CHAPTER THREE

Temperate woodlands in Victoria: distribution,composition and conservation

IAN LUNT1 and ANDREW F. BENNETP

Prior to European settlement, temperate woodlands occupied 32% of Victoria, but by 1987 92% of woodlandshad been cleared, mostly for agriculture, and much of what remains is fragmented and often degraded. Theremaining woodlands contain a wide variety of ecosystems dominated by a range of tree species, and thesewoodlands harbour a diverse flora and many rare and threatened plant species. A distinctive woodland faunacannot be identified due to overlaps with adjacent vegetation types. but it is clear that the fauna has been greatlyaffected by the widespread loss of habitat and fragmentation. Impacts were greatest on the medium-sized ground-dwelling mammals, many of which became extinct. Other faunal groups have also been impacted to variousdegrees. Conservation of woodlands in Victoria will require the protection of existing natural areas to preventfurther degradation and loss, the management of habitats to enhance conservation values, and the long-termrestoration of the ecosystem.

INTRODUCTION

BEFORE European settlement, temperatewoodlands occupied large areas of northern,western and southern Victoria. Nowadays,most woodlands have been transformed toagricultural pastures and crops, and theconservation status of virtually all woodlandcommunities is extremely poor. In this chapter,we provide an overview of the distribution,composition and conservation status oftemperate woodlands in Victoria.

Most temperate woodlands in Victoria occurin flat or gently undulating landscapes on soilsof moderate to high fertility. The groundstratum is generally dominated by herbaceousspecies (usually grasses), and relatively fewspecies of sclerophyllous shrubs are abund-ant. "Typical" Australian families such asMyrtaceae, Proteaceae and Epacridaceae arepoorly represented (Frood and Calder 1987;Lunt 1991; Department of Conservation andEnvironment 1992).

The Victorian Department of NaturalResources and Environment (DNRE) recentlymapped the pre-European and presentdistributions of 28 "broad vegetation types"across Victoria (Department of NaturalResources and Environment 1997a). Nine ofthe 28 "broad vegetation types" typically formtemperate woodlands (coastal grassy wood-lands, sedge-rich woodlands, plains grassywoodlands, herb-rich woodlands, montanegrassy woodlands, riverine grassy woodlands,rainshadow woodlands, Mallee woodlands andWimmera mallee woodlands). In addition,

many areas mapped as grassland supportedan open savannah woodland at the timeof European settlement (McDougall andKirkpatrick 1994), and shared a similar plantcomposition with nearby woodland ecosystems.Figure 1 shows the distribution of temperate.woodlands and grasslands in Victoria, basedon DNRE's map of broad vegetation types.Temperate woodlands were most widespreadacross northern and western Victoria, onalluvial, riverine, basaltic and coastal plains(Fig. 1).

The above circumscription of temperatewoodlands excludes box-ironbark forests,which traditionally have been viewed as adistinctive ecosystem (e.g., Frankenberg 1971;Frood and Calder 1987; Conn 1993). Morerecently, the term "box-ironbark woodlands"has been interpreted more broadly (usuallyby zoologists) to include both "box-ironbarkforests" on infertile auriferous soils and"box woodlands" on the adjacent agriculturalplains; only the latter ecosystem is hereregarded as a temperate woodland ecosystem.

Prior to European settlement, temperatewoodlands occupied over 7 million ha inVictoria, 32% of the state. Grasslands andwoodlands collectively occupied 9 million ha,or 40% of the state (DNRE unpubl. data,May 1997: Table 1). Today, little of theoriginal woodland estate remains. By 1987,92% of woodlands had been cleared (mostlyfor agriculture), and only 575000 ha remained.The 1987 estimates were obtained from aerial-photo analysis of tree stands, and do notreflect the condition of the understorey,

IThe Johnstone Centre. Charles Sturt University. P.O. Box 789. Albury, New South Wales. Australia 2640.'School or Ecology and Environment. Deakin University. Rusden Campus. 662 Blackburn Road. Clayton. Victoria, Australia 3168.Pages 17-31 i. TEMPERATE EUCALYPT WOODLANDS IN AUSTRALIA: BIOLOGY, CONSERVATION, MANAGEMENT AND RESTORATION ed byR. J. Hobbs and C. J. Yates. Surrey Beauy Ie Sons. Chipping Norton. 1999.

18 TEMPERATE EUCALYPTWOODLANDS IN AUSTRALIA

• Woodland

• GrassJand

DOther

Fig. 1. Distribution of temperate woodland and grassland in Victoria at the time ofEuropean settlement. Simplified from Department of Natural Resources andEnvironment (1997a).

which is grossly degraded in most woodlandremnants.

Source: Flora Section, Department of Natural Resourcesand Environment, and Natural Resource Systems, May1997.

Tabk 1. Area of Victoria occupied by temperate woodlandbroad vegetation types, in 1750 (before Europeansettlement) and in 1987.

The amount of habitat loss since Europeansettlement varies dramatically among thedifferent woodland vegetation types (Table 1).However, the ecosystems which were originallythe most widespread have suffered the greatestdestruction (Fig. 2). This pattern reflects thewholesale conversion of woodlands on thenorthern and western plains to agriculture.Three of the four ecosystems which wereoriginally the most widespread (plains grassywoodland, grassland and Wimmera malleewoodland) have fared the worst, with lessthan 3% remaining. At the other extreme,woodlands in eastern Victoria have faredrelatively well, with over 50% of rainshadow

60

50

atC 40"i:'iie! 30~ •

20 • •10 •

•00 2 3 4 5

OrIginal__ (millon he)

Fig. 2. Original area of broad woodland vegetation typesversus the percentage remaining in 1987. Source:Flora Section, Department of Natural Resourcesand Environment, and Natural Resource Systems,May 1997.

70

woodland and montane grassy woodlandremaining. Again, remnants of both ecosystemsare often extremely degraded, especially bystock and rabbit grazing (e.g., Clayton-Greeneand Ashton 1990). Based on these figures,riverine grassy woodlands now account for34% of the remaining temperate woodlandestate in Victoria (Table 1).

