Fish responses to low flows in lowland streams: a summary of findings from the Granite Creeks system, Victoria Nick Bond Australian Rivers Institute, Griffith University Low flows report series – June 2012 NATIONAL WATER COMMISSION — Low flows report series ii © Commonwealth of Australia 2012 This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be reproduced by any process without prior written permission. Requests and enquiries concerning reproduction and rights should be addressed to the Communications Director, National Water Commission, 95 Northbourne Avenue, Canberra ACT 2600 or email bookshop@nwc.gov.au. Online/print: ISBN: 978-1-921853-75-3 Published by the National Water Commission 95 Northbourne Avenue Canberra ACT 2600 Tel: 02 6102 6000 Email: enquiries@nwc.gov.au Date of publication: June 2012 An appropriate citation for this report is: Bond N 2012, Fish responses to low flows in lowland streams: a summary of findings from the Granite Creeks system, Victoria, National Water Commission, Canberra. Disclaimer This paper is presented by the National Water Commission for the purpose of informing discussion and does not necessarily reflect the views or opinions of the Commission. NATIONAL WATER COMMISSION — Low flows report series iii Low flows report series This paper is part of a series of works commissioned by the National Water Commission on key water issues. This work has been undertaken by Griffith University in consultation with the Victorian Department of Sustainability and Environment, and the Victorian Environmental Protection Agency, on behalf of the National Water Commission. NATIONAL WATER COMMISSION — Low flows report series iv Contents Executive summary Report context 1. Introduction 2. The Granite Creeks system 2.1. Hydrology 2.2. Fish population dynamics 2.3. Habitat dynamics 2.4. Dispersal and population connectivity 2.5. Putting it all together – modelling population dynamics 2.6. Summary References vii ix 1 2 2 4 6 9 11 12 13 Tables Table 1: Summary of system-wide pool persistence in the Granite Creeks for the period 2007–09. .............................................................................................................................. 7 Figures Figure S1: Context of reports produced for the Low Flow Ecological Response and Recovery Project. Each circle represents the location of individual case studies and the size of each circle represents the spatial extent of each case study. ............................................................................................................................................. ix Figure 1: Plot showing the location and layout of the Granite Creeks stream network. .......................................................................................................................................... 2 Figure 2: Plotted flow-duration curves for two of the Granite Creeks showing changes in runoff patterns under natural, current and a potential (2030 median) climate change scenario. .................................................................................................. 3 Figure 3: Proportion of streamflow gauges displaying perennial (+’ve) and intermittent (-‘ve) streamflow in Victoria. Note pre-1930s data is unreliable due to just one or two gauges being operational. Trend line displays moving average of the two proportions and ranges between 0.5 (all perennial) and 0.5 (all intermittent). ........................................................................................................................ 4 Figure 4: Total abundances of native fish in relation to local a) dissolved organic carbon, and b) dissolved oxygen concentrations in contracting summer refuge pools. From McMaster and Bond (2008). ........................................................................... 5 Figure 5: Trends in the abundance of western carp gudgeons (H. klunzingeri) at four fixed sites on Faithful Creek over the period 2004–09............................................................ 5 Figure 6: Trends in total abundance of western carp gudgeons (H. klunzingeri) from 2007–09 in Faithful Creek. ..................................................................................................... 6 Figure 7: Trends in total abundance of western carp gudgeons (H. klunzingeri) from 2007–09 in Faithful, Castle and Honeysuckle creeks. ........................................................... 7 Figure 8: Maps showing the distribution of persistent waterholes in 2007, 2008 and 2009. Note the greater number of waterholes in 2008, which is reflected in Table 1. ....................................................................................................................................... 8 Figure 9: Plot showing the number of fish caught at sites during spring as a function of distance from permanent water at the end of the previous dry period. ........................................................................................................................................... 