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The response of rare floodplain plants to an environmental watering event at Hattah Lakes, Victoria.


In 2014 and 2015, botanical surveys were undertaken at Hattah Lakes, Hattah-Kulkyne National Park, in semi-arid north-western Victoria, to study the response of understorey vegetation to a strategic environmental watering event. During the surveys, 23 plant species listed on the Victorian Government Advisory List of Rare and Threatened Species were recorded; with two of these species also listed under the Flora and Fauna Guarantee Act 1988 (Vic). Four of these species were recorded for the first time in the Park, and for a further 11 species the number of records were, at least, doubled within the Park. The 23 rare species represented five Water Plant Functional Groups. The cover of two functional groups and two species were significantly different before and after inundation. Although species and functional group responses to inundation differed, a key outcome from this study was the increase in rare species records for Hattah-Kulkyne National Park. (The Victorian Naturalist, 134 (1) 2017, 19-27)

Keywords: environmental watering, water plant functional groups, threatened flora


The Hattah Lakes complex, which forms part of the Hattah-Kulkyne National Park (480 km(2)), is located on the floodplain of the Murray River, in north-western Victoria (Murray-Darling Basin Authority (MDBA) 2012). It lies in the southern section of the Murray Basin, a major geological unit in south-eastern Australia that covers approximately 300 000 km(2) of New South Wales, Victoria and South Australia (White et al. 2003). Hattah Lakes is a semi-arid environment composed of a 13 000 ha complex of over 20 lakes (10 to 200 ha) and associated waterways and floodplains (Murray Darling Basin Authority (MDBA) 2012). Twelve lakes are listed under the Ramsar Convention as internationally important wetlands (Dept of Sustainability and Environment (DSE) 2003; MDBA 2012). The Hattah Lakes area is defined by the extent of the 1956 flood event, the largest known for the region (MDBA 2012).

Hattah Lakes has a long history of human occupation which has shaped the vegetation communities and their composition. For tens of thousands of years, the Murray River floodplain and the vegetation mosaic of Hattah Lakes sustained indigenous people, including the Latji Latji and Nyeri Nyeri people (Donati 2008). Many cultural sites exist within the Park, some of which have been dated to over 15 000 years old (DCNR 1996; Donati 2008).

Since European settlement, the vegetation has been altered by overgrazing, soil erosion, fire, water regulation and recreational activities (DCNR 1996). In 1847, James Crawford established 'Kulkyne', a 20 mile pastoral run, starting the pastoral industry which peaked in the 1860s (DCNR 1996; Donati 2008). At the same time (and later) the area was also heavily affected by the timber trade, producing fuel for paddle steamers and settlement (e.g. fence posts, housing; DCNR 1996, Donati 2008). In 1915, a native game reserve was created around some of the lakes and in 1924 the Kulkyne State Forest was established, incorporating most of the Crawford pastoral run (Donati 2008). Hattah Lakes National Park was established in 1960, with the state forest incorporated in 1980 to form the current Hattah-Kulkyne National Park (DCNR 1996).

The vegetation mosaic on the Hattah Lakes floodplain relies on occasional flooding (natural or environmental watering) to maintain ecosystem function and allow many plant species to complete their life cycles (e.g. germination, flowering). Diminished connectivity between the Hattah Lakes and Murray River, together with the extraction of water for agriculture, industry and urban use, and severe drought over the last decade, have negatively affected vegetation communities of the Hattah Lakes ecosystem that depend on flooding (MDBA 2012). As a result, the environmental health of the floodplain ecosystem and its habitat value for fauna and flora has declined (Cunningham et al. 2009). In 2005, a program of environmental watering was implemented to manage the reduced frequency of natural flooding and to inundate Hattah Lakes (MDBA 2009). The delivery of environmental water is seen as an important factor in the maintenance and improvement of ecosystem health and biodiversity values (DSE 2003; MDBA 2012).

In this study, we investigated the response of 23 rare plant species to an environmental watering event. Rare plant species were determined by their listing on the Victorian Department of Environment, Land Water and Planning (DELWP) Advisory List of Rare or Threatened Plant Species (2014). We compared their abundance before and after the watering event, and evaluated the responses of species associated with different water plant functional groups.


