, at the same tine, includedonly one such community, A study area of 10 ha was chosen on the basis ofthe apparent gross honogeneity of the environrnent ard vegetation, the rnaturityof the vegetation and a mininun of nan-nade disturbance or invasion by weedy

30

annuals. Figure 1. isthe vegetat ion.

A . V . M i l e w s k i

an aerial view

& C. Davidge

of the area and

N o . 5

figure 2 i 11ust rates

Fig. 1. Aerial view of Jandakot Airport., showing location of study area(arrowedJ.

Fifty two sanple plots, 2 x I m, were placed within a prinary studysi te of t ha (corresponding to the s i te used by Davidge 1979a, ard R1 ofDavidge 1979b), using an 8 x 8 m grid. The first pair of nurnbers from atable of randon numbers gave the co-ordinate of a point on the grid, Twornore pairs of numbers gave the direction and distance to walk frorn this pointto the f i rs t p l "ot . The process was-repeated to locate the other p lots . Thearea assessed in th is way was 100 nz of 1% or the tota l area of the s i te ,

Al l shrub species (perennia ls lower than 3 n) , present as l iv ing p lantsin each plot, were recorded between February and May 1978. The foliagepro ject ive cover of each species was est imated for each hal f o f each pLot .Mean fo l iage pro ject ive cover was then calcu lated for a l l species in theprimary site. The site was exarnined thoroughly for species undetected by

N o . 5 Phenology of BanksLa Woodland

Fj.g. 2. General view of the vegetation in the study area, showing the upperstratun of low trees of Banksi.a spp, (8. menzi,es'Li left foreground,B. atlteyluata background) ,

the sanpl ing method (on1y three were d iscovered in th is nanner) , and thei rfoliage projective cover over the whole prinary site was estinated. Thecover of trees was neasured from colour and infra-red aerial photographs ofthe site. The proportion of cover contributed by each tree species was thenestimated in the field. The prinary site was revisited from the end ofAugust to rnid-November to estimate the cover of species of annual-s and herbswhich lose thei r fo l iage seasonal ly .

. The cover of each perennia l p lant species in three 0.5 ha secondarystudy s i tes (1, 2 and 3, corresponding to R2, R3 and M2 respect ive ly ofDavidge 1979b) in other par ts of the study Srea-was es i inated re lat ive tothat in the primary study site in order to assess floristic variation withinthe area.

5 1

Soi l prof i les were obta inedin a l l s i tes and to 4.5 m in thedescribed in the field by nanualwith standard charts.

when the soi l was nois t by auger ing to 1.5 npr inary s i te . The exposed soi l wastexture assessment and conparison of colours

Specles were identified at the Western Australian Herbariurn by one ofus (AVM) or by herbariul staff. Voucher specinens and photographs werelodged as a pernanent record at the School of Environmental and LifeSciences . lfurdoch Uni-versitv.

32 \ r i r ̂ , . , - 1 , i . a n ^ . : t ^ 6r s w r ^ r q L . u d v r u t , v N o . 5

PHYSICAL ENVI RONMENT

The monthly distribution of average annual precipitation and averagernaxinum and mininurn tenperatures for the study area are given in Table 1.The clinate is mediterranean with 57% of the average annual precipitation of860 run fa l l ing in winter (June-Aug.) . This corresponds to var ious a l terna-t ive c l imate types such as Thornthwai ters (1948) . sub-humid, wam, great-nois ture-def ic iency- in-surnmer (CB's) c l inate, Kcjppen's (1956) t ransi t ionbetween a warn clinate with long hot summe r (Csa) and a warn clinate withlong temperate sunmer (csb) , and the UNESCO-FAO (1963) At tenuatedThemone dit erranean climate .

Winds are usual ly gent le and temperatures rareLy reach f reezing point ,possib ly as se ldon as once per year . The nean n in imum for the coldest nonthof the average year is about 9oC, while the difference between nean winterand rnean suffner tenperatures is only about 10oC (Bureau of Meteorology 196b) .Armual rainfall is very variable, rangi-ng fron 500 rnn to nore than 1,300 run.A wal ln to hot dry season is usual for 7 nonths of the year (Table 1) . InOctober , prec ip i ta t ion and evapot ranspi rat ion are equal (Speck 1952) . Inthe following weeks precipit.ation decreases rapidty and increasingtenperatures raise the evapotranspi ration . Although plants initially naydraw on water s tored in the soi1, the upper layers of so i l reach wl l t ingpoint by early Decenber and contj-nue to dry out until the sol1 water isreplenished in May (Speck 1952) .

