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Freshwater Biology (2015) 60, 2620–2638 doi:10.1111/fwb.


Ecological effects of extreme climatic events on riverine

ecosystems: insights from Australia
*Irstea, UR MALY, Villeurbanne, France

Department of Biological Sciences, Macquarie University, North Ryde, NSW, Australia

Institute for Applied Ecology, University of Canberra, Bruce, ACT, Australia
School of Biological Sciences, Monash University, Clayton, Vic., Australia

Australian Rivers Institute, Griffith University, Nathan, Qld, Australia
**School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham, U.K.

1. Climate extremes and their physical impacts – including droughts, fires, floods, heat waves, storm
surges and tropical cyclones – are important structuring forces in riverine ecosystems. Climate
change is expected to increase the future occurrence of extremes, with potentially devastating effects
on rivers and streams. We synthesise knowledge of extremes and their impacts on riverine ecosys-
tems in Australia, a country for which projected changes in event characteristics reflect global trends.
2. Hydrologic extremes play a major structuring role in river ecology across Australia. Droughts alter
water quality and reduce habitat availability, driving organisms to refugia. Extreme floods increase
hydrological connectivity and trigger booms in productivity, but can also alter channel morphology
and cause disturbances such as hypoxic blackwater events.
3. Tropical cyclones and post-cyclonic floods damage riparian vegetation, erode stream banks and
alter water quality. Cyclone-induced delivery of large woody debris provides important instream
habitat, although the wider ecological consequences of tropical cyclones are uncertain.
4. Wildfires destroy catchment vegetation and expose soils, increasing inputs of fine sediment and
nutrients to streams, particularly when followed by heavy rains.
5. Research on the impacts of heat waves and storm surges is scarce, but data on temperature and
salinity tolerances, respectively, may provide some insight into ecological responses.
6. We identify research gaps and hypotheses to guide future research on the ecology of extreme
climate events in Australia and beyond. A range of phenomenological, experimental and modelling
approaches is needed to develop a mechanistic understanding of the ecological impact of extreme
events and inform prediction of responses to future change.

Keywords: biota, climate change, climate extremes, ecosystem functioning, river

could alter the biodiversity and ecosystem functioning of

rivers (Paine, Tegner & Johnson, 1998; Jones, 2013).
The frequency and intensity of extreme climatic events is Understanding the ecological effects of these events is
increasing (IPCC, 2012). Extreme events are statistically essential to predict ecological responses to future change.
rare or unusual weather or climatic occurrences, such as In this study, we synthesise the knowledge of extremes
extremes of precipitation or temperature, which can have and their impact on riverine ecosystems in Australia, a
severe natural impacts on the environment, including continent that has experienced multiple record-breaking
major floods, hydrologic droughts and fire (IPCC, 2012). events in the 21st century (Steffen, Hughes & Karoly,
Shifts in the regimes and coincidence of extreme events 2013).

Correspondence: Catherine Leigh, Irstea, UR MALY, 5 rue de la Doua, CS70077, 69626 Villeurbanne Cedex, France.