Study region

This study was undertaken in the Hattah Lakes system of Hattah-Kulkyne National Park (480 km(2)) on the Murray River floodplain in northwestern Victoria. Ten Ecological Vegetation Classes (EVCs; White et al. 2003; MDBA 2012) occur across the park, three of which occur as a mosaic in areas that are intermittently flooded: Lake Bed Herbland, Intermittent Swampy Woodland and Riverine Chenopod Woodland.

Lake Bed Herbland is dominated by species adapted to drying mud within lake beds on floodplains (Fig. 1). This vegetation type has two distinct temporal stages: with photosynthetic vegetation and without photosynthetic vegetation (i.e. the species present occur only in the soil seed store), from open water to bare mud. Floods are intermittent but water may be retained for several seasons leading to active growth at the 'drying mud stage' (White et al. 2003). Intermittent Swampy Woodland has a canopy dominated by River Red Gum Eucalyptus camaldulensis (Fig. 2) and Eumong Acacia stenophylla and sometimes Black Box E. largiflorens and Tangled Lignum Duma florulenta with a variable shrubby and rhizomatous sedgy-turf grass ground stratum (White et al. 2003). The vegetation is dominated by flood-stimulated species, together with species tolerant of inundation. Flooding is unreliable but may be extensive (White et al. 2003). Riverine Chenopod Woodland has a canopy dominated by E. largiflorens above a characteristic chenopod shrub stratum. Commonly associated species include Nitre Goosefoot Chenopodium nitrariaceum, Nodding Saltbush Einadia nutans, Ruby Saltbush Enchylaena tomentosa var. tomentosa and Hedge Saltbush Rhagodia spinescens; Acacia stenophylla may also occur (White et al. 2003).

Study design

In April 2014, 20 sites were established and baseline floristic data were collected across 10 lakes (Fig. 3), prior to an environmental watering event. The aim of watering was to inundate the floodplain vegetation to 45 m elevation (asl) (replicating a one-in-eight-year flood event; MBDA 2012) with a particular focus on targeting the treed plant communities and their associated network of permanent and semipermanent wetlands. Eighty-eight gigalitres (GL) of water were delivered to the main lakes between 19 May and 11 September 2014, and 16 GL were delivered to Lake Kramen between 7 September 2014 and 18 January 2015. The sites were resurveyed in April 2015, following the watering event.

At each site, a transect running perpendicular to the lake edge of April 2014 onto the floodplain, was used to measure biological and environmental attributes across the elevation-moisture gradient. Nine of the surveyed lakes contained water in April 2014, while the tenth (Lake Kramen) did not. The lake edge in that instance was estimated based on vegetation composition and species that are known to colonise dry lake beds at Hattah-Kulkyne. Floristic information was recorded in 1 m(2) quadrats along either 50 m or 100 m transects (14 and 6 sites, respectively). Twenty-five 1 m(2) quadrats were surveyed along the 50 m transects and thirty 1 m(2) quadrats were surveyed along the 100 m transects. In addition, each 50 m transect had one 15 x 15 m quadrat and each 100 m transect had two 15 x 15 m quadrats. The percentage of live foliage projective cover of each vascular plant species in each quadrat (irrespective of size) was estimated to the nearest 5%.

Rare species were defined as those listed on DELWP's Advisory List of Rare and Threatened Species (2014). The Victorian Flora and Fauna Guarantee Act 1988 (FFG Act) and Commonwealth Environment Protection and Biodiversity Conservation Act 1999 were also searched for additional rare species found during our surveys. The status of these species is recorded as 'poorly known', 'rare', 'vulnerable' or 'endangered' depending upon their risk of disappearing from wild populations. This risk is based on the number of populations, the area over which populations occur and/or their ability to return from disturbance, including natural events (e.g. drought, fire, flood, landslip; DELWP 2014). Plant taxonomy follows Walsh and Stajsic (2007).

Water Plant Functional Groups

We used Water Plant Functional Groups (e.g. Merritt et al. 2010; Casanova 2011; Campbell et al. 2014) to compare species' responses to watering. These functional groups are based on how species respond to flood events and their water requirements over their lifetime (Brock and Casanova 1997). Five functional groups (Brock and Casanova 1997; Casanova 2011; Campbell et al. 2014) were considered here.