Table 1, The nonthly distribution of average annual preci-pitation and averagenaxinun and ninj_mun tenperatures for the study area, after Bureauof Meteoro logy (1966) . Month ly prec ip j - ta t ion yalues were obta inedby nul t ip ly ing the values for Frenant le by a factor (1.1) der ivedfrom the total arnual precipitation for Frenantle and the isohyetwhich includes Jandakot t864 runl .

Ternperature (oC)

Month

Averageprecip i ta t ion

(run)Average

tnax .Absolute Average Absolute

nax. mln. rntn.

FebruaryMarchApri IMayJuneJuly'lUB(,rs L

Septernber0ctoberNovemberDecember

8 . 01 0 . 81 9 . 14 6 . 7

1 2 7 . 81 8 6 . 11 7 3 . 01 3 4 . 0

7 7 . L4 7 . 81 9 . 61 3 . 6

2 9 . 42 9 . 72 7 . 72 4 . 52 0 . 61 8 . 1L 7 . T1 7 . 81 9 . 32 L . 02 4 . 52 7 . 2

4 3 . 94 4 . 44 L . 33 7 . 6

2 7 . 62 4 . 62 7 . 83 3 . 33 7 . 24 0 . 34 2 . 2

1 7 . 57 7 . 77 6 . 41 4 . 01 1 . 6

8 . 88 . 89 . 0

1 0 . 0

t 3 . 71 q 0

9 . 28 . 77 . 74 . 0r . 71 . 67 . 21 . 92 . 64 . 4q q

8 . 6

Year 8 6 5 . 6 2 3 . 1 - 4 4 . 4 L 2 . 9 7 . 2

No. 5 Phenology of Banksia Woodland Ss

The area lies between the Bassendean and Spearwood Dune Systens(McArthur 6 Bet tenay 1960) , in a gent ly unduJ.at ing p le is tocene landscape ofwel l -dra lned, deep, coarse, s i l iceous, podsol ized, whi te over ye l low sandon the Swan Coasta l P la in (Tables 2a F" b) . The a l t i tude is 49-52 m abovenean sea level . The average depth of the water tab le, which is v i r tua l lyf la t here, is 24-28 n below ground sur face (water tab le naps held by theMetropol i tan Water Board) .

The study area covers part of a singl.e dune and the substrate variess l ight ly in d l f ferent s i tes as regards a l t i tude, s lope angle, the presenceof a partly organic surface layer and the depth below the surface it "hi ct.,ye l low sand and i ts associated band of sof t concret ions occurs, In soneplaces the upper , whi te sand is 4 m deep, whi le in one s i te (RJ) the f ineryel low sand reaches the sur face (Table 2a & b) .

The deep sandy soi l fa l1s wi th in the.Jandakot ser ies ( i ron-hunus pod.zols)of the Bassendean Associat ion (Bet tenay et aL, 1960) , but corresponds ioSpeckrs (1952) Karrakat ta Sands, perhaps t ransl t ional to h is Muchea Sands(par t o f h is Bassendean Associat ion) . Speck descr ibed both of these ash a v i n g a p H o f 5 . 7 - 6 . 6 n e a r t h e s u r f a c e a n d 7 . 0 - 7 . 9 a t d e p t h . T h e s e s o i l shave excellent infiltrati-on and drainage and are very tealhed. They comprise85% coarse sand, 11% f ine sand and L% c lay and s iLt (Speck 1952) . b t l ler -d e t a i l s a r e : C a C O J = 0 , 5 - 1 . 3 % n e a r t h e s u r f a c e , d e c r e a s i n g t o l - e s s t h a n0.2e" at depth; organic carbon = 3.4-5.7% near the sur face, decreasing to\ . ? ! : 8 % a t d e p t h ; P 2 0 5 = 9 . 9 9 1 2 n e a r t h e - s u r f a c e ; K 2 0 = 0 . 0 0 5 % ; a n d N =0 . 0 0 5 % ( S p e c k 1 9 5 2 ) .

