Distribution of Clonal and Non-clonal Wetland Plants at Clydebank Morass, Gippsland Lakes, in Relation to Elevation and Salinity Gradients

2008·2.4 MB·English

Research Reports
Distribution ofclonal and non-clonal wetland plants
at Clydebank Morass, Gippsland Lakes,
in relation to elevation and salinity gradients
Matthew J Hatton1, Paul 1 Boon 1,2and Randall W Robinson1
InstituteforSustainabilityandInnovation,StAlbanscampus,VictoriaUniversity
P.O.Box 14428,MelbourneCityMailCentre,Victoria8001
:Authorforcorrespondencee-mail:paul,[email protected]
Abstract
A review ol the published literature suggested that plantswith aclonal growth habit dominated the
vegetation ofwetlands in manyparts ofthe world. Totestwhetherthispattern held inAustralia, the
distribution ofplantswith clonai and non-clonal growth habits wasexamined in Clydebank Morass,
a brackish-water wetland ofthe Gippsland Lakes in south-eastern Victoria. Nineteen ofthe twenty
speciesofaquaticorsemi-aquaticplantspresent in thewetlandwereclonal. Intermsofbothspecies
numberandpercentagecover,clonal plantsdominatedthevegetation in wetand intermittentlydamp
parts ofthe Morass whereas non-clonal plants were progressively more common as elevations
increased.Thiselevationaleffectwasduemoretochangesin soil moisturecontentthan in soilsalin-
ity. These results not only confirmed the prediction that clonal plants were the dominant growth
habit in the wetland but were consistentwith predictions made in the 1960s as to likely vegetation
changes as the Gippsland Lakes became progressively salinised. Understanding the dominance of
wetlands by clonal plants has implications forassessments ofplant fitness and the maintenance of
plant biodiversity and habitat heterogeneity; it is central also to improving the success with which
degradedwetlandsarerehabilitated.(TheVictorianNaturalist125(1),2008, 11-18)
Introduction
Clonal plants are those that can reproduce clonal. Clonal plants also have been shown
asexually and spread as a population of to dominate the floristics of wetlands in
semi-independent parts ofwhat was origi- Central Europe (Klimes et al. 1997; L.
nally a single seedling (Harper 1977). Klimes, pers. comm. 2005) and China
Clonal growth and vegetative reproduction (Songand Dong2002).
may occur in many ways, including the There have been no comparable studies
production of rhizomes, stolons, tubers, undertaken to examine the importance of
turions and plantlets, fragmentation of the clonality in the vegetation ofwetlands of
plant body, and node rooting. Many of temperate south-eastern Australia, but a
these features are evident in well-known review ofplant descriptions in the three
andwidely distributed wetland andaquatic most important monographs on Australian
plants - for example willows Salix spp., aquaticandwetlands plantsinthisregion-
Eel Grass Val/isneria spp., Common Reed Aston (1977), Romanowski (1998) and
Phragmitesaustralis andCumbungi Typha Sainty and Jacobs (2003) - indicated that
spp. Indeed, a number of international 288 ofthe 450 described taxa (64%) were
studies have indicated that wetlands are clonal and capable ofvegetative reproduc-
often dominated by clonal plants, which tion. The ratio of approximately 2:1 for
can reproducevegetativelyaswell as sexu- clonal to non-clonal aquatic plants was
ally by seed when environmental condi- consistent across all three references: 65%
tions permit (Miihlberg 1982; Cook 1990; in Aston (1977), 61% in Romanowski
Grace 1993; Obomy and Bartha 1995; van (1998) and 71% in Sainty and Jacobs
Groenendael et al. 1997; Crow and Barre (2003). This result would suggest that
Hellquist 2000). For example, 61% and Australian wetlands, like wetlands in the
68% of the aquatic or wetland plant Northern Hemisphere, are likely to be
species in North Carolina and Florida, dominated by clonal plants. Nevertheless,
respectively, are clonal (Beal 1977; Tarver to our knowledge there have been no stud-
et al. 1978). Spencer Jones and Wade ies that have attempted empirically to
(1986) reported that 88% of vascular quantify the incidence ofclonality- either
aquatic plantspecies in the British Isles are floristically or in terms ofcover or abun-
Vol. 125 2008
(1) 11
Research reports
dance - in an Australian wetland. for waterbird breeding and hunting and
Moreover, it is uncleareven from overseas habitat values. The wetland has been con-
studies as to what environmental factors nected permanently to Lake Wellington
are most important in controlling the rela- sinceabout 1990, when high waterlevels in
tive distributions of clonal and non-clonal the lake breached the low-lying boundary
plants in wetlands. betweentheMorassandLakeWellington.