In most regions, most indigenous vegetationhas been lost, and pastures and crops with fewnative species and a high exotic cover,prevails.Thus, the traditional "habitat fragmentation"

Area (I OOOsha)

1750 1987

4140 124997 201862 2691 68224 61173 3577 4359 3926 2

I 882 8

9133 584

Broad vegetation type

Plains grassy woodlandRiverine grassy woodlandWimmera mallee woodlandHerb-rich woodlandMallee woodlandCoastal grassy woodlandRainshadow woodlandMontane grassy woodlandSedge-rich woodlandGrassland

TOTAL

LUNT and BENNETI': TEMPERATE WOODLANDS IN VICTORIA 19

model is. generally more applicable thanthe "habitat variegation" model, which hasrecently been developed in the New EnglandTablelands in New South Wales (Mcintyreand Barrett 1992). Without doubt, the mostwidespread woodland landscapes in northernand western Victoria are highly fragmented,not variegated. Within this fragmentedlandscape, a "habitat segregation" modelhas received considerable attention fromVictorian botanists. Under this model, sitesmanaged in different ways (e.g., burningand grazing exclusion vs grazing and fireexclusion) often conserve different portionsof the original regional species pool (Lunt1995, 1997; Foreman 1997). The segregationmodel focuses only on the best qualityremnants, rather than the entire landscape,but emphasizes the divergent effects ofdifferent management regimes on native plantspecies. It is probably a more useful predictorfor plant distributions than for larger verte-brate fauna, since it assumes that managementhistory is a more critical component ofplant composition than landscape issues suchas remnant size, shape or connectivity, ashas been found elsewhere (Kirkpatrick andGilfedder 1995; Prober and Thiele 1995). Itis worth noting, however, that managementhistory can have important local effects onresources required by certain animals, suchas tree hollows (Bennett et al. 1994), and inthis way can influence the 'composition anddistributional pattern of the fauna.

FLORA OF TEMPERATE WOODLANDS

Ecosystem descriptions

A wide variety of distinctive woodland eco-systems occurs in Victoria (Table 2), and thefollowing descriptions are adapted fromFrood and Calder (1987), Department ofConservation and Environment (1992) andConn (1993). In semi-arid, northwesternVictoria, Slender Cypress Pine Callitris preissiiand Buloke Allocasuarina luehmannii formedwoodlands on sandy lunettes (Table 2). BeforeEuropean settlement, the ground layer wasprobably dominated by perennial Spear-grasses (Stipa species) but the community hasusually been heavily grazed, and introducedspecies, mostly annuals, now dominate theground layer at most sites. Black BoxEucalyptus largiflorens, Buloke Allocasua~inaluehmannii and Bull Mallee Eucalyptus behnanaoriginally formed extensive woodlands onthe Wimmera plains. Only scattered remnantsof these vegetation types now exist. Theoriginal ground-layer is thought to have beendominated by Kangaroo Grass Themeda triandra,Common Wallaby-grass Danthonia caesPitosaand Spear-grasses Stipa species (Morcom andWestbrooke 1998).

The original vegetation of the NorthernPlains has been extensively modified by agri-culture, as in the Wimmera, and only smallremnants now occur. River Red Gum Eucalyptuscamaldulensis was the original dominant in

Table 2. Common (or once-common) dominant tree species in temperate woodlands in Victoria.

Alectryon oleifolius Cattle-BushAllocasuarina luehmannii BulokeAllocasuarina verticillata Drooping She-oakBanksia marginata Silver BanksiaCallitris glaucophylla White Cypress PineCallitris preissii Slender Cypress PineCasuarina pauper BelahEucalyptus albens White BoxEucalyptus behriana BuIl MalleeEucalyptus blakelyi Blakely's Red GumEucalyptus bridgesiana Apple BoxEucalyptus camaldulensis River Red GumEucalyptus camphora Mountain Swamp GumEucalyptus laTgiflorens Black BoxEucalyptus leucoxylon YeIlowGumEucalyptus melliodora YeIlowBoxEucalyptus microcarpa Grey BoxEucalyptus ovata Swamp GumEucalyptus pauciflOTa Snow GumEucalyptus rubida CandlebarkEucalyptus stellulata Black SaIleeEucalyptus tereticomis Forest Red Gum xEucalyptus viminalis Manna Gum x x

Regions: MAL = mallee, WIM = Wimmera, NP = northern plains, WE = western plains, GIP = lowland Gippsland,EA = montane eastern Victoria. These six regions are simplified from the 16 "natural regions" identified by Conn(1993). MAL = Conn's Murray Mallee and Lowan Mallee regions; WIM = Wimmera region; NP = Riverina andnorthern Midlands regions; WE includes Wannon, Grampians and Victorian Volcanic Plain regions; GIP = GippslandPlain, and EA = parts of East Gippsland natural region (includes rainshadow woodlands).

Scientific name Common name MAL

xx

xxx

x

xx

WIM

x

x

x

xx

x

NP

x

x

x

x

x

xx

WE

xxx

x

xx

xxx

GIP

xx

xx

x

xxx

EA

x

x

x

xxx

20 TEMPERATE EUCALYPT WOODLANDS IN AUSTRALIA

regularly flooded, riverine areas. The under-storey in elevated, irregularly flooded areaswas dominated by Kangaroo Grass Themedatriandra, Wallaby-grasses Danthonia species,Spear-grasses StiPa species and Windmill GrassChloris truncata. Considerable uncertaintyexists on the original dominant grassesthroughout most of northern Victoria, o~ngto the extremely rapid, and virtually complete,elimination of C4 grasses such as KangarooGrass (Moore 1953; Robinson and Mann1996; Morcom and Westbrooke 1998). Inthe west of the region, E. largiflorens wasdominant in woodlands on the drier marginsof the River Red Gum woodlands, withKangaroo Grass Themeda triandra, Wallaby-grasses Danthonia species and Spear-grassesStiPa species and a variety of chenopod shrubsin the ground layer. Extensive woodlands ofGrey Box Eucalyptus microcarpa occurred acrossmuch of the Northern Plains, often withWhite Cypress Pine Callitris glaucophylla,Buloke Allocasuarina luehmannii and, at times,Yellow Box Eucalyptus melliodora. White BoxEucalyptus albens was often dominant inthe east. The ground layer of relativelyundisturbed remnants is usually dominated byWallaby-grasses Danthonia species and Spear-grasses StiPa species, although Kangaroo GrassThemeda triandra is likely to have dominatedin many areas before grazing by Europeanstock.