2620 © 2014 John Wiley & Sons Ltd

Extreme events and riverine ecosystems 2621
Australia spans multiple climate zones (Fig. 1a–c) and However, long-term research on the European summer
its rivers have a correspondingly wide range of flow heat wave of 2003 indicated these events can have
regimes, from predictably perennial to unpredictably severe, lasting impacts on benthic assemblages (Mou-
intermittent (Kennard et al., 2010). Although many spe- thon & Daufresne, 2006). In Australia, the prospect of
cies in these systems are adapted to natural variability unprecedented thermal extremes has renewed interest in
in climate and river flow, the persistence of taxa, partic- upper thermal limits (UTLs) of aquatic biota (e.g. Stew-
ularly those with restricted geographic and climatic art et al., 2013a). UTLs are typically determined experi-
ranges, may be threatened by climate extremes (Hughes, mentally using dynamic, non-lethal, behaviour-based
2011; Morrongiello et al., 2011). As the observed and methods (e.g. critical thermal method) or static, lethal
projected occurrence of extreme events in Australia methods (e.g. lethal temperature), with the former gen-
reflects that in many other regions of the world (IPCC, erally considered the better approximation to field con-
2012), ecological responses in Australia can help under- ditions. UTLs and the vulnerability of biota to heat
stand events elsewhere. waves and hot days are likely to depend on a variety of
We focus on six types of extreme events identified by factors, including antecedent ambient temperature and
the Australian Climate Commission as a threat to Aus- the rate and duration of warming in the environment,
tralian ecosystems (Steffen et al., 2013): heat waves and but data are scarce (Beitinger, Bennett & McCauley,
hot days, fires, droughts, heavy rainfall and floods, trop- 2000; Dallas & Rivers-Moore, 2012).
ical cyclones, and storm surges and coastal flooding. We Air temperatures in Australia regularly exceed the
examine their effects as reported in the literature, iden- UTLs of many aquatic taxa during the warmest months
tify key knowledge gaps and propose hypotheses to of the year, although the periods of exceedance may
stimulate research. only span a few hours in a daily cycle (Davies et al.,
2004), and this may also be the case during heat waves.
Fully aquatic species and those with long aquatic life
Heat waves and hot days stages may be less vulnerable to extreme air tempera-
tures than semi-aquatic species or those with extended
Definitions, observations and projections
aerial life stages, particularly where water temperatures
Heat waves, sometimes referred to as warm spells, are remain cool, or at least below UTLs, due to evaporative
periods of abnormally hot weather, but more precise cooling (Bogan et al., 2006), riparian shading or ground-
definitions vary regionally (IPCC, 2012; Perkins & water inputs. However, water temperatures that
Alexander, 2013). In Australia, heat waves are defined approach or exceed UTLs during heat waves, particu-
as at least three consecutive days of high maximum and larly those of extended duration or extreme intensity,
minimum temperatures that are unusual for a location and in hydrologically disconnected, shallow or still
(BOM, 2014a). High temperature extremes also include waters, could eliminate sensitive taxa (e.g. Matthews,
hot and very hot days (>35 °C and >40 °C, respectively) Surat & Hill, 1982). The projected intensification of heat
and warm days and nights (temperatures > 90th percen- waves threatens the survival of many riverine species
tile for any 5-day window across the annual cycle) (Fig. 2), and persistence may depend on adaptive man-
(BOM, 2014b). The duration and frequency of heat agement such as riparian revegetation and provision of
waves have increased in Australia since the 1970s, and cool-water refuges (Robson et al., 2013).
the hottest days of heat waves have become hotter (Per- Research to determine UTLs for stream invertebrates
kins & Alexander, 2013). The warmest year on record has revealed marked contrasts in vulnerability to
(2013) included a national summer heat wave c. 3 weeks extremes both within and among taxonomic groups. For
long (BOM, 2013). Climate projections indicate that the instance, UTLs of Ephemeroptera and Plecoptera
duration of heat waves and the number of hot days will nymphs are generally lower than those of Odonata and
continue to increase across Australia, consistent with Coleoptera larvae (e.g. Stewart et al., 2013a), although
projected global trends (Alexander & Arblaster, 2009; the mean UTL for each order varies substantially among
IPCC, 2012). geographic regions (e.g. southern versus northern taxa;
Davies et al., 2004). Some Ephemeroptera from south-
east Queensland can survive water temperatures >30 °C
Ecological effects
for at least 72 h (L. Luke, unpubl. data), which is well
The ecological effects of heat waves and hot days are above the global mean UTL for this group (c. 22 °C;
less well understood than those of hydrologic extremes. Davies et al., 2004; Stewart et al., 2013a).
© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
2622 C. Leigh et al.