Terrestrial dry (Tdr) species are essentially terrestrial plant species which do not require flooding, but will germinate in damp soil following a flood event. They may invade riparian zones and wetland edges if these remain dry for an extended period (e.g. episodic lakes; Casanova 2011). We expect the response of terrestrial dry species to environmental watering to be variable. That is, species may disappear from areas inundated too long (e.g. the soil seed bank cannot withstand prolonged inundation) but may increase in areas where inundation is short (e.g. leaving the soil seed bank intact and the soil suitably damp for germination).

Terrestrial damp (Tda) species may germinate following flooding or high rainfall and need the soil to remain damp for approximately three months. They cannot tolerate flooding but exist in dried up puddles, drainage lines and wetland edges (Casanova 2011). We expect terrestrial damp species to be present in any year if there has been sufficient rainfall to provide damp soil in which they can germinate, and limited flooding while they are in their vegetative state. Amphibious fluctuation responder - plastic (ARp) species respond to changes in water levels morphologically; for example, rapid growth, and are able to survive on damp and drying soil (Casanova 2011).We expect amphibious responder species present in the soil seed bank to germinate in response to an environmental watering event.

Amphibious fluctuation tolerators - low-growing (ATl) species can germinate on either saturated soil or under water but must be above water in order to flower and set seed (Casanova 2011). We expect low-growing amphibious tolerator species to be present aboveground both before and after an environmental watering event, although they may be more common after inundation as they require shallow flooding for approximately three months.

Amphibious fluctuation tolerators - emergent (ATe) species can survive in saturated soil or shallow water but most of the plant must be above water (Casanova 2011). We expect emergent amphibious tolerator species to be present before and after the environmental watering event, although they may be more common after inundation as they need water to be present for approximately 8 to 10 months of the year.

Data analysis

We searched the Victorian Biodiversity Atlas (VBA; DELWP 2015) for previous records of rare species within Hattah-Kulkyne National Park, to determine how many additional records our study added. The quadrat data (1 m(2) and 15 x 15 m) were used to determine the total number of records for each of the rare species recorded. Each quadrat occurrence was counted as one record.

Each rare species was allocated to a Water Plant Functional Group (Brock and Casanova 1997; Casanova 2011; Campbell et al. 2014). For species not allocated to a functional group by Campbell et al. (2014), we followed the key in Casanova (2011) to allocate species to an appropriate group.

Wilcoxon signed rank tests in R 3.2.3 (R Core Team 2015) were used to investigate if there was a difference in species or functional group cover before and after the environmental watering event. Wilcoxon tests were run for emergent amphibious tolerator, terrestrial damp and terrestrial dry functional groups comparing paired quadrats for each species before and after the watering event. Unlike the number of records, here we considered only the 1 m(2) quadrats. To be included in the analysis, species had to be present in each quadrat in only one year. Amphibious responders and low-growing amphibious tolerators were not tested because only one species (Ammannia multiflora, Centipeda nidiformis respectively) was present in each of these functional groups. Ammannia multiflora was present only in 2015 and Centipeda nidiformis in less than 10 quadrats across both years, thus a meaningful comparison could not be made between the years. In order for a species to be analysed it had to have at least 10 records within the 1 m(2) quadrats across both years. Of the 23 rare species, Wilcoxon tests were carried out for five species (Alternanthera sp. 1 (Plains), Calotis cuneifolia, Cyndon dactylon var. pulchellus, Eragrostis lacunaria, Phyllanthus lacunellus). For the remaining 18 species, there were either <10 records from both years or the species was present in one year only, thus meaningful statistical comparisons could not be made.


The surveys resulted in 319 records of 23 rare species, in the Hattah Lakes system (Table 1). Four species (Alternanthera sp. 1 (Plains), Cardamine moirensis, Crinum flaccidum, Cyperus squarrosus) were recorded for the first time within Hattah-Kulkyne National Park. Eleven species were recorded in both survey years, while 12 species were recorded only in 2014 or 2015 (Table 1). Of the 23 species, five are poorly known, eight are rare, nine are vulnerable and one is listed as endangered (DELWP 2014). Two species, Crinum flaccidum (vulnerable) and Cyperus rigidellus (endangered), are listed under the FFG Act.