FLORISTIC COMPOSITION

a. General descr ip t ion

The vegetat ion, in co l loquia l , terns, is , ' low woodland" ( t ree height5-6 m, canopy cover 20%) over "heath" (height 0.5-1.0 * , .unopy cover 60%).The tree stratun in the primary study site consists nainly oi'Banksiaat teruata (Proteaceae) , a l though f ive other species are present . Eucalyptusnarginata (Jarrah) and. CasuarLnct ft"as er,mta are represented by only two irthree t rees in th is s l te . The shrub stTatun compi ises g0 perennia l speciesin 25 families, of which Beaufortia elegmrs (lvtyrtaceae) ,id Lnu.opogoikirqianus (Epacridaceae) are particulatly pto*i'rerrt. Most species ;ccurwi th low cover va lues, of less than 2% (Tables S E 4) .

The 0,5 ha s i tes d i f fer s l ight ly f ron the pr i rnary s i te in the conposi_t ion of both the t ree and shrub st rata. The sarne species are genera l lypresent but have d i f ferent cover va lues (Table 3) . In one s i te 1U. ;C, f raseTana occurs as severa l large t rees; in th is and anotheru i i " tn , IE. marginata is nore conmon than in the prinary site. The shrubs A. ilZgurroand' L. kingimtus have lower cover in the secondary sites than in the prirnarysite, their place being taken by e,g, Leucopagon conostephioides, Erelmaeapauciflora" Casuan"Ltu hwnilis, Stirlingio tatlfolia and Htbbertiahyp-erLcoides.. Several species, e.g. Mesomelaetut stygia, Dat;iesia nudtfloraand Hibbeytia racemosa, are restricted to one slte (i2), apparentlyassociated wi th patches where yel low sand reaches gro-und sur face.

' In

addi t ion, secondary s i te Mj has a re lat ive l .y h igh -over ( to ta l 20%) otperennial monocotyledons, nai"nly pateysonLa (Iridaceae) , phlebocctt taa(Haemodoraceae) , LyginLa (Rest ionaceae) and, Anrphipogo, ' lpoaceae) ( iab1e J) .

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35Phenology of Banksi,a Woodland

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No, 5 Phenology of Banksia Woodland 37

Table 3, Average fo l iage pro ject ive cover of perennia l p lant species, inorder of decreasing cover, in the prinary study site (1 ha ofBanksi ,a woodland) . + = cover of less than 0.1%. Also g iven arethe p lant species found in three 0.5 ha secondary s tudy s i tes (1,2 and 3 corresponding to sites R2, Rg and M2 respectively ofDavidge 1979b). For these s i tes, fo l iage pro ject ive cover wasonly est inated and is g iven for those species which e i ther ( i ) havecover greater than 2% i-n at least one si-te, or (il") are absent fronat least one s i te ; the presence of other species is denoted by anaster isk. The t ree st ratum was def ined as a l l p lants ta l ler that3 n. Total foliage projective cover in the primary site wasest i -mated to be 60-65% f rom aer ia l photographs and f ie ld assessment .

Secondary s i tes

SpeciesPrinary

TREE STRATUM1. Banksia attenuata R.Br.2. Banksia menziesi i R.Bt.3. EucaLyptus todtiarn F. Muell.4. Nuytsi.a fLottLbundn (Labill.)

R . B r .5. Eucalgptus maz:ginata Sn.6 . CastnzrLrn fz,aserana l'liq.7 . Banksia iL ieLfol ia R.Br.

SHRUB STRATUMU. Beaufortia eLegans Schau.9. Leucopogon corns tep?riotdes DC.

10. Leucopogon kirqianus (F. Ituel1.)C .A . ca rdn ,

11. Hi.bber,tla hgperLcoides (DC.)Benth.

t2. St i r l ingia Lat i fo l ia (R.Br. )Steud.