(a) Darwin (d)


Alice Springs QLD

WA Brisbane
Perth NSW
Adelaide VIC Canberra,
ACT (e)

Hobart, TAS
Grassland Arid and
Desert Semi-arid Subtropical
Temperate, Mediterranean Tropical and
and Subalpine Equatorial

2500 5th percenle
Annual rainfall (mm)

2000 Median
95th percenle







Alice Springs

35 Daily minima
30 Daily maxima
temperature (°C)

Annual mean






Alice Springs

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
Extreme events and riverine ecosystems 2623

Droughts Knowledge gaps

Heat waves & hot days
(see Fig. 3)
Knowledge of thermal tolerance in aquatic organisms is
limited to coarse levels of taxonomic resolution, and less
↑ Air & water temperature is known about thermal tolerance among closely related
species or across populations (Chessman, 2009, 2013;
↑ Risk that upper thermal ↑ Risk of sub -
Stewart et al., 2013a). Organismal responses to extreme
limits of biota (including lethal effects of
temperature may be context dependent (Caissie, 2006;
aquac, riparian & terrestrial hot extremes
Verberk & Bilton, 2013), varying among habitats, and
taxa and life stages) are on biota
exceeded contingent upon the modifying influence of other
extreme events or stressors. For instance, thermal
Disrupon or
Δ Emergence rates impacts on biota may be exacerbated during drought
insgaon of
Death of of aquac when instream temperatures are less buffered against
spawning or
some biota macroinvertebrates temperature changes in air (Davies et al., 2004; Fig. 2),
dispersal cues in
some biota but data are scarce. More research is needed to identify
heat wave and hot day refugia for the most sensitive
Δ Reproducon taxa to direct management efforts, and there is a critical
& recruitment absence of research specifically examining effects of heat
waves and hot days at the community and ecosystem
Δ Aquac community composion
& ↑ threat to species’ persistence

Fig. 2 Effects of heat waves and hot days on rivers, as based on Droughts
links among drivers and responses identified in the Australian lit-
erature (see text for detail). Responses may be affected by the inten- Definitions, observations and projections
sity, duration and/or timing of heat waves and hot days. Solid
arrows show direct links, broken arrows where one event often Hydrological drought is a period of below-typical flow
coincides with or increases the likelihood or intensity of the linked that is unusual in its duration, extent or intensity
event. Ovals show where effects are hypothesised (event-specific
(Humphries & Baldwin, 2003). Drought is seasonal in
evidence scarce in the Australian literature). D, change in.
many systems, but here, we focus solely on ‘supra-sea-
sonal’ events: unpredictable droughts extending beyond
Vulnerability to heat waves can be influenced by seasonal periods, sometimes lasting >10 years (‘mega-
behavioural or physiological responses to warming. droughts’) (Lake, 2003; IPCC, 2012). Drought is a key
For example, high temperatures may influence the element of climatic and hydrological variability in Aus-
dispersal of some species (e.g. Hassall & Thompson, tralia. The Millennium Drought in south-eastern Austra-
2008), limiting escape of intolerable conditions. Sublethal lia was one of the most severe hydrological and
effects could also alter sex ratios in surviving populations. meteorological mega-droughts on record, lasting
For instance, temperature extremes forecast for the year between 1997 and 2009 (Gallant & Gergis, 2011; Timbal
2100 simulated in a mesocosm experiment increased & Fawcett, 2012; Fig. 1g). Mega-drought has also
emergence rates of male Ulmerophlebia pipinna mayflies affected the south-western region of Western Australia,
more so than females, implying a change in the sex ratio which has been in drought since the mid-1970s, with
that could have demographic and fitness consequences rainfall 10–20% below long-term averages (NCC, 2010).
and cause local extinctions (Thompson et al., 2013a). Climate projections indicate southern Australia will

Fig. 1 Australia has a range of climate zones (a), with regional variation in spatiotemporal patterns of temperature and rainfall (b, c).
Examples of extreme events include Cyclone Larry in March 2006 (d: aerial image of Upper North Johnstone River, post-event, showing
treefall and leaf denudation in the riparian zone; credit S. Turton), flood across much of south-east QLD in January 2011 (e: Gregors Creek,
post-event, during which flood waters reached the mature treeline above river banks; credit L. Luke), the 2003 wildfires in the ACT
(f: aerial image of Cotter River, post-event, showing burned vegetation across the catchment; credit T. Nelson), and the Millennium Drought
(g: a drying waterbody in the Ovens River catchment, 2009; credit S.S. Ho). Climate-zone mapping data and summary statistics sourced
from, with statistics calculated for the reference period 1 January 1961 to 31 December 1990. ACT, Australian Capital
Territory; NSW, New South Wales; NT, Northern Territory; QLD, Queensland; SA, South Australia; TAS, Tasmania; VIC, Victoria; WA,
Western Australia.