The 23 rare species occurred across five functional groups (Table 1). Two of these groups were represented by one species each, amphibious responders (Ammannia multiflora) and low-growing amphibious tolerators (Centipeda nidiformis). Emergent amphibious tolerators were represented by four species (Cyperus rigidellus, Cyperus squarrosus, Isolepis australiensis, Lipocarpha microcephala). The dominant functional groups were terrestrial dry (11 species) and terrestrial damp (six species).

Terrestrial dry species exhibited a significant difference before and after the watering event with cover generally higher in 2014 (Table 2). Individually, the 11 terrestrial dry species showed variable responses to the environmental watering event with two species (Lotus australis var. australis, Sida fibulifera) present in 2014 only. In the case of L. australis var. australis the single site at which it was present in 2014 was still inundated in 2015. Another three species (Atriplex lindleyi subsp. lindleyi, Triraphis mollis, Wahlenbergia tumidifructa) were present only in 2015. The remaining six species (Calotis cuneifolia, Cynodon dactylon var. pulchellus, Eragrostis lacunaria, Phyllanthus lacunellus, Sclerolaena patenticuspis, Swainsona microphylla) were present in both years. Of these six species, the cover of Calotis cuneifolia was significantly higher before the watering event (2014) while the cover of Phyllanthus lacunellus was higher following the watering event (2015; Table 2).

The covers of terrestrial damp species were significantly higher in 2014 prior to the watering event (Table 2). Individually the six terrestrial damp species showed different responses to the environmental watering event, with Bergia trimera and Rorippa eustylis present only in 2014 and Cardamine moirensis present only in 2015. The remaining three species (Alternanthera sp. 1 (Plains), Austrobryonia micrantha (Fig. 4), Crinum flaccidum) were present in both years with the presence of Crinum flaccidum similar in both years, while the other species fluctuated. Alternanthera sp. 1 (Plains) exhibited a significantly higher cover before the watering event (2014; Table 2).

Ammannia multiflora (amphibious responder) was recorded at one site only following the watering event. Centipeda nidiformis (lowgrowing amphibious tolerator) was recorded in more quadrats following the environmental watering event than before it (6 vs. 2 quadrats) and at an additional two sites following inundation. Two of the four emergent amphibious tolerator species (Cyperus rigidellus and Cyperus squarrosus) were present in 2014 only, and one (Isolepis australiensis) in 2015 only. The fourth species (Lipocarpha microcephala) was present in both years.


The environmental watering event in late 2014 simulated a one in eight year flood event. Overall expectations were that vegetation community condition would improve (e.g. increased canopy health), flood respondent species would germinate and the abundance of species would increase as a result of increased soil moisture post-inundation. Water plant functional groups can be used as a means to predict how plant functional groups as a whole and individual species will respond to flooding (natural or otherwise). Generally, we would expect species in the five functional groups (terrestrial dry, terrestrial damp, amphibious responders, low-growing amphibious tolerators, emergent amphibious tolerators) to be present before and after the watering event. Their presence above-ground before and after would be dictated by available soil moisture and, for the amphibious tolerators, presence of shallow water. In addition, the length of inundation may influence how some species survive the watering event and whether those species in the soil seed bank are triggered to germinate.

Vegetation surveys prior to the environmental watering event detected 16 rare species in four functional groups (terrestrial dry, terrestrial damp, both amphibious tolerators). Following the watering event, an additional seven rare species were recorded, including one species in an additional functional group (amphibious responders). Of the 16 species recorded in 2014, 11 were still present in 2015 while the remaining five species were no longer present in the above-ground vegetation. There was considerable variability in how species within each functional group responded to the watering event.