13. Eremaea paueLflora (End1 . ) ttruce!4. Scholtzia inuolucrata EndI .15. Hi.bbertia subuagirwta (Steud. )

F . Mue l l .16. Melaleuca thgmoides Labi l l ,17 . LygLn ia bo rba ta R .Br .18. AcaaLa pulcheLLa R.Br.19. Corns tepfuLun pen&)lton Benth,20, DasApogan broneLiaefolizzs R. Br.2L. CaLy tu"Lr flauescern A. Cunn.22. Eaemaea sp. aff. fi..nbr"Lata Lindl.23. Petr'ophiLe Lineaz"Ls R.Br.24. Dauieeia juncea Sm.25. Bosstaea eriocatpa Berfth.26 , Hemicrndya purryens R.BT.27, Lommtdta caespitosa (Benth. )

Ewart28. Loaocatya flexuosa (R.Br.) Benth,

6 . 92 . 97 . 2

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2 . 7

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3 8

Table 3 (cont. )

A.V. Mi lewski & C. Davidge N o . 5

Secondary s i tes

SpeciesPrj-mary

s i t e

29. Phlebocary a ciLiata R.Br.30. Schoenus cunifolito lR.Br .)

Benth.J l . 5LA L1d l ' Ln repenB R .Br .32. Cassytha sp. indet,33. Casuarina hwniLis Otto et Dletr.34. Cornstylis aculeata R.Br.35. Gompho Lobiwn tomentosLrn Labill.36, Hypo calynrna z:obustwn EndL.37. HApoLaene exsuLca R.B'r.38. JacksonLa t 'urcel laLa (BonpL) DC.39. La:nnanni,a rotnosa LindL.40. Leucopogon stTL ctus Belth.41,. LomftidTa endL.Lchet"L (F. Mue1l.)

Ewart42, MacrozatrLa ztLedlei (Gaud.)

C ,A. Gardn.43. Phlebocarya fnlifolia (F. Ituell.)

Benth.44. Rest i ,o sp. indet.45. Schoenus by.euisetLs Benth.46. StgLi&LLon b rLtlontanwn Benth.47. Acac:La stenopte[a Benth.48. Anrphipogon tta;l:inatus R.Bt.49. CaLy trLr fyuserL A. Cunn.50, ErLos temon spicatus A. Ri.ch.51. Hibbe:r,tia aurea Steud,.52. Hibbertia huegeli.i (EndL.)

F. ]vfuel1 .55. Hibbey,tia pachgz,rhi.za Steud.54, Ho,ee. tr"Lsperma Benth.55. Larmenni.a squarrosa. Lindl.56. LechenauLtia flor"Lbunda R.Br.57. Lepidospezma cLngus taturn R.Br.58. Ly si,nema ciltatun R.Bt.59. MeLaleuca seriata Lindl.60. 0rg Lobiwn capitatum Benth.61. Pate?s orxLa occidentalis R.Br.62. Platgsace conry)Tessa (Labi11.)

Nornan63. Xanthosia tn tegel i i (Benth.)

Steud.64. Acacia huegeLi,i Benth.65. Arrto cr"Lnum preissii Lehn,66. Calectasia cAdnea R.Br.67. ConostgLis aurea LindL.68. Conostg Lis juncea EndL.69. Conostylis seti.ger(r R.Br.70. Danpiez,a Lineayl,s R.Bt:.

0 . 5

0 . 30 . 50 . 20 . 2o . 20 . 20 . 20 . 20 . 20 . 20 . 2

0 . 2

0 . 2

0 . 2o . 20 . 20 . 20 . 10 . 10 . 10 . 10 . 1

0 . 10 . 10 . 10 . 10 . 10 . 10 . 10 . 10 . 10 . 1

0 . 1

0 . 1

++

+

+

0 . 3 0 . 3 1 . 5

4 . 0

o_ ,

q n , 1 a

o 1 n ?

+

0 . 1

0 . 5

+

0 . 6 1 . 3

1 . 61 . 0

r-

N o . 5

Table 3 (cont . )

Phenology of Bml<sia Woodland 39

Secondary site

SpeciesPrinary

site

7L. Dnosera.palea72. Hensmania turbinata (Endf . )

W . V . F i t z g .7 3 , LeptomerLa erpetrLfonnts lt':..q.74. Leucopogon puLcheLlus Sond.