10 Figure 10: Examples of individual runs from the demographic model illustrating the high inter-annual variability in population size in response to changing habitat availability. Note in all cases that declining population sizes in response to dry spells (grey boxes) are always lagged. From Perry and Bond (2009). ................................................................................................................................. 11 Figure 11: Plot showing relative population change in relation to the availability of wetted habitat along the channel from one year to the next. From Perry and Bond (2009). ................................................................................................................................. 12 NATIONAL WATER COMMISSION — Low flows report series v Acknowledgements Numerous staff and students helped collect field data as part of the Granite Creeks project. In no particular order thanks are extended to Tom Daniel, Matthew Johnson, Damien McMaster, Zoe Squires, Tim Dexter and Darren Giling. Paul Reich, Rob Hale and Sam Lake also shared resources from the Murray-Darling Basin Authority (MDBA)-funded Riparian Restoration Experiment being carried out on some of the same sites, and Paul Reich and Sam Lake contributed to ongoing discussions throughout the course of the work. eWater CRC also provided funding for much of the work and for Nick Bond during the course of the data collection. NATIONAL WATER COMMISSION — Low flows report series vi Executive summary The latest incarnation of the Granite Creeks project was established in 2006. It aimed to understand the impacts of low-flow stress on fish population persistence at a landscape (whole-of-river-system) scale in streams draining the Granite Creeks region in central Victoria. The project included four primary areas of activity: 1. Empirical observation of fish population trajectories at individual sites 2. Empirical observation of habitat dynamics (hydrology and pool persistence) at a landscape scale 3. Measurement of fish dispersal via the application of genetic approaches 4. Spatially explicit modelling of fish population persistence. The purpose of the current document is not to present a detailed analysis of the data collected as part of the Granite Creeks project, but instead to summarise some of the key findings and to draw on these in presenting some of the lessons that have emerged through the course of the research. A summary of the research program has yielded a number of key lessons presented below. Lesson 1 Perhaps with the exception of perennial streams and larger river systems with a relatively high level of hydrologic connectivity and low transmission losses, available gauging data are frequently unsuitable to support questions relating ecological dynamics to spatial and temporal patterns of hydrologic variability. Lesson 2 Farm dams can have a dramatic effect on flow permanency in small unregulated rivers, with significant consequences for aquatic ecosystems, including the permanent loss of low-flowsensitive fish and invertebrate taxa. Lesson 3 A number of native fish species found inhabiting drying pools in the Granite Creek system show high levels of persistence at local scales even when exposed to very harsh environmental conditions. Lesson 4 While datasets such as the Granite Creeks dataset have provided some useful insights into the effects of low flows, in highly variable climates such as Australia, long-term datasets are essential to understand how populations are influenced by low-flow periods. Such datasets are still uncommon. Lesson 5 During low flows, dispersal distances of small-bodied native fish may be extremely limited, leading to slow and constrained recolonisation of sites from which local populations have been lost. This has implications for recovery rates following prolonged periods of low flow. NATIONAL WATER COMMISSION — Low flows report series vii Lesson 6 Consideration of the influence of climate variability on habitat persistence, together with the influence of demographic characteristics in determining the lags in population response to wet-dry cycles, provide a useful way of illustrating the potential thresholds in population growth trends. Further work is needed to validate these predictions. NATIONAL WATER COMMISSION — Low flows report series viii Report context This report is part of a larger series of reports produced for the National Water Commission’s Low Flow Ecological Response and Recovery Project (Figure S1). This report presents one of 11 hydro-ecological case studies. The purpose of the case studies is to test hypotheses that relate ecological process and function and biological traits to key hydrological measures that are affected by low flows. A summary of the findings in this report and the other case studies are contained in Synthesis of case studies quantifying ecological responses to low flows (Marsh et al. 2012). Guidance on ecological response and hydrological modelling for low-flow water planning Low-flow hydrological classification of Australia Review of literature quantifying ecological responses to low flows Early warning, compliance and diagnostic monitoring of ecological responses to low flows