The research reported in this paper had
threeaims. The firstwastotestthehypoth- Vegetationsurveys
esisthat clonal plants dominatedthe aquat- Floristic surveys were undertaken in late
ic and semi-aquatic vegetation in summer (March 2004) and winter (June
Clydebank Morass, a brackish-water wet- 2004) to quantify the relative proportions of
land of the Gippsland Lakes in eastern clonal and non-clonal species. These two
Victoria. This question was addressed by surveyperiods were chosen to maximisethe
undertaking vegetation surveys attwo con- number ofplant species identified in the
trasting times ofyear to quantify the rela- wetland, within the constraints ofthe time
tive incidence ofclonality in the wetland’s available for field-based assessments. To
aquatic and semi-aquatic vegetation. The determinewhethertheratioofclonal tonon-
second aim was to determine whether the clonal taxa varied with elevation, 20 belt
distributions ofclonal and non-clonal plant transects (50 cm wide) were run in July
species varied with environmental condi- 2004 from thewater-line to the most elevat-
tions, especially elevation and soil water ed level where terrestrial vegetation bor-
content and salinity. The third aim was to dered the Morass. Transects varied in length
use these data on floristics, vegetation from50to 100mandweredivided intocon-
m
cover and environmental conditions to tiguous 1 long quadrats. Transects were
address a number ofpredictions made in positioned systematically rather than ran-
the 1960s as to the likely historical trajec- domly because systematic samples are par-
tory ofplant communities in wetlands that ticularly well-suited to detecting vegetation
fringe the Gippsland Lakes, with particular changes along environmental gradients and,
reference to increasing salinity. In a series in such cases, random sampling would gen-
of pioneering papers by ECF Bird in the erate an arbitrary set ofvalues for species
1960s (Bird 1961, 1962, 1966), it was pre- abundance(Kershaw 1973).
dicted that wetlands and fringing vegeta- Plant cover for vascular plant species in
tion along the Gippsland Lakes would be each quadrat was estimated using the
affected by increasing salinity arising from Braun-Blanquet scale(0= 1-5% cover; 1 =
the artificial opening at Lakes Entrance. 5-10%; 2 = 10-25%; 3 = 25-50%; 4 = 50-
Bird (1966) specificallyraisedthepossibil- 75%; and 5 = 75-100%). An inclinometer
ity that inexorably increasing salinity was used to estimate transect elevations:
would cause the replacement of reed elevations are given in terms of vertical
swamp (Common Reed Phragmites aus- height above the water level and not with
tralis, then known as P. communis by reference to the Australian Height Datum.
swamp scrub (Swamp Paperb)ark This decision was necessitated by the lack
Melaleuca ericifolia and eventually ofsurvey data or a digital elevation model
swamp scrub by even )more salt-tolerant forthe site. Using known measurements of
saltmarsh communities. transect lengths and differences in eleva-
tion (calculated from inclinometer read-
Methods ings), each ofthe 20 transect gradients was
Fieldsite broken into 0.25 m elevations to allow the
Clydebank Morass (38°02’50”S, zonation width of different taxa to be
147°14’00”E) is a shallow (zmax = 1-2 m), mapped in terms of elevation above the
brackish-water wetland of about 1420 shoreline. Plant nomenclature follows that
hectares in Gippsland, Victoria. It is one of ofRossand Walsh (2003).
a chain of Ramsar-listed, brackish-water
wetlands along the perimeter of Lake Soilcharacteristics
Wellington, the westernmost waterbody of Soil cores were taken in September 2004
the Gippsland Lakes. Clydebank Morass is to determine whether there were consistent
a State Game Reserve, managed primarily variations in soil water or salt contents
12 The Victorian Naturalist
Research reports
along the elevational gradients or from Table 1. Incidence ofclonality in plants at
zones with contrasting vegetation along a Clydebank Morass, Gippsland Lakes, south-
single transect. Three replicate soil cores easternVictoria.