Much of western Victoria supported opengrasslands and grassy woodlands (Fig. 3)dominated by a variety of species, includingRiver Red Gum Eucalyptus camaldulensis,Manna Gum E. viminalis, Yellow Box E.melliodora, Swamp Gum E. ovata, Grey Box E.microcarpa, Drooping She-oak Allocasuarinaverticillata and a tree form of Silver BanksiaBanksia marginata. The latter species hasdisappeared from vast areas in which it wasoriginally a co-dominant. Kangaroo GrassThemeda triandra again dominated the groundlayer in most well-drained areas, with Tussock-grass Poa labillardieri in poorly drained sites.

The lowland plains of eastern Victoriasupported grassy woodlands dominated byManna Gum Eucalyptus viminalis, Black She-oak Allocasuarina littoralis and, in some places,Snow Gum Eucalyptus pauciflora. Forest RedGum Eucalyptus tereticornis formed extensivewoodlands and open forests on the lowlandGippsland plain. Regular burning byAboriginals may have helped maintain anopen woodland structure in many lowlandcoastal areas, and there are many instancesof tree and shrub invasion since Europeansettlement. Small, but distinctive, outliers ofgrassy woodland occur on fertile soils ininter-montane basins in montane eastern

Victoria, for example at Buchan, Omeo,Delatite and Wulgulmerang. The dominanteucalypts included Snow Gum Eucalyptuspauciflora, Candlebark E. rubida, Yellow BoxE. melliodora and Mountain Swamp Gum E.camphora. The ground layer was originallydominated by Kangaroo Grass Themedatriandra and Tussock-grassesPoa species. Manyof these woodlands are similar in compositionto woodlands on the Southern Tablelands ofNew South Wales.

Plant diversityThere are no published estimates of plant

species diversity in temperate woodlandsin Victoria. However, unpublished quadratdata held by the Victorian Department ofNatural Resources and Environment, supple-mented by recent species lists (Lunt 1992;McDougall and Kirkpatrick 1994; Robinsonand Mann 1996), include nearly 1 000 nativeplant species. This data set includes manygrassland quadrats, but as virtually all nativegrassland species also occur (or occurred) inwoodlands (McDougall 1994), the figureremains a valid estimate for plant diversityin grassy woodlands. Assuming that an extra100 species (if not more) might be addedfrom more exhaustive surveys, then the totalnative plant diversity of Victorian temperatewoodlands might be tentatively estimated asat least 1 100 species, which is 33% of the3 289 native plant taxa recorded from Victoria(Ross 1996).

Whilst considerable, this figure is notespecially high for an ecosystem that onceoccupied 40% of the state. By contrast, asimilar number of species (almost one-thirdof Victorian native plant species) occur inthe 167000 ha Grampians National Park inwestern Victoria (Calder 1987). The relativelylow plant species diversity across grassywoodlands in Victoria probably reflects theenvironmental uniformity of most woodlandhabitats, and low species turnover betweensites in most woodland regions. Most wood-land species were widespread before habitatdestruction, and few local endemics are knownto occur (Scarlett and Parsons 1993; Prober1996).

Despite a relatively low regional plantdiversity,small-scaleplant richness is extremelyhigh in some woodlands. Up to 45 vascularplant species have been recorded from a singlesquare metre, and 93 species from 128m2, inwoodlands in south-west Victoria (Lunt 1990).At this scale, these woodlands are among therichest terrestrial plant communities in theworld. Small-scale richness in these woodlandsis comparable to that of species-rich kwonganvegetation in south-west Western Australia

LUNT and BENNETT: TEMPERATE WOODLANDS IN VICTORIA 21

(Lamont et al. 1984) and chalk grasslands inEurope (Lloyd et al. 1971; During and Willems1984, see also Chapter 7).

Rare and threatened plant speciesThe status of threatened plant species in

Victorian temperate woodlands has been com-prehensively reviewed (Scarlett and Parsons1982, 1993; Department of Conservation andEnvironment 1992; Lunt 1992). Scarlett andParsons (1993) calculated the number of rareand threatened plant species in all majorbiomes in Victoria, including "grasslands,grassy woodlands and open forests of thefertile lowlands", which is largely synonymouswith "temperate woodlands" as defined here.These lowland grassy ecosystems contained 48nationally rare or threatened plant species(Table 3), about half the number in the biomewith the most rare or threatened species("shrubby open-forests of the ranges, heath-lands and shrubby open forests of the coastalplains", with 97 species), but similar to thatof alpine and sub-alpine c.ommunities (47species). However, lowland grassy ecosystemscontained many more nationally endangeredplant species than all other ecosystems (16 cf.8 in shrubby open forests and one in thealps). Lowland grassy ecosystems contained142 of 688 species (21%) which were rare orthreatened in Victoria (regardless of theirstatus elsewhere in Australia), and againincluded the greatest number of plant specieswhich were endangered in Victoria: 33endangered species compared to 31 in theMallee, and 14 in shrubby open forests(Table 3). Other reviews (Department ofConservation and Environment 1992; Lunt1992) showed essentially the same patterns:

temperate woodlands contain a relatively smallproportion of the total number of rare andthreatened plant species in Victoria, but asizeable proportion of those most at risk ofextinction in Victoria, the endangered species.