or .L. polymorphus Sond,.75, Leucopogon y'acemulostrs DC.76. Lomandya pz'eissi.i (Endl .) Ewaxt77. MeLaLeuca scabra R.Br.78. Pe:rsoonta saccatq. R.Br.79. Pt)thoeazpa puLchelLa Lindl,80. Poaceae sp, indet. 181. Poaceae sp. indet. 282. Scaeuola paludosa R.Br ,83 , Sty Li.&twn juncewn R.Br.84. StglidLwn pili,ferwn R.Bt.85. Sty LidLwn schoenoi.des DC.86 . ThA sanotLtB tt tmdrus (Labil1 . ) R. Br,87, I rLcoryne eLat ior R.Br.88. Buttonia conferta DC.89. Cyperaceae sp. indet,90 . Dauiesi.a ru,tdifloz,a Meisn.91. Hibbez'ti,a racenosa. (Endf,) Gi1g.92. MesomeLaenn s@rgria (R,Br.) Nees93. HneLea sulphu.z,ea Meisn.94. RegeLi.a tnops Schar:.95. Schoenus sp. i ndet .96. Sgnaphea spirutlosa (Burm. f. )

Merr i l l97. Xo:nthornhoea prei.ssii End,I.

++

+

++

++

+

+

+

0 . 7

U . J

40

Tab le 4 .

A.V. I t i lewski € C. Davidge N o . 5

Est imated re lat ive perennia l cover for a l l p lant fan i l ies occurr ingin the study area, in order of decreasing covef. Nunbers inparentheses refer to species as l is ted in Table S. Tota l fo l iagepro ject ive cover was obta ined by adding cover va lues of species;no account was taken of overlap between canopies in thecalcu lat lon.

iI

FarnilyTota l fo l iage

pro ject ive cover (%)

M y r t a c e a e ( 5 , 5 , 8 , 1 3 , 1 4 , 1 6 , 2 L , 2 2 , 3 6 , 4 9 , 5 9 , 7 7 , 9 4 )P r o t e a c e a e ( 1 " , 2 , 7 , 1 2 , 2 3 , 7 8 , 9 6 )E p a c r i d a c e a e ( 9 , 1 0 , 1 9 , 4 0 , 5 8 , 7 4 , 7 5 )D i l l e n i a c e a e ( 1 1 , 1 5 , 5 1 , 5 2 , 5 5 , 9 1 )R e s t i o n a c e a e ( 1 7 , 2 8 , 3 7 , 4 4 )F a b a c e a e ( 2 4 , 2 5 , 3 5 , 3 8 , 5 4 , 6 0 , 8 8 , 9 0 )X a n t h o u h o e a c e a e ( 2 Q , 2 7 , 4 7 , 6 6 , 7 6 , 9 7 )Casuar inaceae (6, 3 3)Minosaceae (L8,47,64)H a e m o d o r a c e a e ( 2 9 , 3 4 , 4 3 , 6 7 , 6 8 , 6 9 )Loranthaceae (4)C 4 r e r a c e a e ( 5 0 , 4 5 , 5 7 , 8 9 , 9 2 , 9 5 )S t y l i d i a c e a e ( 5 1 , 4 6 , 8 3 , 8 4 , 8 5 )Lamiaceae (26 )L i l i a c e a e ( 3 9 , 5 5 , 6 5 , 7 2 , 8 6 , 8 7 )P o a c e a e ( 4 8 , 8 0 , 8 1 )Apiaceae (62,63)Goodeniaceae (56,70,82)I r idaceae (61)Lauraceae (32)Rutaceae (50)Zaniaceae (42)Asteraceae ( 79)Droseraceae ( 71)Santa laceae (75JThynelaeaceae (9 3)

1 9 . 01 5 . 07 2 . 6

L 7

1 . 61 . 31 1

1 . 10 . 90 . 80 . 70 . 60 . 60 . 40 . 40 . 20 . 10 . 10 . 10 . 10 . 10 . 1

++

A specj-es present in the study area but absent fron a1l study sites isAdern:nthos ca gnorwn, which occurs as a stand of tall shrubs between second.arys i t e s N 4 J a n d R r .

The vegetation of the study area does not conespond precisely to anypreviously described comrnunity, although it has been described in a qeneralsense by Die ls (1906) , Seddon (L972) and Heddle, Loneragan and Havel (1979,I tBassendean - centra l and south" vegetat ion complex) and corresponds toSpeckrs (1952) I 'Banksia, Casuar ina, Eucal )?tus Associat ion ' , , and Beard 's(1979) Bassendean Systen: southern or per th sect ion. The vegetat ion d i f ferss l ight ly f rom that nor th of Per th, e .g. t fpe F of Havef (1976) , in lack inge.g. Jacksonia florLbmda and rnenbers of the genera Boronia, Astroloma andConospeTrnwn. The Banksia woodland described by Bel1 et aL, (1979) occurson weakJ-y leached sand associated with linestone outcrops and differs in theconposl t ion of the shTub st ratun.