(5 cm diameter x 10 cm deep) were taken %
at each quadrat; in total 228 cores from 76 Characteristic Noof
different quadrats were analysed. Soils species
were sampled to only 10 cm because the Totalnumberofplantspecies 90 100
species present, even Swamp Paperbark, Nativespecies 58/90 64
are quite shallow rooted. Soil cores were Exoticspecies 32/90 36
oven driedat 105°C to determine moisture NCaltoniavlecslpoencailesspecies 4354//9508 5509
contents (MC%, expressed as a percentage Putativeguerillagrowthhabit 24/45 53
ofdry-soil weight). To determine salt con- Putativephalanxgrowthhabit 21/45 47
tents, subsamples were groundwith amor- Aquaticspecies 20/90 22
tar and pestle, made into slurries (1:5 Clonalaquaticspecies 19/20 95
soikwater), then shaken by inversion for
onehourat25°C (Raymentand Higginson Of the 20 species found in aquatic or
1992). The temperature-adjusted electrical semi-aquatic environments in Clydebank
conductivity ofthe slurries was converted Morass, 19 were easilyclassified as clonal.
tvoerssailotncofnaccteonrtroafti0o.n6sth(emngtLab'u)luastiednginaucnoint-s TsphuerroenyeSrpeermgauilnairnigasmpeecdiieas,wGarseaatnerexSoetai-c
ofgL' ofsoil porewater. wnahteirveeittotyEpuircoalpley agnrdowNsorasthaApmeererninciaa,l
Results above the water-line. At this study site,
Relative proportions ofclonal and non- however. GreaterSea-spurreywas found at
clonalspecies the water-line and displayed a growth and
Ninety plantspecies from 31 plantfamilies senescence cycle typical of an annual
were identified during thetwo floristic sur- species. As there was no evidence ofnode
veys. Exactly one halfofthe species pre- rooting or other asexual growth, Greater
sent were clonal, though this figure Sea-spurrey was classified as non-clonal.
increased to 59% when only native taxa Accordingly, 95% of the wetland plants
were considered (Table 1). Native species present in Clydebank Morass were unam-
accounted for 64% ofthe species and non- biguously clonal.
natives 36%; the non-natives were mostly
introduced annuals. Guerilla and phalanx Variations withelevationandsoilsalinity
clonal growth habits (sensu Lovett-Doust Clonal species completely dominated the
1981) wereboth well represented at 53 and vegetation ofClydebank Morass at lower
47%, respectively. Clonal plants with a elevations, and the proportion of clonal
guerilla growth habit tend to have long species declined consistently with increas-
stolons or rhizomes, which allow them to ing elevation (Fig. 1). Clonal species
penetrate deeply into new habitats. In con- accounted for more than 70% ofthe total
trast, plants with the phalanx growth form numberofspecies in the first metre ofver-
tend to have short stolons or rhizomes and tical elevation and approximately 50% of
colonise new habitats on a broad front, the species up to an elevation of2.5 m. Atele-
metaphor reflecting a massed body of vations greater than 3 m from the water-
infantry drawn up in close order and line, the number ofclonal taxa declined to
advancingslowly intonew areas. Ofthe 90 around 20% and remained at this value
species present in themorass, only20 were until the highest elevation (6 m) examined
classified as aquatic or wetland plants in this study. Thedominance ofclonal taxa
(Table 2). The definition ofaquatic plants was evident not only in terms of floristic
was fairly broad and followed the descrip- criteria (Fig. 1) but also in terms ofper-
tions given by Best (1988) and Sainty and centage cover: up to an elevation ofabout
Jacobs (2003). This definition allows for 2 m, clonal plants provided a cover of
the accommodation ofwoody tree species, >75%(Fig. 2).