Synthesis of case studies quantifying ecological responses to low flows Figure S1: Context of reports produced for the Low Flow Ecological Response and Recovery Project. Each circle represents the location of individual case studies and the size of each circle represents the spatial extent of each case study. NATIONAL WATER COMMISSION — Low flows report series ix 1. Introduction High inter-annual variability in runoff is a key feature of Australia’s aquatic ecosystems (McMahon et al. 2007). This manifests itself as cycles of floods and droughts, giving rise to a boom and bust ecology (Lake et al. 2006; Bunn et al. 2006; Jenkins & Boulton 2003). These dynamic cycles have captured intense interest in large inland rivers but more recently they have been considered in a wider context, largely driven by the recent protracted drought in southeastern Australia. This has focussed considerable research effort on understanding the impacts of low flows episodes in a much wider range of river types (Bond et al. 2008) (Robson and Matthews 2004, Lind et al. 2006) (Chessman 2009). The nature of dry cycles (and droughts) – their slow onset and large spatial scale – mean that their effects are best studied over long time and large spatial scales (Lake et al. 2008). Yet most traditional research is focused at the site-scale, both in terms of the scale at which samples are collected and the scale at which relationships are assessed. The few studies conducted at landscape scales demonstrate quite clearly the value of such a perspective. For example, in a review of the consequences of flow variability in dryland rivers, Bunn et al (2006) demonstrated the impacts of lengthy dry periods in isolating refuge waterholes over many thousands of square kilometres. Thus, while much has been learnt from studying assemblage composition and ecological processes within individual isolated waterholes (Arthington et al. 2005; Balcombe et al. 2005; Marshall et al. 2006; Balcombe et al. 2007), it is landscape-scale processes of waterhole persistence that ultimately set the context for longerterm patterns and dynamics. Similar findings have come from work conducted over smaller scales examining the influence of low-flow stress on fish populations inhabiting intermittent streams in the prairie streams of the United States (e.g. Labbe & Fausch 2000). Indeed, the influence of landscape-scale habitat dynamics (patterns) on local-scale ecological processes is at the very heart of landscape ecology (Turner 1989), which during the past 20 years has transitioned from a sub-discipline to a core area of ecological research (Turner 2005). It is in this context that the most recent incarnation of the Granite Creeks project was established in 2006, with the aim of understanding the impacts of low-flow stress on fish population persistence at a landscape (whole-of-river-system) scale. The project included four primary areas of activity: 1. Empirical observation of fish population trajectories at individual sites 2. Empirical observation of habitat dynamics (hydrology and pool persistence) at a landscape scale 3. Measurement of fish dispersal via the application of genetic approaches 4. Spatially explicit modelling of fish population persistence. The purpose of the current document is not to present a detailed analysis of the data collected as part of the Granite Creeks project, but instead, to summarise some of the key findings and to draw on these in presenting some of the lessons that have emerged through the course of the research. NATIONAL WATER COMMISSION — Low flows report series 1 2. The Granite Creeks system 2.1. Hydrology The Granite Creeks system consists of a series of neighbouring tributaries that arise in the Strathbogie Ranges in central Victoria and flow northward into the Goulburn River (Figure 1). In their upper reaches all of the streams, including Pranjip, Creightons, Castle, Sevens, Faithful and Honeysuckle creeks, are perennial, largely due to the persistence of spring-fed baseflows during summer. Only half of the creeks are gauged, and available time series are of short duration and poorly correlated with the regime experienced along the length of each creek. Figure 1: Plot showing the location and layout of the Granite Creeks stream network. Lesson 1: Perhaps with the exception of perennial streams and larger river systems with a relatively high level of hydrologic connectivity and low transmission losses, available gauging data are frequently unsuitable to support questions relating ecological dynamics to spatial and temporal patterns of hydrologic variability. In light of this shortcoming, Monash University contracted Sinclair Knight Merz (SKM) to develop rainfall-runoff models for each of the Granite Creeks mentioned above, the aim being to provide modelled flow data that could be used to help support the interpretation of hydrologic influences on ecological processes in this system (see Bell & Ramchurn 2011). The modelling included the development of runoff sequences for ‘non-consumptive’, ‘current’ and ‘future’ scenarios, and these are particularly instructive in showing the potential