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
2624 C. Leigh et al.
experience future droughts of increased duration and chemistry (e.g. Sheldon & Fellows, 2010). Pools recede,
intensity (Risbey, 2011; Steffen et al., 2013). Similar and surface water can disappear completely (Fig. 1g).
changes may also occur in parts of Europe, the Mediter- Declining groundwater levels and increasing salinity in
ranean, the Americas and Africa (IPCC, 2012, 2014). groundwater-dependent systems and floodplains can
lead to loss of riparian and floodplain vegetation (e.g.
Murray et al., 2003; Fig. 3). In floodplain forests of the
Ecological effects
Murray–Darling Basin (MDB), stands of drought-tolerant
Supra-seasonal river droughts are ‘ramp’ disturbances, Eucalyptus camaldulensis suffered increased mortality
increasing in strength and spatial extent over time (Lake, during the Millennium Drought (Horner et al., 2009).
2003). In the initial stages of drought, riparian habitats Similarly, drought-related low flows have increased
disconnect from channels and littoral habitat dries (Boul- salinity levels in the lower lakes of the MDB (Mosley
ton, 2003). Lateral and longitudinal hydrological connec- et al., 2012) and contributed to the loss of vulnerable fish
tivity is reduced or lost as flow declines, creating populations (Nannoperca obscura; Wedderburn, Hammer
isolated pools (Fig. 3). In downstream river sections and & Bice, 2012). Ultimately, there is a shift from aquatic to
estuaries, reduced flow can limit the delivery of nutri- terrestrial habitat as organisms from surrounding ripar-
ents from upstream, alter productivity and trophic struc- ian areas move into dry channels (Steward et al., 2012).
ture and increase sediment accumulation (Lake, 2011). Biotic responses to drought vary depending on many
Once surface flows cease, many rivers enter a lentic factors, but in general, organisms that inhabit non-peren-
phase associated with major changes in water physico- nial systems have adaptive traits of resistance or resil-
ience to dry periods and may be more likely to survive
supra-seasonal drought than those from perennial sys-
tems (Lake, 2003; Rolls, Leigh & Sheldon, 2012; Fig. 3).
Fish species that tolerated the Millennium Drought in
↓Flow & flow Groundwater the MDB tended to have traits associated with adapta-
permanence decline tion to warm environments (e.g. high spawning temper-
atures), the ability to switch diets and life-history
↓ Connecvity ↑ Mortality of strategies including delayed maturation, short spawning
↓ Size &/or
of aquac riparian &
availability of
seasons and high fecundity (Chessman, 2013). The Mil-
aquac habitats habitats
vegetaon lennium Drought also altered macroinvertebrate com-
& refugia
↑ Buildup of organic & munities in Victorian streams, with conditions favouring
non-organic material in
↓ Influx of taxa tolerant of low flow and poor water quality (Rose,
dry channels &
nutrients from disconnected pools Heavy rainfall Metzeling & Catzikiris, 2008; Thomson et al., 2012). Des-
upstream to & floods iccation-resistant life-history stages also enable some
downstream (post-drought)
reaches (see Fig. 5) taxa (e.g. zooplankton and aquatic plant species, Brock
et al., 2003) to survive drought, although survival can
depend on drought duration. Cladoceran egg banks
↓ or Δ water ↑ Risk of hypoxic
from Australian dry land rivers became depleted when
quality blackwater
events drought lasted more than 6 years, which likely slowed
recovery of microinvertebrate communities and altered
Δ Community composions ↑ Risk of fish post-drought food webs (Jenkins & Boulton, 2007).
(↓sensive & flow- kills & threat
dependent, ↑ tolerant biota to other Drought survival depends on the type, quality and
& terrestrial biota) aquac taxa quantity of refugia and the hydrological history and con-
nectivity of habitats (Robson, Chester & Austin, 2011;
Fig. 3 Effects of droughts on rivers, as based on links among driv- Bogan, Boersma & Lytle, 2014). The temporal availability
ers and responses identified in the Australian literature (see text for
detail). Responses may be affected by the duration, extent and/or