N o . 5 Phenology of BanksLa Woodland

Eastwards fron the Karrakatta/Bass endean tnarsition, "as the lfuchea(Bassendean) sands becone nore typlcal the Jarrah trees become fewer, arereduced to stunted trees and finally are absent altogether" (Speck 1.952) .It is as part of this "Jarrah-Banksla ecotonefi Speck 1952) that the studyarea can best be descr ibed, t ransi t ional as i t is between stands wi th acanopy of E. matgi,rnta and, CasLrarLnq fraserana over snaller tlcees of Battksia,and those containing onLy Banksia and, E. todtiana (Speck !952, Beard L967,Seddon 19721 . The shrub stratrn appears to represent a slrnilar transitionfrom yellow to white saad, as shown by the rare local occurrence of yellowsand species such as Mesomelaentt stAgia (Havd, 7976).

b. Honogenei ty

The prinary study site (1 ha) appears to tcepresent a single plantcorununity, as shown by a species-area curve (Fig. 5). The very smalL varia-tions in floristic cornposition here nay be attributed to the chanceoccurrence of local ly rare species in cer ta in par ts of the s i te (Table 5) .

NO. 2m'z PIOTS

Fig. 5. Cunulative total nunber of shrub species found in consecutive 2 n2sanple plots in the t ha pr i rnary si te. Note that the sanple plotswere not adjacent, but seattered through the si te.

Over the ent i re area, as represented by the secondary s i tes, f lor is t icconposition varies slightly from place to place in association with subsrratedifferences. In view of the large area studied, this variation is relativelysnal1, being found nainly in the cover of species rather than in theirpresence or absence. Such variation seens unavoidable in view of therequirenent of an area large enough to achieve a minimally acceptable sarnple12735-4

4 I

I

!c(J

z

42 A . V . M i l e w s k i & C . D a v i d g e N o . 5

studies of vegetation structure and the anirnal conmunity as well as forf lor is t ic s tudy.

PHENOLOGY

Table 5 g ives l year 's observat j -ons on the f lower ing and f ru i t ing t inesof 81 perennia l p lar t species found in the study area. These data werecol lected f ron March 1978 to February 1979, an unusual ly dry year . Theprevlous sunmer (1977-78J was ext remely hot and dry and the ra ins arr ivedabout a month Later than usual (Met . Bureau records) . Thus the phenologica ldata presented do not refer to ar average yeat : . For instance, in thefo l lowing year nany p lants were observed f lower ing in the study area atleast a rnonth ear l ier than recorded in 1978. However, the data do a l lowsome general izat ions to be nade.

The per lod of maximul f lower ing, i .e . when the nost species are infu l l b loon, is October . However, sone species f lower at the cold t ine ofyear and others, such as the phys iognoni cally dominant Banksia attenuata and,Beaufoz'tia elegons, flower in sunner. "For a tine the epacrids doninate thewho1e, the c l imax of the i r d isp lay being of ten reached in Mayr ' (Diefs 1906) .Banksia menziesii, Lyginia baybata and. Calytt"ir fnaserL are examples ofspecies having prot racted f lower ing seasons inc luding par t o f sunner ,a l though no species was recorded f lower ing throughout the year , The legurnes(Fabaceae and I ' t inosaceae) are of specia l in terest as a source of seeds forants, b i rds and rodents (Majer 1978, Bra i thwai te & Gu1lan 1978) . The podsof legurnes in the study area (e.g. species of BossLaea, GotnpthoLobiun,Jctcksonla and )ry Lobium) reach naturity, dehisce and release seed on to theground nain ly in suruner (Dec.-Feb.) , var ious species fo l lowing each other rntu1n.

Table 6 g ives the t ines of appearance of species wi th annualfo l iage, to ta l l ing 25 addl t ional species not l is ted in Tab1e 3. Severa lof these specles have recent ly been ln t roduced to Austra l ia . Both ind igenousand introduced annuals (mainly Poaceae and Asteraceae) and other herbs(mainly Orchidaceae and Haernodoraceae) are apparent only in winter andspr ing and wi ther by December.