impacts NATIONAL WATER COMMISSION — Low flows report series 2 of anthropogenic water use on low-flow characteristics in unregulated systems. For example, Castle Creek has already undergone an extensive change from a naturally perennial to now quite intermittent river system, with an average runoff reduction of 40 per cent as a result of farm dams (Bell & Ramchurn 2011). In contrast, Creightons Creek has undergone far less dramatic changes in runoff (20 per cent) due to lower dam densities in the upper catchment. These findings are consistent with work in other regions of south-eastern Australia, yet the effects of farm dams on aquatic ecosystems are still largely ignored in a policy context. Lesson 2: Farm dams can have a dramatic effect on flow permanency in small unregulated rivers, with significant consequences for aquatic ecosystems including the permanent loss of low-flow sensitive fish and invertebrate taxa. 10000 Castle Creek Flow (ML/d) 1000 100 10 1 0.1 0% 20% Natural 10000 40% 60% 80% 100% Percentage Exceedance Current 2030 Med CC 10th Percentile Creightons Creek Flow (ML/d) 1000 100 10 1 0.1 0% 20% Natural 40% 60% 80% 100% Percentage Exceedance Current 2030 Med CC 10th percentile Figure 2: Plotted flow-duration curves for two of the Granite Creeks showing changes in runoff patterns under natural, current and a potential (2030 median) climate change scenario. NATIONAL WATER COMMISSION — Low flows report series 3 Even where farm-dam impacts have had a minimal impact on streamflows, periodic droughts can have a large impact on patterns of cease-to-flow in river systems such as the Granite Creeks, as evidenced by Figure 3, which shows long-term trends in the proportion of Victorian streamflow gauges exhibiting permanent flow each year. Note in particular the droughts of 1937–45, 1965–68, 1982–83 and post 2000, all of which caused a sharp spike in the occurrence of intermittent flows. Figure 3: Proportion of streamflow gauges displaying perennial (+’ve) and intermittent (-‘ve) streamflow in Victoria. Note pre-1930s data is unreliable due to just one or two gauges being operational. Trend line displays moving average of the two proportions and ranges between 0.5 (all perennial) and -0.5 (all intermittent). 2.2. Fish population dynamics With flow intermittency comes a rapid fragmentation and contraction of habitat (Stanley et al. 1997) and rapid declines in water quality, including increasing temperatures, decreasing oxygen concentrations, and in many lowland rivers in Australia, rapid increases in respiration rates and concentrations of dissolved organic carbon (DOC) from falling eucalyptus leaves. The Granite Creeks project has examined both local-scale impacts of declining water quality and whole-of-stream-scale changes in connectivity and habitat availability. At a local scale, McMaster and Bond (2008) showed that a number of native fish species occurring in lowland sections of the Granite Creeks system (including southern pygmy perch (Nannoperca australis), mountain galaxias (galaxias olidus) and western carp gudgeon (Hypseleotris klunzingeri)) were widely distributed in relation to low oxygen concentrations and high DOC concentrations (Figure 4), appearing in lower numbers even where dissolved oxygen (DO) concentrations fell below 2 mgl-1 (McMaster & Bond 2008). They subsequently observed similar tolerances of adult fish in a series of laboratory experiments, but speculated that metabolic costs were still being borne by individuals living in these environments, or that other life stages such as eggs and larvae were more susceptible. More recently Morrongiello NATIONAL WATER COMMISSION — Low flows report series 4 et al. (2011) confirmed sub-lethal effects of DOC on southern pygmy perch (N. australis), with a reduced number of females reaching sexual maturity in experimental treatments consisting of high DOC concentrations. Figure 4: Total abundances of native fish in relation to local a) dissolved organic carbon, and b) dissolved oxygen concentrations in contracting summer refuge pools. From McMaster and Bond (2008). These findings regarding fish survival in drying stream pools have been further confirmed by multi-year surveys of fixed sites in the Granite Creeks system, which include a mix of permanent and semi-permanent waterholes. One notable finding is the potential consequence of sampling a small number of fixed sites, or simply choosing sites randomly with surface water present on each sampling date. For example, sampling at two fixed sites on Faithful Creek, which first dried completely in 2006, suggests a complete loss of western carp gudgeons (H. klunzingeri) from this system (Figure 5). Figure 5: Trends in the abundance of western carp gudgeons (H. klunzingeri) at four fixed sites on Faithful Creek over the period 2004–09. However, a wider survey of sites across Faithful Creek encompassing several permanent waterholes suggests a somewhat different pattern – although abundances at these sites were also low (Figure 6). NATIONAL WATER COMMISSION — Low flows report series 5 Figure 6: Trends in total abundance of western carp gudgeons (H. klunzingeri) from 2007–09 in Faithful Creek. Across a much larger set of sites (n=25) in Faithful, Honeysuckle, Castle and Sevens creeks over the same period, H. klunzingeri abundances showed a quite different pattern (Figure 7), with a notable increase in abundance in autumn 2009. These broader trends in fish abundance are still under investigation, with sampling continuing up until the present (although data are not currently available for inclusion here). Lesson 3: A number of native fish species found inhabiting drying pools in the Granite Creek system show high levels of persistence at local scales even when exposed to very harsh environmental conditions. 