such as Swamp Paperbark and willows, as The distribution of individual plant taxa
well asherbaceoustaxa. also varied with elevation. Common Reed
Vol. 125(1) 2008 13
Research reports
Tabic 2. Range ofclonal attributes possessed by wetland plants at Clydebank Morass, Gippsland
mLaakressh,psloauntth-beyasBtreirdngeVwiacttoerriae.ta*li(n1d9i8c1a)t.esan exotic species, “indicatesaspecies identifiedas a salt-
Family Speciesname Common name Clonalattribute
Aizoaceae DisphymacrassifoliumSM PurpleNoon-flower Noderoots
Asteraceae CotulacoronopifoliaSM Water-buttons Noderoots
Leptinellalongipes CoastCotula Stolons
Caryophyllaceae Spergulariamedia* GreaterSea-spurrey Non-clonal
Chenopodiaceae SarcocorniaquinquefloraSM BeadedGlasswort Stolons
Cyperaceae Baumeaarthrophylla SoftTwigrush Rhizomes
Botboschoenuscaldwellii SeaClub-rush Rhizomes
Eleocharisminuta* VariableSpike-rush Stolonorrhizomes
Eleocharispusilla SmallSpike-rush Rhizomes
IsolepisnodosaSM KnobbyClub-rush Rhizomes
Goodeniaceae SetHeraradicansSM ShinySwamp-mat Noderoots
Juncaceae Juncuspallidus PaleRush Rhizomes
Juncuskraussiisubsp. SeaRush Rhizomes
australiensisSM
Juncaginaceae TriglochinstriatumSM StreakedArrow-grass Rhizomes
Triglochinprocerum Water-ribbons Tubersandrhizomes
Myrtaceae Melaleucaericifolia Swamp Paperbark Suckersandrhizomes
Poaceae Phalarisaquatica* ToowoombaCanary- Rhizomes
grass
Phragmitesaustralis Common Reed(Djarg) Rhizomes
Scrophulariaceae Mimulusrepens CreepingMonkey-flower Noderoots
Typhaceae Typhadomingensis Cumbungi, Bulrush Rhizomes
Phragmites australis for example, was nificantly different (one-way ANOVA:
,
tpiroensaelntgroadnileyntast,tuhpe twoetatbeosutte0n.d5 omfeelleevvaa-- aEc5r,7o4ss=th1e.4t7r4a,nsPect>s.0.T0h5e)hwiigthhestelaevveartaigoen
tion (data notshown). In contrast, asecond soil salinity was recorded beneath Swamp
set of plant taxa, consisting of Sea Rush Paperbark(21 gL1) in thepeatyephemeral
Juncus kraussii. Purple Noon-flower ecotone, whereas the lowest average soil
DanidspBhlyumeaTcurassssoicfko-lgiurma,ssSwPaomappoPiafpoerrmbiasr,k Csaolimnmitoyn(1R7eegd,La1t) wthaeswreetcteosrtdeedndbeonfeatthhe
was present in the ephemeral ecotonal gradient. Because ofthe nature ofthe con-
zone with mean elevations extending to version from electrical conductivity to
approximately 1.25 m above the water salinity, these salt concentrations should be
level. Vegetation in the highest, and driest, regardedasonlyapproximate.
zone consisted of Australian Salt-grass
Distichlis distichophylla and exotic annual Discussion
species, mostlypastureescapes. Oursurveys atClydebankMorass indicated
This pattern of plant distributions was that 19 ofthe 20 aquatic or semi-aquatic
related more to variations in soil moisture species present were clonal, and the single
content than the salinity ofthe soil pore remaining species that was not clonal,
water. There was a gradual decrease in the Greater Sea-spurrey, was an exotic native
mean soil moisture content from the to Europe and North America, where it
wettest zone colonised by Common Reed typically grows as a perennial above the
(93% w/w soil-moisture content) and Sea water-line. This empirical analysis con-
Rush (85%) to values of 61% in soils firmsthepattern presented in the main tax-
beneath Blue Tussock-grass stands and onomic monographs covering wetland and
50% under Purple Noon-flower. A one- aquatic plants in southern Australia (Aston
way analysis of variance (ANOVA) 1977; Romanowski 1998; Sainty and
showed a significant difference (F = Jacobs 2003), as well as a large number of
9.138, P < 0.0001) in mean soil-moi5st74ure comparable studies undertaken on the inci-
content across this gradient. In contrast to dence ofclonality in wetlands in Europe,
the results obtained with soil-moisturecon- theAmericasandAsia(see Introduction).
tents, average soil salinities were not sig-
14 The Victorian Naturalist
Research reports
Fig. 1. Percentage ofclonal plants, as a proportion oftotal species, in relation to elevation at
Clydebank Morass, Gippsland Lakes, south-eastern Victoria. Means and standard errors are shown,
n=20.