Both terrestrial dry and terrestrial damp species showed variable responses to the environmental watering event. While species of both groups respond to increased soil moisture following flooding or high rainfall, they cannot tolerate flooding for extended periods of time (Casanova 2011). Three terrestrial dry species (Atriplex lindleyi subsp. lindleyi, Triraphis mollis, Wahlenbergia. tumidifructa) were present in 2015 only, in quadrats which were inundated during the environmental watering event, suggesting they may have germinated in response to increased soil moisture resulting from inundation. Bergia trimera and R. eustylis (terrestrial damp) were present in 2014 and not 2015, suggesting the above average rainfall (Bureau of Meteorology 2015) in the 12 months preceding the April 2014 surveys was sufficient to trigger germination of these species. In addition, terrestrial species lack tolerance for extended periods of inundation (Casanova 2011) and this may be the reason for their absence in 2015, as the quadrats in which B. trimera and R. eustylis were present were inundated for approximately 2-3 months during the environmental watering event. Furthermore, the presence of terrestrial species following the watering event may be a result of the interaction between inundation and site specific factors (e.g. soil type and moisture, competition from more robust species, presence in the soil seed bank, other disturbances) plus the inherent rareness of suitable habitat for these species (Casanova and Brock 2000; Capon 2003; Raulings et al. 2010).

The amphibious functional groups have slightly different responses to inundation, with amphibious responders responding morphologically (Casanova 2011), and the two amphibious tolerator groups germinating in response to inundation. Thus, we expected amphibious responder species to germinate from the soil seed bank following inundation and the two tolerator groups to be more common following inundation as they require water to be present for extended periods of time (Casanova 2011). Both the single amphibious responder (Ammannia multiflora) and low-growing amphibious tolerator (Centipeda nidiformis) were more common following the environmental watering event than before it, suggesting inundation triggered germination. Nevertheless, this interpretation should be treated with caution as A. multiflora was present at only one site.

Emergent amphibious tolerators were more variable in their response to the environmental watering event. Two species (Cyperus rigidellus, Cyperus squarrosus) were present in 2014 but not 2015. As we expected emergent amphibious tolerators to be present in both wet and dry years, it was an unexpected result to 'lose' these two species from the landscape following the watering event as we would have expected them to increase in abundance. While these species require water for approximately 8 to 10 months of the year (Casanova 2011), it is possible that the period of submergence was too long for C. rigidellus and C. squarrosus to survive. There are also other factors that may have contributed to this result, such as seed presence or absence in the soil seed bank, other unknown germination triggers, competition from more robust species and the inherent rareness of suitable habitat. Lipocarpha microcephala was present at two of three sites in both years with a strong increase in cover at one of those sites (2% to 12% cover) indicating a response to the environmental watering event.

Water plant functional groups provide a useful tool to examine species' responses and make predictions as to how species within the groups will respond to an environmental watering event. While our expectations were broadly met, there was also much variation between individual species in each group. This is not an unusual outcome and highlights the need to appreciate that local influences may differ widely, affecting how species react to an environmental watering event. In addition, limitations associated with studying rare species often result in a paucity of data, as was the case with some species in this study. While functional group and species responses to the environmental watering event differed, a key outcome from this study was the increase in rare species records for Hattah-Kulkyne National Park.


This work was commissioned and funded by the Mallee Catchment Management Authority through the Federal Government's The Living Murray Initiative. The authors would like to thank Andrew Greenfield (Mallee Catchment Management Authority) and Parks Victoria (Hattah-Kulkyne National Park) staff, in particular Shane Southon and Damian Kerr. Doug Frood assisted with plant identification in 2014. David Cameron and Neville Walsh assisted with plant identification in 2015. Brad Farmilo and Garry Cheers assisted with field work in 2015. Mike Duncan, Brad Farmilo, Matt White, Andrew Bennett and David Cheal provided comments on earlier versions of the manuscript that greatly improved readability.