Most rare and threatened species intemperate woodlands had a wide range (andpresumably large total populations) prior toEuropean settlement (Scarlett and Parsons1982, 1993). Their present rarity is a functionof habitat destruction and unsuitablemanagement (especially grazing) in survivingremnants. Despite the large number ofthreatened species in these ecosystems, fewhave received specific ecological attention.Button Wrinklewort Rutidosis leptorrhynchoides isa notable exception, being perhaps the moststudied endangered plant species in south-eastern Australia (Gray 1979; Scarlett andParsons 1990; Leeton and Fripp 1991; Morgan1995a,b, 1997, 1999; Young et al. 1999).

The Victorian Flora and Fauna Guarantee Act(1988) allows listing of threatened species andcommunities. Threatened communities aredefined at a finer resolution than the "broadvegetation types" listed above. By June 1997,just one woodland community was listedunder the Act: Forest Red Gum Grassy Wood-land (in Gippsland). The paucity of listedwoodland communities reflects the difficulty ofnominating communities for listing (whichrequires data usually inaccessible to the public,such as detailed floristic descriptions), ratherthan the true conservation status of woodlandcommunities (Fig. 4). No Action Statementsfor woodland communities have appeared todate. Many threatened species in temperatewoodlands are listed under the Act, and

Table 3. Numbers of rare and threatened plant taxa in major ecosystems in Victoria.

Taxa rare or threatened in Victoria (but not necessarily in other states)Fertile lowlands 9 33 54 43Shrubby forests 9 14 81 142Tall wet forests 5 19 26Alpine 6 4 24 66Mallee 7 31 70 80Other 3 3 10 22

Conservation statusE V R

Taxa rare or threatened throughout AustraliaFertile lowlands 3 16Shrubby forests 5 8Tall wet forestsAlpineMalleeOther

Ecosystem type x

19222793

55953452

K Total

5 483 971 84 474 191 6

3 1427 253

501 1012 190

38

Threat codes: X = extinct, E = endangered, V = vulnerable, R = rare, K = poorlyknown. Ecosystem types: fertile lowlands = grasslands, grassy woodlands and open-forests of the fertile lowlands; shrubby forests = shrubby open-forests of the ranges,heathlands and shrubby open-forests of the coastal plains; tall wet forests = tall open-forests and closed-forests (rainforests); alpine = alpine and sub-alpine communities.Data from Tables 10.3 and lOA in Scarleu and Parsons (1993).

22 TEMPERATE EUCALYPT WOODLANDS IN AUSTRALIA

Action Statements have been published forwoodland species such as Hairy-pod WattleAcacia glandulicarpa (Venn 1996), Hairy AnchorPlant Discaria pubescens (Humphries 1993),Wedge Diuris Diuris dendrobioides Uohnsan1992, as D. cuneata), Small Psoralea Psoraleaparva (Muir 1991), Austral Toad-flax Thesiurnaus/rale (Scarlett et al. 1994), and others.

FAUNA OF TEMPERATE WOODLANDS

In parallel with the extensive destructionand modification to temperate woodlandvegetation in Victoria (Table 1), the fauna ofthis ecosystem has also experienced enormouschange over the past 150 years. Informationconcerning the occurrence and distribution ofthe vertebrate fauna of temperate woodlandsin Victoria is available from four main sources.

• Historical records and reports from earlysettlers (e.g., Dawson 1881; Curr 1883),from naturalists, and from specimen recordsin museums. Because most of the changesto vegetation (and hence to faunal habitats)occurred prior to systematic faunal surveys,these historical records are invaluable inreconstructing faunal assemblages and inassessing changes that have occurred (e.g.,Seebeck 1984; Bennett et al. 1989, 1998;Robinson 1993).

• Regional faunal surveyscarried out primarilyon public land to obtain information

Fig. 3 (righl): Herb-rich grassy woodland at Langi GhiranState Park in western Victoria. Dominant treesare Eucal),ptU5 camaldulensis River Red Gum andEucalyptus meliiodoTa Yellow Box. Species richnessof native plants is very high at the small-scale: atthis site, 79 plant species were recorded from asingle 16 m2 plot. Photo: Ian Lunt.

relevant to land use planning by the formerLand Conservation Council (e.g., LandConservation Council 1981, 1983). Regionalfaunal surveys have provided data forthe Western Plains (Emison et at. 1975,1978), Northern Plains (Bennett et at.1998), MaBee (Emison and Bren 1989;Robertson et at. 1989), Wimmera (Menk-horst and Beardsell 1982), Gippsland Plains(Norris el at. 1983) and East Gippsland(Land Conservation Council 1985).

Fig. 4 (left): The only known,intact remnant of Alloca5uarinaverticil/ata Drooping She-oakwoodland on the LowlandGippsland Plains in easternVictoria; a 0.4 ha rail-lineremnant at West Sale. Flowersinclude Burchardia umbel/ataMilkmaids (white), Bulbinebulbosa Bulbine Lily andChrysouphalum apiculatumCommon Everlasting (yellow).Photo: Ian Lunt.

LUNT and BENNElT: TEMPERATE WOODLANDS IN VICTORIA 23

• Faunal surveys and reviews of existingdata for local areas, for selected faunalgroups, or for environmental features ofinterest (e.g., Menkhorst and Gilmore 1979;Bennett 1982; Chesterfield et al. 1984;Middleton 1984; Seebeck 1984; Haddenand Westbrooke 1996; Robinson and Mann1996).

• Surveysand ecological studies of threatenedspecies that occur in temperate woodlands,such as the Superb Parrot Polytelis swainsonii(Webster and Ahern 1992), Bush Stone-curlew Burhinus ml!-gnirostris Oohnson andBaker-Gabb 1994), Grey-crowned BabblerPomatostomus temporalis (D. Robinson et al.,in prep), Squirrel Glider Petaurus norfolcensis(Menkhorst et al. 1988), and Striped LeglessLizard Delma impar (Coulson 1990)..