Fig. 2. Percentage coverofclonal plantspecies along 10 elevational gradients atClydebank Morass,
Gippsland Lakes, south-eastern Victoria. The Braun-Blanquet scale is used as the index ofcoveron
they-axis(0=1-5%cover; 1 =5-10%;2= 10-25%;3=25-50%;4=50-75%;and5=75-100%).
Vol. 125(1) 2008 15
Research reports
It seems that Clydebank Morass is similar notrestricted to a single season orraretime
to wetlands in other parts ofthe world in ofyear having the requisite water regime.
that its aquatic and semi-aquaticvegetation Furthermore, as seedordiasporeproduction
is overwhelmingly clonal. The dominance is often a function ofbiomass, long-term
ofclonal plants has a number ofimportant persistencethrough clonal offspringoffersa
implications for the understanding and greaterprobability ofcontinual seed output,
rehabilitation ofwetland vegetation. First, sothatviableseedwill bepresentwithinthe
it has implications for assessing plant fit- wetland system if suitable conditions for
ness. Theoccurrenceofseeds is commonly germinationdooccur.
used as a measure offitness for plants, as Second, the ubiquity ofthe clonal growth
seed production through outcrossing habit has implications for the production
ensures genetic recombination and creates and maintenance of habitat diversity in
a diverse and resilient gene pool (Eriksson brackish-water wetlands. Large clonal
1997). Although fitness istraditionally cal- plants, such as Swamp Paperbark, that
culated by forecasting the expected seed grow mostly at the lower elevations ofthe
output ofa plant over its lifetime, estimat- Morass provide more structural and topo-
ing the lifetime seed production for clonal graphical complexity in the landscape than
plants can be very difficult as these plants do non-clonal species. At high elevations,
are potentially immortal and may produce structural complexitymaybedriven largely
innumerable offspring in the form ofclon- by season, as temporarygaps are created in
al propagules and ramets, as well as seeds the vegetation with the rapid germination,
(Wikberg 1995; Santamaria 2002). Indeed, maturation and death ofannual species. In
Pan and Price (2002) argued that an contrast, at lower elevations tall phalanx
emphasis on seed production has meant species such as Common Reed, Sea Rush
that clonal growth has been ignored as a and Swamp Paperbark border one another
critical componentofplantfitness. and are often carpeted beneath by low-
Botanists have traditionally seen clonal growing guerilla species, such as Streaked
growth ability as an adaptation to comple- Arrow-grass Triglochin striatum, Shiny
ment the dispersal ofseeds (Grace 1993). Swamp-mat Selliera radicans, Creeping
There are a number ofreasons indicating Monkey-flower Mimulus repens and
that this view should be reversed when Purple Noon-flower. The combination of
wetland plants are considered, and that the tall phalanx and shortguerillaclonal archi-
obverse view - that sexual reproduction tecture creates a rich mosaic of structure
plays the complementary role to vegetative andmicrotopographicalrelief.
reproduction - should be erected in its Finally, the prevalence ofclonality has
place. In wetlands, recruitmenteventsfrom implications for wetland rehabilitation.
seed are irregular and highly unpredictable Seed banks feature prominently in the sci-
and seed germination is often dictated by entific literature on wetland revegetation
subtle changes in hydrological conditions and rehabilitation (e.g. Brock and
(Brock and Britton 1995; Rea and Ganf Casanova 2000). If, however, many wet-
1994; George et al. 2005; Capone and land plants recruit, disperse and colonise
Brock 2006). This means that conditions new areas primarily by vegetative means,
suitable for sexual recruitment may occur wetland managers will have to reconsider
atrelativelyfewtimes oftheyearor, in the the strategies for rehabilitating degraded
case ofwetlands with variable hydrologi- wetland sites. Three issues seem most
cal regimes, only in a few years everycou- important and their resolution may help
ple of decades. Moreover, the seeds of redress the general failure ofwetland reha-
even dominant wetland plants may be bilitation in southern Australia reported by
short-lived and this can prevent the forma- de Jong (1997). First, seed may not be the
tion ofa soil seed bank; such a pattern has most appropriate mechanism for re-intro-
been demonstrated recently for Lignum ducing plants into degraded sites:
Muehlenbeckiaflorulenta by Chong and Chambers et al. (1995) noted that vegeta-
Walker (2005). In contrast to sexual repro- tive recolonisation (e.g. with rhizomes)
duction, the period suitable for vegetative was the best form ofpropagation for a