2.3. Habitat dynamics An important question being asked of this data is to what degree longer-term and landscapescale changes in abundance reflect changes in total habitat availability. For example, there were approximately twice as many pools that persisted through the dry season of 2007–08 (Table 1). A further indication of the differences in habitat persistence is provided in Figure 8. This increase in habitat may have reduced population densities and increased net reproductive output of individuals that survived to breed successfully in 2008–09. This is one potential explanation for the rise in overall abundances in the autumn 2009 surveys. At this stage this hypothesis remains extremely speculative, although there is evidence from the literature that severe drought years can increase density-dependent mortality and reduce reproductive output (Bell et al. 2000; Elliott 2006). Notably, this evidence comes from a 30year study of population dynamics in a hydrologically stable region in England. The uniqueness of the study by Elliot (2006) illustrates the importance of long-term data. Lesson 4: While datasets such as the Granite Creeks dataset have provided some useful insights into the effects of low flows, in highly variable climates such as Australia, long-term datasets are essential to understand how populations are influenced by low-flow periods. Such datasets are still uncommon. NATIONAL WATER COMMISSION — Low flows report series 6 Figure 7: Trends in total abundance of western carp gudgeons (H. klunzingeri) from 2007–09 in Faithful, Castle and Honeysuckle creeks. Table 1: Summary of system-wide pool persistence in the Granite Creeks for the period 2007–09. Year Mean % Total number Mean distance Max distance Rainfall in flowing length of pools between pools between preceding year per stream (m) pools (m) (mm) per stream 2007 12.1 744 595.0 32877 286 2008 15.1 1796 243.0 9520 562 2009 12.1 624 661.0 37243 516 NATIONAL WATER COMMISSION — Low flows report series 7 2007 2008 2009 Figure 8: Maps showing the distribution of persistent waterholes in 2007, 2008 and 2009. Note the greater number of waterholes in 2008, which is reflected in Table 1. NATIONAL WATER COMMISSION — Low flows report series 8 2.4. Dispersal and population connectivity The preceding section on habitat dynamics illustrates that not only does habitat availability change dramatically between years even during low-flow periods, but so too does the extent of habitat isolation. Isolation of refuge habitats is an important element to predicting rates of recolonisation and population recovery following low-flow disturbances (Lancaster & Belyea 1997); however, documenting dispersal distances in aquatic ecosystems is fraught with challenges (Downes & Reich 2008; Hughes 2007). In the Granite Creeks system, our complete mapping of permanent refuge habitats has provided a unique opportunity to estimate minimum dispersal distances by fauna colonising previously dry sites following the resumption of flow. The seasonal predictability of the dry period allowed us to map pools close to the end of the dry season, and hence gather a reliable estimate of the location of potential source populations. It is important to note that this approach provides estimates of minimum distances only. Nevertheless, the results are illuminating. In short, of the three native species caught in the surveys, only one, G. olidus, a notably good swimmer, was found to have dispersed more than several hundred metres over the six-month period of hydrologic connectivity. The species was found to have moved a maximum distance of 2000 m (Figure 9). This observation of limited dispersal was further supported by genetic data (Hughes & Schmidt, unpublished data), although this information is not presented here. It is further worth noting that more recent data from a flood year (2010; Dexter, unpublished data), suggests that downstream and upstream movements of G. olidus may be much greater under conditions of high flow. Lesson 5: During low flows, dispersal distances of small-bodied native fish may be extremely limited, leading to slow and constrained recolonisation of sites from which local populations have been lost. This has obvious implications for recovery rates following prolonged periods of low flow. NATIONAL WATER COMMISSION — Low flows report series 9 150 100 50 0 total gudgeon catch in Spring Total gudgeon catch in Spring 200 a) H. klunzingeri 0 2000 4000 6000 8000 10000 12000 Minumum