There is now an extensive data base ofdistributional records of vertebrate wildlifefrom temperate woodlands upon which landmanagers and the community can draw.However,while there is reasonable knowledgeof the distributional patterns of many verte-brate species (although data are limited forprivate land that comprises the majority of theformer temperate woodlands), there is littledetailed knowledge of the ecology of mostspecies or of the processes that influence theirpresent distribution and abundance.

Composition of the woodland fauna

Temperate woodlands span a broad rangeof elevations, rainfall zones and floristiccompositions, and consequently representa wide range of habitats for fauna. Wood-lands of Casuarina and Allocasuarina in thedry north-west, for example, have a faunadominated by species associated with theEyrean zoogeographic region of the inland,while montane eucalypt woodlands in theranges have a fauna with strong Bassianaffinities (Emison 1982). Thus, faunal assem-blages in temperate woodlands display muchvariation in composition and there is no singlewoodland fauna. In most areas, the faunalassemblage overlaps. substantially with that ofadjacent vegetation types (such as mallee, dryforests, moist forests), as well as includingsome characteristically woodland species.

The faunal component most distinctivelyassociatedwith woodlands in Victoria are thosespecies whose distribution is centred on theextensive woodland habitats of the WesternPlains, Northern Plains, and Wimmera.Discussion of woodland fauna in this chapterprimarily relates to temperate woodlands inthese regions, rather than that in southerncoastal or highland regions (e.g., montanewoodlands, rainshadow woodland). First, there

is a suite of species whose distribution inVictoria extends (or formerly extended) ina broad zone across the inland side ofthe Great Dividing Range. Examples includethe Yellow-footed Antechinus AntechinusflaviPes, Squirrel Glider Petaurus norfolcensis,Brush-tailed Phascogale Phascogale tapoatafa,Diamond Firetail Stagonopleura guttata, BushStone-curlew, Superb Parrot, Turquoise ParrotNeophema pulchella, Fuscous HoneyeaterLichenostomusfuscus, Black-chinned HoneyeaterMelithreptus gularis, Woodland Blind SnakeRamphotyphlops proximus, Dwyer'sSnake Unechisdwyeri, Olive Legless Lizard Delma inornataand Sloane's Froglet Ranidella sloanei (fordistributional maps see Emison et al. 1991;Coventry and Robertson 1991; Hero et al.1991; Menkhorst 1995).

A second group of woodland species inVictoria are those that occur (or formerlyoccurred) more widely in inland Australia,but in Victoria are primarily associated withdry woodlands in the north. These includespecies such as the Red-tailed PhascogalePhascogalecalura, Pig-footedBandicootChaeropusecaudatus, Bridled Nail-tail WallabyOnychogaleafraenata, Apostlebird Struthidae cinerea,Cockatiel Nymphicus hollandicus, Red-cappedRobin Petroicagoodenovii, Curl Snake Suta suta,Hooded Scalyfoot Pygopus nigriceps andBoulenger's Skink Morethia boulengeri.

Although not an integral part of the wood-land fauna, it is notable that the Western,Northern and Wimmera Plains also supportmany species that favour wetland habitats.The flat plains generally have limited drain-age and so numerous seasonal wetlands occurin low-lyingareas and depressions within thewoodland ecosystem (Corrick 1995).Adjoiningaquatic and terrestrial environments areimportant for frog species, and also for birdssuch as herons, egrets, ibis, and kingfishersthat feed in wetlands and roost or nest inwoodlands. A number of these wetlands aresignificant nesting areas for waterbirds; forexample, those in the riparian floodplains ofBarmah and Gunbower forests for colonialwaterbirds, and shallow freshwater marshesof the open plains for the Brolga Crusrubicundis (Land Conservation Council 1983;Chesterfield et al. 1984; Corrick 1995).

The structure of temperate woodlands andtheir climatic regime are two attributes thatinfluence the composition of the fauna.Temperate woodlands frequently have an opencanopy of large-crowned mature trees, anda relatively open ground layer dominatedby grasses and herbs with scattered shrubs.Tree hollows in mature trees provideimportant nesting sites for a suite of obligate

24 TEMPERATE EUCALYPT WOODLANDS IN AUSTRALIA

hollow-using species. Parrots and cockatoos(Families Psittacidae and Cacatuidae), anotably conspicuous and diverse group intemperate woodlands, use tree hollows fornesting and a number of these species foragein the ground layer for seed-bearing grassesand forbs, culms and bulbs. The Sulphur-crested Cockatoo Cacatua galerita, Long-billedCorella Cacatua tenuirostris, Galah CacatuaroseicaPilla, Eastern Rosella Platycercus eximiusand Red-romped Parrot Pseplwtus haematonotus,are common in most woodland regions(Emison et al. 1991) while, in the driernorth-west, woodlands provide nesting areasfor the Mallee Ringneck Barnardius barnardi,Blue Bonnet Northiella haematogaster, MulgaParrot Pseplwtus varius and Major Mitchell'sCockatoo Cacatua leadbeateri (Land Conser-vation Council 1987; Emison and Bren 1989).Riverine woodlands are also important nestingareas for these families, especially for thethreatened Superb Parrot and Regent ParrotPolytelis anthopeplus, which move betweenwoodlands and nearby foraging habitatsduring the breeding season (Burbidge 1985;Webster and Ahern 1992).

The open ground layer of temperatewoodlands provides foraging habitat for otherseed-eating species (e.g., Diamond Firetail,Peaceful Dove Geopelia striata, BudgerigarMelopsittacus undulatum, Crested Pigeon Ocyphapslophotes), as well as numerous ground-foragingbirds that feed on invertebrates (e.g., Red-capped Robin, Hooded Robin, White-wingedChough Corcorax melanoramphos, Apostlebird,Restless Flycatcher Myiagra inquieta, Yellow-romped Thornbill Acanthiza chrysorrhoa,Australian Magpie Gymnorina tibicen).