The phenology of p lants in the study area nay be largely expla ined byra infa l l , temperature and photoper iod (Speck 1952) . There are two t ines ofyear when these factors night retard plant growth. In winter, nininuntemperatures and photoperiod are reached, while in summer lack of noisture .r- sthe l j .n i t ing factor . This d iv ides the year in to two per iods favourable toplant growth. The shor ter of these star ts in la te Apr i l or May wi th thef i rs t ra ins, and the longer per iod inc ludes October . possib ly speciesflowering during the long sunmer drought are able to do so becauie of deeproot systens which penetrate to the underground water (Speck 1952) ,However, the physio loglca l adaptat lons which a l low the t iny, shal low-rootedperennial herb, Stg Li&Lum yepens, to flower in sumrne r remain a puzzle.

The only other s tudies in Austra l ia of p lant phenology in areas ofs in i lar c l imat ic and edaphlc condi t ions are those of pat ton (1939) , Spechtand l{ayson (1957) and Groves and Specht (1965) , The October peak of flower_ing in the study area agrees with that found by patton (193SJ and Groves andSpecht (1965) for heath ln coasta l V ic tor ia , but is a nonth ear l i -er than thatobserved by Specht and Rayson (1957) for heath in eastern South Australie,.Fewest species flower i,n April and May in the Victorian heath, agreeing with

forthe

43Phenology of Banksia Woodl and

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L 4 +

tir ri .-{ Ln sS LD S'|o Ln <.

N sl- N f.l fO fO fQ Ln fO- r | Q N | Q N f O | o L n | o

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Phenology of Banksia Woodland

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46 A.V. I r t i lewski & C. Davidge N o . 5

Table 6. Phenology of annuals and herbs bearing foliage only in winter andspring and wilting by December. The maxinurn cover of each specieswas less than 0.1e" of the study area. * indicates perennial species.

Fanily and Time of peakflowering

APIACEAETracLtgmene pilosa Sm.

ASTEMCEAEHypo choez"Ls sp. indet. (introduced)Mi l lot ia terwifol ia Cass.Podotheca chrg santha Benth.PodoLepis spp. lndet.ureinia anthemoides (L.J Gaertn, (introduced)

CENTROLEPI DACEAECentrolepis dtunnnondLi (Nees) Hieron

CRASSUT"ACEAECt:assuLa sp. indet.

CYPERACEAESaLrpr,Ls sp . indet .

DROSERACEAE* DTosera er:y thronhiza LindL,*Wosera nenziesi i , R.Br. ex DC.

HAEMODORACEAT* AwLgozanthos hwntlis LindL .* Anlgozanthos mm'glesii o. Don

IRIDACEAE* GladtoLus catyophy LT.acezrs (Burm. f . ) Poir. (introduced)

LI LIACEA!* Buy,chat: itLa wrbeLlata R.Bt.* Sotoenbaea LcwLflora Lindl.

ORQII DACEAE* Caladenia flaua R.Bt .* CaLadsni,a dLs coidBa Lil,dl.* Pter:ostylis rea)ruq ? Benth.* Pterostylis u1:ltltata Lindl.* ThelynrLtxa cattp anulata LindL.

POACEA-EAira caz,g ophg LLea f. (introduced)BvLza mqs'Lma L. (introduced)Stipa sp. i\det .D a a c a c a i n . l A f

ear ly Oct .ear ly-n i .d Oct .ear ly-n id Oct .ear ly Oct .

mid Sept .

; ; ; ; :

ear ly oct .

ear ly oct .

ear ly Oct .ear ly Oct .ear ly Sept .ear ly Sept ,n id Oct .

eanly Oct .ear ly Oct .mid Oct .

No, 5 Phenology of Banksia Woodland 47

the present s tudy, In Vic tor ia , where there is a more even d is t r ibut ion ofrainfall through the year than in the study area, a sllghtly higher propor-tion (35% compared wi-th 14%) of perennial species flower in sunmer, includingsevera l which f lower throughout the year (Pat ton 1935, Groves & Specht 1965) .However, Myrtaceae flower rnainly in spring in Victoria (Groves fi Specht 1965).,although those in the study area (e.g. species of Melaleuca, CaLy tr"Lx andScholtzia) reach their peak in suruner. These genera have also been foundf lower ing in sunrner e lsewhere on the Swan Coasta l P la in (Die ls 1906) .