growthandpropagation bywetlandplants is wide range ofimportant wetland plants in
16 The Victorian Naturalist
Research reports
south-western Australia, including Jointed Southern Ocean by a shifting and intermit-
Twig-rush Baumea articulata. Bare Twig- tent outlet through the sand barriers
rush Baumeajuncea. Pithy Sword-sedge between Cunninghame and Red Bluff at
Lepidosperma longitudinale and River the easterly part of Lake King. Although
Club-sedge Schoenoplectus validus. they would open to the sea during large
Second, the spatial heterogeneity created floods, to improve navigability an artificial
by large clonal plants may provide oppor- entrance was cut to the ocean in 1889 at
tunities for improving the success of Lakes Entrance, about 5 km from the nat-
revegetation trials. In earlier field trials the ural entrance. It is believed that a major
survival of Swamp Paperbark tubestock consequence of opening the artificial
was improved markedly by planting entrance has been to increase the salinity
seedlings into raised hummocks of Water ofthe Gippsland Lakes, which previously
Couch Paspalum distichum (Raulings et were relatively fresh, being fed by the
al. 2007). This experimental finding is rivers flowing into Lakes Wellington and
consistent with field observations that King and having only an intermittent link-
Swamp Paperbark seedlings and juveniles age with the ocean. Bird (1966) predicted
occur mostly on hummocks that offer that, because ofthis progressive and inex-
respite against the stressful combination of orable increase in salinity, the fringing
waterlogging, salinity and soil acidity vegetation of the Gippsland Lakes would
occurring in surrounding sediments. Third, change as salt-intolerant taxa, such as
the incidence ofclonality may force a re- Common Reed, were replaced by taxa tol-
assessment of the number of plants that erant of the higher salinities, such as
need to be introduced per unit area ofwet- Swamp Paperbark and, ultimately, salt
land during revegetation trials. Taking a marsh species. The findings of our study
lead from terrestrial precedents, revegeta- indicate not only that soils in Clydebank
tion trials in wetlands commonly involve Morass were highly saline (17-21 g L1 at
planting on close (e.g. 1 m) centres. Since the times of sampling in 2004) but that
many wetland plants are not only clonal taxa typical of salt marsh communities
but have short-lived seed (e.g. willows: were abundant in the wetland. Table 2
Sainty and Jacobs 2003; Lignum: Chong identifies those species present in the
and Walker 2005), most ofthe plants ofa Morass that were listed in the monograph
given species in a wetland were probably on salt marshes of southern Australia by
derived from oneorfew individuals. Itmay Bridgewateretal. (1981): itis obviousthat
be more appropriate to plant large, long- a large number ofthe taxapresent in 2004
lived clonal species at far lower densities were, indeed, salt-tolerant salt marsh repre-
than is the current common practice and let sentatives. Thusthedatapresentedhere are
them develop into discrete but extensive consistent with the historical trajectory
stands over the subsequent decades. This predicted for fringing and wetland vegeta-
recommendation is, however, not to under- tion along the shores of the Gippsland
estimate the importance ofthe prodigious Lakes by Bird (1966). Further historical
amounts of seed that can be produced by analysis is required to determine whether
some wetland taxa, e.g. Typha spp., and these taxa were present before salinities in
the problems this causes for the spread of the Gippsland Lakes apparently commen-
unwanted species into newly constructed cedtheirrecentincrease.
orrehabilitatedwetlands.
prAosbawbellylwaisdetlhyesaeppilmipclaibcleataicornosssthsaotutahr-e AWceknthoawnlkedPagrekmseVnitcstoria for allowing us to
undertake fieldwork at Clydebank Morass.
eastern Australian wetlands, the research Thanks particularly go to Andrew Schulz (Sale
findings throw light also on the predictions Office) and Manfred Ratzmann (Loch Sport
made over 40 years ago by ECF Bird Office). Thanks are also extended to Leos
(1961, 1962, 1966) for fringing and wet- Klimes (Institute ofBotany, Dukelska, Czech
land vegetation ofthe Gippsland Lakes. Republic)forhelpwith European flora interpre-
When the Gippsland region was first set- tfaiteilodnwso,rka.nd to Sharyn Leahy forassistance with
tled by Europeans in the 1840s, the
Gippsland Lakes were linked with the
Vol. 125 (1)2008
17
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Research reports
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