The most widespread woodlands occur indrier, warmer parts of the state, climaticconditions that are favourable to reptiles.There is a trend of increasing species richnessand taxonomic diversity of reptiles towardsthe warmer north-west of Victoria (Brownand Bennett 1995). Thus, temperate wood-lands generally support diverse assemblagesof reptiles, with a greater abundance andrichness in dry woodlands of the Wimmeraand Mallee regions. Families such as geckoes(Gekkonidae), legless lizards (Pygopodidae)and blind snakes (Ramphotyphlopidae) arepoorly, or not represented, in mesic forestedregions of southern Victoria, but are a moreprominent component of the fauna in warmdry woodland habitats (e.g., Brown andBennett 1995; Hadden and Westbrooke1996).

The geographic location and climate oftemperate woodlands make them an attractivedestination for migratory birds. The less-marked seasonal climate of the woodlands

contributes to a more reliable year-roundsupply of food than in the forested rangeswhere colder winter conditions contribute toseasonal food shortages (Nix 1976). Thisencourages winter migration of forest birdssuch as the Golden Whistler Pachycephalapectoralis, Crimson Rosella Platycercus elegans,White-naped Honeyeater Melithreptus lunatus,Pied Currawong Strepera graculina and FlameRobin Petroica phoenicea to the woodlands ofthe plains. In spring and summer, the wood-land avifauna is swelled by breeding migrantsfrom northern Australia, such as RufousWhistler Pachycephala rufiventris, Horsfield'sBronze-CuckooChrysococcyxbasalis, Tree MartinHirundo nigricans and Sacred KingfisherTodiramphus sanctus, and in some years byan unusually large influx of inland speciessuch as the Budgerigar, Cockatiel, BlackHoneyeater Certhionyx niger and White-browedWoodswallowArtamus superciliosus, dependingon climatic conditions in inland Australia.An additional group of migratory birds moveto and from woodland habitats in response tothe flowering patterns of eucalypts. Seasonalfluxes of nectarivorous birds track heavyflowering of Grey Box E. microcarpa, WhiteBox E. albens and YellowBox E. melliodora onthe plains, and also the prominent floweringof ironbarks and other eucalypts in adjacentBox-Ironbark forests (e.g., MacNally andMcGoldrick 1997).

Conservation status and historical changes tothe fauna

It is not possible to remove 92% of temper-ate woodlands (Table 1) without havingprofound effects on the biota. Not surprisingly,there have been major changes to thefauna, including global extinction of species,decline and loss at the regional scale, andimpoverishment and simplification of faunalassemblages in many localities (e.g., Seebeck1984; Environment Conservation Council1997; Bennett et al. 1998). Indeed, temperatewoodlands now have a greater number ofextinct, endangered, vulnerable or rare speciesthan any other broad ecosystem (aquatic,coastal, forests, mallee) in Victoria (Depart-ment of Conservation and Environment 1992).The decline in, and change to, the fauna ismost severe in areas formerly occupied byplains grassy woodland and grasslands on theWestern, Northern and Wimmera Plains. Incontrast, relatively little change has occurredto the fauna of temperate woodlandsassociated with the highlands of southernVictoria (e.g., rainshadow woodland, montanewoodland) where much habitat remains intact(Table 1). The loss of fauna in temperatewoodlands is not unique to Victoria, but is anissue of great concern throughout southern

LUNT and BENNETI: TEMPERATE WOODLANDS IN VICTORIA 25

Australia (Robinson 1993; Barrett et al. 1994;Saunders and Ingram 1995; Robinson andTraill 1996; Bennett and Ford 1997).

MammalsThe mammalian fauna of temperate wood-

lands has been massively altered by habitatchanges over the last 150 years. The ef!ectshave been particularly severe on medmm-sized (0.1-5 kg) ground-dwelling mammals,such as wallabies, rat-kangaroos, bandicoots,quolls and rodents, which have all butdisappeared (e.g., Bennett et al. 1989, 1998).At least 14 of the 19 species of mammalsknown to be extinct in Victoria (Conservationand Natural Resources 1995) are believedto have regularly occurred in woodlandhabitats. Four are now globally extinct (EasternHare-wallaby Lagorchestes leporides, LesserStick-nest Rat Leporillus aPicalis, Pig-footedBandicoot Chaeropus ecaudatus and White-footed Rabbit-rat Conilurus albiPes), whilethe others, such as Bridled Nail-tail WallabyOnychogaleafraenata, Rufous Bettong Aepyprymnusrufescens, Red-tailed Phascogale P'!asc~galecalura and Eastern Quoll Dasyurus vzvemnus,persist in greatly reduced ranges in otherstates (Menkhorst 1995). It is difficult forpresent-day observers to appreciate t~ediversity and composition of the terrestnalmammal fauna recorded by the first Europeanexplorers and settlers in these woodlands(Dawson 1881; Curr 1883; Seebeck 1984;Bennett et al. 1989, 1998).

Fewof the native ground-dwelling mammalsthat remain are common. Even widespreadspecies such as the Eastern Grey Kang~rooMacropus giganteus, Short-beaked EchidnaTachyglossus aculeatus and Common WombatVombatusursinus, are now scarce or absent fromnumerous districts in the former woodlandecosystem (Menkhorst 1995). One species,..t~eEastern Barred Bandicoot Perameles gunnu, ISthe focus of an intensive recovery effort thatis using captive breeding, re-int!oductio~s andhabitat management to restore Its status In thewild (Humphries and Seebeck 1995). Arborealmammals have generally fared better, but twotypically woodland species, the Brush-tailedPhascogale and Squirrel Glider, are rega~edas threatened in Victoria and face dauntIngconservation problems due to the paucity ofremaining woodland habitat (EnvironmentConservation Council 1997). Bats are thegroup most resilient to habitat change, andnow make up a large proportion of theextant mammal fauna (e.g., 45% of mammalspecies in the Northern Plain). Their mobilityallows them to move between disjunct areasof habitat to gain access to resourc~s f?rfeeding and breeding, and thus to maIntaIn

populations even in greatly disturbed environ-ments (Lumsden et al. 1995).