distance of site to pool in Autumn 600 minimum distance of site to pool in Autumn 400 300 200 100 0 total galaxid catch ininSpring Total galaxid catch Spring 500 b) G. olidus 0 2000 4000 6000 8000 10000 12000 Minumum distance of site to pool in Autumn minimum distance of site to pool in Autumn 400 300 200 100 0 total southern perch catch in Spring Totalpygmy southern pygmy perch catch in Spring c) N. australis 0 2000 4000 6000 8000 10000 12000 Minumum distance of site to pool in Autumn minimum distance of site to pool in Autumn Figure 9: Plot showing the number of fish caught at sites during spring as a function of distance from permanent water at the end of the previous dry period. NATIONAL WATER COMMISSION — Low flows report series 10 2.5. Putting it all together – modelling population dynamics A fitting endpoint for this summary of findings from the Granite Creeks project is to consider the longer-term trajectories of fish populations via the application of spatial demographic models. A spatially explicit individual-based population was constructed for the Granite Creeks system to examine the likely long-term population persistence of carp gudgeons in response to increased low-flow stress (Perry & Bond 2009). The model considered stochastic inter-annual variation in rainfall, runoff and habitat persistence, and coupled this with estimates of demographic rates – fecundity, survivorship and (limited) movement patterns – to examine stochastic population dynamics over decadal time scales. Several key findings emerged from this work. First, the model predicted extremely large fluctuations in population density in relation to cycles of high- and low-flow periods (wet/dry climate cycles) Figure 10. Figure 10: Examples of individual runs from the demographic model illustrating the high interannual variability in population size in response to changing habitat availability. Note in all cases that declining population sizes in response to dry spells (grey boxes) are always lagged. From Perry and Bond (2009). Second, the model showed that on average, periods of population growth are restricted to years when greater than 60 per cent of the stream length remained wet (Figure 11). Such periods are already relatively rare due to the effects of farm dams in many rivers in the Granite Creeks system, and are likely to become increasingly rare under future climate change predictions (Figure 2). The model thus suggests that the long-term viability of populations of H. klunzingeri in the Granite Creek system are under long-term threat from the combined effects of catchment interception of runoff by farm dams and climate change. This finding has since received further support from attempts to model range shifts of fish in Victoria in response to climate change (Bond et al. 2012). NATIONAL WATER COMMISSION — Low flows report series 11 Figure 11: Plot showing relative population change in relation to the availability of wetted habitat along the channel from one year to the next. From Perry and Bond (2009). Lesson 6: Consideration of the influence of climate variability on habitat persistence, together with the influence of demographic characteristics in determining the lags in population response to wet-dry cycles, provide a useful way of illustrating the potential thresholds in population growth trends. Further work is needed to validate these predictions. 2.6. Summary Overall, the various strings of empirical research discussed in this summary of the Granite Creeks research have revealed a range of interesting insights into the capacity of fish in the Granite Creeks system to endure low-flow disturbances (presented as a series of ‘lessons’ throughout the text). At the same time, measured responses coupled with longer-term population dynamic models suggest these populations may – to coin a description applied to streams in the Great Plains basins of the United States (Dodds et al. 2004) – be living on the edge. This conclusion is also consistent with the results of the hydrologic classification conducted as part of the NWC low flows project (Mackay et al. 2012), which identified a number of streamflow classes likely to be particularly sensitive to additional low-flow stress. NATIONAL WATER COMMISSION — Low flows report series 12 References Arthington AH, Balcombe SR, Wilson, GA, Thoms MC & Marshall J 2005, ‘Spatial and temporal variation in fish-assemblage structure in isolated waterholes during the 2001 dry season of an arid-zone floodplain river, Cooper Creek, Australia’, Marine and Freshwater Research 56:25. Balcombe S, Bunn S, McKenzie-Smith F & Davies P 2005, ‘Variability of fish diets between dry and flood periods in an arid zone floodplain river’, Journal of Fish Biology 67:1552– 1567. Balcombe S, Bunn S & Arthington A 2007, ‘Fish larvae, growth and biomass relationships in an Australian arid zone river: links between floodplains and waterholes’, Freshwater Biology 52: 2385–2398. Bell A & Ramchurn A 2011, Formulation of river health related hydrologic indices, a report to Monash University by Sinclair Knight Merz, Melbourne. 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