of refugia and their position in river networks influence
intensity of droughts. For example, build-up of material on dry macroinvertebrate community resistance and resilience
channels and in pools and the risk of hypoxic blackwater events to drought (Marshall et al., 2006; Sheldon et al., 2010;
increase with duration and extent; disconnection among habitats Chester & Robson, 2011; Fig. 3). The rate of water
increases and the size and availability of habitats decline with
increased duration, intensity and extent; survival of aquatic taxa
decline may also affect the ability of some organisms to
declines with increased duration. See Fig. 2, for explanation of find refugia and influence the availability of basal
arrows, shapes and symbols. resources for riverine food webs (Bunn et al., 2006).
© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
Extreme events and riverine ecosystems 2625
particularly in the south-east, and projections indicate
Knowledge gaps
this trend will continue, with the annual fire season
Although the long-term (decadal) consequences of starting earlier and lasting longer (Lucas et al., 2007;
supra-seasonal drought are still poorly understood CSIRO & BOM, 2014). Projections are similar for New
(Bond, Lake & Arthington, 2008), recent research sug- Zealand and increased frequency of fire risk is projected
gests prolonged events can elicit novel community tra- for much of the Americas (IPCC, 2012).
jectories and extirpate sensitive species (Bogan & Lytle,
2011; Bogan et al., 2014). However, research is frequently
Ecological effects
opportunistic, sporadic and short term (e.g. <1–2 years),
and there is a pressing need for comprehensive monitor- One of the most significant and commonly reported
ing programmes to assess drought impacts and recovery effects of wildfire is the large-scale destruction of
processes systematically across regions and taxa (Rose terrestrial vegetation (Fig. 1f). In severely burned and
et al., 2008; Lake, 2011; Thomson et al., 2012). Research devegetated catchments, exposed soils become hydro-
typically focuses on macroinvertebrates, fish and ripar- phobic, increasing run-off and associated inputs of fine
ian vegetation, with less emphasis on other groups such sediment and nutrients (e.g. Lane, Sheridan & Noske,
as microbes and microinvertebrates. While drought 2006; Fig. 4). For instance, severe fires in Victoria sub-
impacts on the structure of some key assemblages are stantially elevated stream phosphorus concentrations
well described, consequences for ecosystem processes and conductivity (Chessman, 1986). Similarly, severe
and higher levels of biological organisation are less well burning in experimental forests in New South Wales
known (e.g. Ledger et al., 2013; Woodward et al., 2012). increased stream nitrate concentrations by an order of
Incorporating species resistance and resilience traits
into biomonitoring approaches will enhance mechanistic
and predictive understanding of drought impacts (e.g.
Fires Heat waves &
Sheldon, Marsh & Rolls, 2012; Chessman, 2013). In many Droughts hot days
cases, however, trait information is incomplete or held (see Fig. 3) (see Fig. 2)
in separate repositories and literatures (Crook et al., Destrucon of
2010). We also need to know how traits interact and bet- Heavy rainfall catchment & ↓ Habitat for
& floods riparian riparian terrestrial
ter understand relationships between traits and biotic
(post-fire) vegetaon fauna & Δ
interactions (Verberk, Van Noordwijk & Hildrew, 2013). (see Fig. 5) bioc community
For example, a trait such as air breathing that aids sur- composion
vival of anoxic conditions during drought may increase
predation risk where movement to the water surface Burned vegetaon
↑ Erosion ↓ Shading
increases encounter rates (Robson et al., 2011). Water- input
quality guidelines specific to drought periods are also
required to manage and protect surface and subsurface ↑ Fine sediment ↑ Nutrient
drought refugia, thereby supporting species persistence input input
(e.g. Sheldon & Fellows, 2010).
Δ Organic maer
↓ Water
Habitat smothered quality &/or
↓ Periphyton
Definitions, observations and projections cover