BirdsBirds are the most diverse and visible group

of vertebrates in temperate woodlands. Therehave been extensive changes in the status ofbirds but unlike mammals, there have notbeen who'lesale extinctions of species. How-ever. at least 38% (20/53) of bird species listedas e~tinct, endangered, vulnerable or rare inVictoria (Conservation and Natural Resources1995) regularly occur in woodland ecosystems.

Several types of changes to the woodlandavifauna can be recognized. First, there hasbeen a major reduction in the geographicrange of certain species as they have graduallydisappeared from areas where they onceoccurred. The Grey-crowned Babbler, forexample, was formerly widespread intemperate woodlands of the plains, but hassuccessively disappeared from the WesternPlains and much of the Wimmera and Malleeareas, and now persists in substantial numb~rsonly in limited areas on the Northern PlaInswhere there are highly connected syste~ns ofremnant habitat (Davidson and Robmson1992; Robinson et al., in prep.). Similar rangereductions are evident (and appear to beongoing) for other species such as the BushStone-curlew, Superb Parrot and RegentHoneyeater (Menkhorst 1994; Webster andBaker-Gabb 1994).

Second, there is an ongoing decline in therelative abundance and status of numerouswoodland-dependent species throughout theirrange in temperate woodlands of southernAustralia (Robinson 1993; Saunders andIngram 1995; Robinson and. Traill 199~).This is evident from changes m the relativeabundance of species in comparison withhistorical records for various districts, adecreased frequency of reports from localareas, and "gaps" in the distribution of specieswhere heavy clearing of woodlands hasoccurred. Many of the woodland-dependentspecies believed to be dec~ining (e.g., <?,ilbert'sWhistler Pachycephala znornata, DIamo~dFtretail Hooded Robin, Buff-rumped ThornbillAcanthi~a reguloides, Varied Sitella Daphoenosi",:chrysoptera, Black-chinned Honeyeater Melz-threptus gularis) are not yet regarded asthreatened on a statewide basis, but in termsof persistence in temperate woodl~nd regions,these are serious concerns (Robmson 1993;Bennett and Ford 1997).

Third, the extensive clearing of woodlandshas enhanced opportunities for birds of openhabitats and those tolerant of highly-disturbedhabitats, allowing them to expand their range

r26 TEMPERATE EUCALYPT WOODLANDS IN AUSTRALIA

and increase in abundance. Cleared, openfarmland is generally dominated by a handfulof common species such as Australian Magpie,Australian Raven Corvus coronoides, WillieWagtail Rhipidura leucophrys, Galah, Richard'sPipit Anthus novaeseelandiae, and introducedspecies such as the Common Starling Sturnisvulgaris and European Goldfinch Cardueliscarduelis. Even common woodland birds arescarce unless fragments of woodland havebeen retained. In many areas where patchytree cover is retained amongst farmland,the Noisy Miner Manorina melanocephala hasbecome abundant (Loyn 1985; Grey et al.1997). This aggressive native honeyeateractively excludes small insectivorous speciesfrom the habitats it occupies (Greyet al. 1997).Together with the decline of woodland-dependent species, the overall result has beenan impoverishment and simplification of thewoodland avifauna in many rural districts.

ReptilesReptiles are the most poorly surveyed and

least-known group of vertebrates in temperatewoodlands. Further, there are few baselinehistorical data with which to compare presentpatterns of occurrence. There is no evidencethat any woodland reptiles have becomeextinct in Victoria, but present patternssuggest that loss of woodland habitats hasgreatly altered the distributional pattern ofmany species (Environment ConservationCouncil 1997). Some 38% of reptiles regardedas threatened in Victoria (Conservation andNatural Resources 1995) have much or allof their distribution in woodland and grass-land ecosystems. While a few species are ableto persist in highly disturbed environments(e.g., Boulenger's Skink, Blotched BluetongueTiliqua nigrolutea, Grass Skink Pseudemoiaentrecasteauxii), most species of reptiles are nowpatchily distributed with isolated populationscentred on localized areas with suitablemicrohabitats for refuge, feeding and breeding(Bennett et al. 1998). Open grassy farmland,now the dominant vegetation type in theformer woodland ecosystem, supports few orno reptiles, and disturbed woodland remnantsamong farmland may also be poor habitat. Arecent survey in the Northern Plains (Brownand Bennett 1995), for example, recordedno reptiles at a third of sites (65/193) inwoodland fragments of varying size, shape andtree species composition.

Amphibians

Most species of frogs are closely associatedwith streams or wetlands of various types,rather than woodland habitats per se. Frogsneed moisture to breed successfully and so

the availability of water, the soil type, andaccess to microhabitats such as logs, rocks orexfoliating bark that provide refuge fromdesiccation, are likely to be more importantinfluences on these species than the structureor composition of woodland vegetation. Mostspecies appear to be capable of persistingin modified woodland· environments, andsome species in cleared farmland, wherethese requirements for moisture and shelterare met. Artificial water sources associatedwith agriculture, such as farm dams andirrigation channels, are extensively used bycommon species of frogs (e.g., CommonFroglet Ranidella signifera, Spotted MarshFrog Limnodynastes tasmaniensis) and maysupport high density populations (Bennettet al. 1998).

Two species of frogs regarded as threatenedin Victoria (Conservationand Natural Resources1995), the Giant Bullfrog Limnodynastesinterioris and Barking Marsh Frog Limnodynastesfletcheri occur in the woodland ecosystem(20% of listed threatened amphibians). Theiroccurrence in northern Victoria forms thesouthern part of a more extensive range ininland New South Wales (Hero et al. 1991).