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Received 28 April 2016; accepted 6 October 2016

Sally A Kenny, Claire Moxham and Geoff Sutter

Arthur Rylah Institute for Environmental Research, Department of Environment, Land, Water and Planning, 123 Brown Street, Heidelberg, Victoria, 3084 Email:

Table 1. Rare plant species recorded during botanical surveys at
Hattah-Kulkyne National Park including the number of records from the
Victorian Biodiversity Atlas (VBA) and from the current study. Species
in bold are new records for Hattah-Kulkyne National Park.

recorded  Scientific name                    Common name

Both      Alternanthera sp. 1 (Plains)       Plains Joyweed
2015      Ammannia multiflora                Jerry-jerry
2015      Atriplex lindleyi subsp. lindleyi  Flat-top Saltbush
Both      Austrobryonia micrantha            Mallee Cucumber
2014      Bergia trimera                     Small Water-fire
Both      Calotis cuneifolia                 Blue Burr-daisy
2015      Cardamine moirensis                Riverina Bitter-cress
Both      Centipeda nidiformis               Cotton Sneezeweed
Both      Crinum flaccidum                   Darling Lily
Both      Cynodon dactylon var. pulchellus   Native Couch
2014      Cyperus rigidellus                 Curly Flat-sedge
2014      Cyperus squarrosus                 Bearded Flat-sedge
Both      Eragrostis lacunaria               Purple Love-grass
2015      Isolepis australiensis             Inland Club-sedge
Both      Lipocarpha microcephala            Button Rush
2014      Lotus australis var. australis     Austral Trefoil
Both      Phyllanthus lacunellus             Sandhill Spurge
2014      Rorippa eustylis                   Dwarf Bitter-cress
Both      Sclerolaena patenticuspis          Spear-fruit Copperburr
2014      Sida fibulifera                    Pin Sida
Both      Swainsona microphylla              Small Leaf Swainsona-pea
2015      Triraphis mollis                   Needle Grass
2015      Wahlenbergia tumidifructa          Mallee Annual-bluebell

Year      Water plant  VIC     Number of records within Hattah
recorded  functional   ADV(2)
group(1)                       VBA  This   Updated
                                    study  total(3)

Both      Tda          k         0  20      20
2015      ARp          V         2   2       4
2015      Tdr          k         5   2       7
Both      Tda          r         2  13      15
2014      Tda          V         2   4       6
Both      Tdr          r         8  72      80
2015      Tda          r         0   1       1
Both      ATl          r         1   8       9
Both      Tda          v(*)      0  10      10
Both      Tdr          k         2  16      18
2014      ATe          e(*)      1   3       4
2014      ATe          V         0   6       6
Both      Tdr          V        34  30      64
2015      ATe          k         1  15      16
Both      ATe          V        19   9      28
2014      Tdr          k         1   1       2
Both      Tdr          r         7  71      78
2014      Tda          r         8   1       9
Both      Tdr          V         3   9      12
2014      Tdr          V        12   5      17
Both      Tdr          r       150  15     165
2015      Tdr          r         5   1       6
2015      Tdr          r         1   5       6

1: water plant functional groups are: ARp--Amphibious fluctuation
responder--plastic, ATl--Amphibious fluctuation tolerator--low-growing,
ATe--Amphibious fluctuation tolerator--emergent, Tda--Terrestrial damp
and Tdr--Terrestrial dry. 2: species status codes are: r--rare,
v--vulnerable, e--endangered and k--poorly known (DELWP 2014). 3:
updated total number of records for Hattah (i.e. VBA + this study).(*)
indicates species also listed under the Flora and Fauna Guarantee Act

Table 2. The relationship between the environmental watering event and
water plant functional group and species cover. Significant results
are shown in bold and comparisons not undertaken due to limited
records are not shown.

Water plant functional group  Plant species
Terrestrial dry
                              Calotis cuneifolia
                              Cynodon dactylon var. pulchellus
                              Eragrostis lacunaria
                              Phyllanthus lacunellus
Terrestrial damp
                              Alternanthera sp. 1 (Plains)
Amphibious fluctuation
tolerator - emergent
Water plant functional group  V      p

Terrestrial dry               8652    0.01
                               536   <0.0001
                                42.5  0.33
                               179.5  0.93
                               630    0.05
Terrestrial damp               462   <0.0001
                               171    0.0002
Amphibious fluctuation
tolerator - emergent            83.5  0.43
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Author:Kenny, Sally A; Moxham, Claire; Sutter, Geoff
Publication:The Victorian Naturalist
Article Type:Report
Geographic Code:8AUVI
Date:Feb 1, 2017
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