REFERENCES

Beard, J ,S, (1967) . Natura l woodland in King 's Park, Pef th. WestemAustralian N atur.crT:L s t 10: 77 - 84.

Beard, J .S. (1979) . The vegetat ion of the Pin jarra area, Western Austra l ia .Vegetat ion Survey of Western Austra l ia , 1 :250,000 Ser ies: Maps andExplana. tory Memoir . Vegnap Publ icat ions, Per th.

B e l 1 , D . T . , L o n e r a g a n , W . A . & D o d d , J . ( 1 9 7 9 ) . P r e l i r n i n a r y v e g e t a t i o nsurvey of Star Swamp and vicinlty, Western Australia. WesterrL Atntz'alianHerbat+um Resea:rch Nates 2: I-21.

B e t t e n a y , E . , M c A r t h u r , W . M . f r H i n g s t - o n , F . J . i 1 9 6 0 ) . T h e s o i l a s s o c i a t i o nof part of the Swar Coastal Plain, llestern Australia. Soils and LandU s e S e r L e s n o . 3 5 , D i v i s i o n o f S o i 1 s , C , S . I . R . 0 .

Bra i thwai te, R.W. & cul1an, P.K. (1978) . Habi tat se lect ion by snal l maf fnalsin a Victorian heathland. Attstt alia:rL Jourral of EcoLogy 3: 109-27.

Bureau of Meteoro logy (1966) . ClLmat ic su.xDeA, region 15. Metropol i tanWestern Austra l ia . Issued by the Director of Meteoro logy, Melbourne.

Davidge, C. (1979a) . A census of a cornmuni ty of sma11 terrest r ia l ver tebrares.AustnaZi .an Joutnnl o ! tco logg 4: Ib5- I70.

Davidge, C. (1979b) , The Herpetofaurnl Conmtmitg of a Battksia WoodLayLd nea!Perth, Wes ter.rt Australia, Ph.D. Thesis, School of Environrnental & LifeSciences, Ifurdoch Univers ity.

Dle ls , L . (1906) . Die Pf lanzenwel t von West Austra l ien s i jd l ich desWendekreises. Vegetation der Erde 4 (The Plant Wot'Ld of WestemAustyalia South of the Iropics; English translation by W.J. Dakin,L920. Unpubl ished rnanuscr ip t , J . ibrary at W.A. Herbar i r .un, South pef th) .

G r o v e s , R . H . Q S p e c h t , R . L . ( 1 9 6 5 ) . G r o t r t h o f h e a t h v e g e t a t i o n . I . A n n u a lgrowth curves of two heath ecosystens in Australia. AustraLian JourrtaLof Botarry 13: 261-80.

Havel , J .J . (1976) . The potent ia l o f the nor . thern Swan Coasta l p la in forPirnts pirnste" Ait, plantations. Foyests Depa.z,tment BuLletin no. 76.Forests Departnent , Per th.

Heddle, E.M., Loneragan, 0. ! t / , & Havel , J .J . (1979) . Vegetat ion of the Dar l ingSystem. II],: DaTLing Systen Atlas. Envlronmental pt:otection Authority,

. . P e r t h .Koppen, W. (1936) ttDas Geographische Sg stem der Klimatett being Vo1 . 1,

part C, of ttHandbuch dez, Kli.matologiett , Berlin.Majer , J .D. (1978) . Studies on inver tebrates in re lat ion to bauxi te n in i . r rp

in the Dar l ing Range. A rev iew of the f i rs t 18 monthsr research.Enuirownental Besearch Bulletin 3. Alcoa, Westem Australia.

McArthur , W.M. & Bet tenay, E. (1960) . The developnent and d is t r ibut ion ofthe soi ls of the Swan Coasta l P la in, Western Austra l ia . C,S. I .R,A.(Austv,alia), SoiL PubLication no. 16.

Pat ton, R.T. (1933) . Ecologica l s tudies ln Vic tor ia : the Chel tenhan f1ora.Pz'oceed:Lrqs of the Roy aL Society of VictorLa 45: 205-18.

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4 8 A.V. Mi lewski & C. Davidge N o . 5

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