OUTLOOK FOR TEMPERATEWOODLANDS IN VICTORIA

Conservation of the flora and fauna oftemperate woodlands in Victoria poses aformidable challenge because this ecosystemhas already experienced profound change.Remaining natural areas are mostly distributedas fragments amongst intensively settledrural lands and occur as thousands of patchesin farmland, as strips along roadsides, streamsand rail reserves, and as occasional smallconservation reserves. Many woodland frag-ments are on private land or areas of publicland that frequently are licensed to privatelandholders (e.g., stream frontages, unusedroads, bushland reserves). In some areas,remnant woodlands are virtually the onlypublic land, leading to land use conflicts andprobable future losses of biodiversity. Thebiodiversity values of woodland remnantsare threatened by a wide array of processes.Some of these threats, such as rising salinewater tables, weed invasions, tree dieback, andstock grazing operate at the landscape scale,while others, such as fertilizer drift, road-works, quarrying, cropping, pasture improve-ment, grazing by native herbivores,inappropriate burning regimes, soil disturb-ance, ploughing, grading, rubbish dumpingand recreational damage, are usually localizedin effect (Lunt 1991; Department of Conser-vation and Environment 1992; Robinson andTraill 1996).

LUNT and BENNETT: TEMPERATE WOODLANDS IN VICTORIA 27

Three broad elements are particularlyimportant considerations in the future con-servation of the biota of temperate wood-lands: protection of existing natural areasto prevent further degradation and loss;management of habitats to enhance conser-vation values; and long-term restoration ofthe ecosystem.

First, protection of existing natural andsemi-natural habitats is essential to minimizefurther decline in the status of plant andanimal populations. Planning controls inVictoria have reduced the rate of tree clearingin the state but, because so little remains,broadscale clearing is rarely a major issue intemperate woodlands. Rather, incrementalclearing, hectare by hectare, and gradualdeterioration of woodland habitats and lossof species by degrading processes, are themain ways in which temperate woodlandscontinue to decline. There is a need for newapproaches and strategies for conservation ofwoodland habitats. The traditional approachof achieving representative conservationthrough reservation of large areas of publicland as parks and reserves is not now possible.Because most remnant woodlands in Victoriaare in landscapes that are managed primarilyfor purposes other than nature conservation,strategies to conserve woodlands must bemulti-faceted and incorporate education,extension, community input, and research, aswell as the traditional approach of surveysandreservation (e.g., Prober and Thiele 1993;Ross 1995; Binning 1997).

Second, active management is requiredto ensure that remaining habitats achievetheir potential for conservation. A particularchallenge in this regard is to arrest furthermodification and degradation of habitats thatso easily occurs through oversight, neglect,or perpetuation of land use practices thatare not ecologically sustainable. Obviousexamples include: excessive grazing by stockleading to loss of native plants and lack ofregeneration, excessive firewood collection,invasion and spread of weeds, altered nutrientlevels, and loss of mature trees. Often,processes such as understorey clearance,addition of fertilizers, alteration to drainagepatterns, gross soil disturbance and damagefrom recreational use, pose the mostimmediate threats for many remnants andspecies.Manyof these processesare avoidable,although not necessarily easy to alter. Littlework has been undertaken on ranking threatsto woodland remnants, either in terms ofubiquity, potential damage or irreversibility,and so assessment of the relative importanceof various threatening activities is largelyobservational.

The woodland fauna depends upon sym-pathetic on-site management to maintainhabitat componentsand required resources(e.g.,diverse ground and litter layer, logs, shrubs,large trees with hollows). However, manage-ment of temperate woodlands for fauna mustalso adopt a landscape and regional perspec-tive in order to strategically maintain systemsof habitat. To maintain viable populations,many species require much greater areas ofhabitat than are available in single isolatedwoodlands, while other species (notably batsand some birds) regularly move between anumber of woodland patches to obtain neces-saryresources.Consequently,effectiveconserva-tion of the woodland fauna generally requiresextensive interconnected systems of habitatthroughout the landscape (Bennett et at. 1998).

Last, because so much change has alreadyoccurred, there is a need for long-termstrategic, and visionary, restoration of thetemperate woodland ecosystem. In recentyears, a number of strategies have appearedin Victoria from government agencies (e.g.,Department of Conservation and Environment1992) and non-government bodies (e.g.,Robinson and Traill 1996), and a range ofimportant initiatives is under way. Theseinclude, for example, ongoing support forprotection and restoration of natural habitatsby landholders through the Land for WildlifeScheme (Platt and Ahern 1995), a majorreview of resources and land use on publicland in box and ironbark forests and wood-lands across central Victoria (EnvironmentConservation Council 1997), and numerouslocal actions to protect and expand habitatsof threatened species such as the SuperbParrot (Raven 1995) and Brolga (McIntyre1995).The recent publication of a biodiversitystrategy for Victoria (Department of NaturalResources and Environment 1997a, 1997b,1997c) outlines a broad direction for conser-vation and identifies a further range ofinitiatives that have the potential to makea major contribution to conservation oftemperate woodlands in the state.

The time frame for restoration will, ofnecessity, be long - decades and centuriesrather than years. There is growing com-munity support for an integrated approachto sustainable land management in ruralenvironments, in which nature conservationand productive use of land for agriculture areboth viewed as important and complementarygoals. However,if restoration of the temperatewoodland ecosystem is to be achieved, itrequires a sustained politicalwill and enhancedcommitment of resources to implement con-servation strategies that do more than simplyslowthe tide of decline.

28 TEMPERATEEUCALYPTWOODLANDSIN AUSTRALIA

ACKNOWLEDGEMENTSBill Peel (Department of Natural Resources

and Environment) and Steve Farrell (NaturalResource Systems) kindly provided unpub-lished data on the extent of broad vegetationtypes in Victoria, and Annette Muir and PaulForeman (DNRE) provided unpublishedquadrat data from grassy woodlands. Researchon the fauna of woodlands in Victoria overrecent years has been generously supportedby the Department of Natural Resourcesand Environment (Arthur Rylah Institute), theLand and Water Resources Research andDevelopment Corporation (Project DUV2) andthe Australian Research Council (Grant No.A19531268). Helpful comments on themanuscript were provided by two referees.

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TelDperate Eucalypt Woodlandsin Australia

Biology, Conservation, Managementand Restoration

Edited by

Richard J. Hobbs and Colin J. Yates

Published by

Surrey Beatty & Sons