Much of Australia is vulnerable to wildfire due to the

Δ Composion of Δ Aquac community ↓ Abundance & ↑
dry, hot weather and volatile compounds in many of the funconal feeding composion (↓ sensive, mortality of
native plant species. In the north, the fires generally groups of consumers ↑ tolerant species) sensive species
occur in winter and spring (the dry season in northern
Australia), whereas in the south, they generally occur in Fig. 4 Effects of fires on rivers, as based on links among drivers
summer and autumn (BOM, 2009). Hot, dry and windy and responses identified in the Australian literature (see text for
detail). Responses may be affected by the intensity and/or extent
conditions increase fire danger (Clarke, Smith & Pitman, of fires, for example by increasing the extent and quantity of vege-
2011), linking heat waves, drought and strong winds to tation destroyed or reducing the survival or abundance of species.
fire. Since the 1970s, extreme fire weather has increased, See Fig. 2, for explanation of arrows, shapes and symbols.

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
2626 C. Leigh et al.
magnitude (Mackay & Robinson, 1987). Changes to fire, although there is also some uncertainty regarding
riparian zones may also affect incident light and alter short-term impacts, which may be underestimated when
temperature regimes. post-fire sampling is delayed (Verkaik et al., 2014).
The ecosystem consequences of fire can be exacer- Research on the influence of land use on post-fire recov-
bated by heavy rains (Cooper et al., 2014), with the com- ery is rare, even though this likely determines recovery
bined, interactive impacts of these two extremes creating rates. In the Lower Cotter River catchment, for example,
a compound event (sensu IPCC, 2012) characterised by stream turbidity recovered more quickly in naturally
intense sedimentation and turbidity (Fig. 4). In the Cot- regenerating, native forest than in plantation pine forest
ter River catchment of the Australian Capital Territory, a (Harrison et al., 2014). Research on interactions between
large storm 1 month after severe wildfire raised stream fire and other extreme events such as drought and flood
turbidity to an unprecedented level of 3000 NTU (White is also needed. The trajectory of community recovery
et al., 2006). By contrast, fires not followed by heavy rain from fire during drought periods may be altered by
tend to have less impact on rivers, as was the case in burning of dewatered benthic sediments (Cowell, Mat-
Australia’s wet–dry tropics, where fires occurring early thews & Lind, 2006) or ash physicochemistry (Verkaik
in the dry season had little effect on water quality et al., 2014), but our understanding of such interactions
(Townsend & Douglas, 2000). is incomplete.
Declines in abundances and richness of instream
fauna are often observed following wildfire (Fig. 4). For
instance, macroinvertebrate communities in the most Heavy rainfall and floods
severely burned areas of the Cotter River catchment
Definitions, observations and projections
were impoverished compared with those in unburned
areas (T. Nelson, unpubl. data). However, the scale of Heavy rainfall events are periods of prolonged and/or
the effects depends on the intensity of fire and post-fire intense rainfall within a specific location relative to
rainfall (Papas, 1998). A range of conditions in post-fire recorded averages and are often defined by expected
streams, including heavy sedimentation and poor water return intervals over decades or centuries (Steffen et al.,
quality (e.g. dissolved oxygen sags, chemicals from 2013). The events are often associated with inland flood-
burned material), has also been associated with reduced ing because surface water accumulations and run-off
survival and abundance of fish (Carey et al., 2003; Lyon can rapidly inundate dry lands and floodplains. Austra-
& O’Connor, 2008). lian indices used to describe these events include heavy
Fires also affect the availability and quality of trophic and very heavy precipitation days (≥10 and 30 mm,
resources. In south-eastern Australian streams, sodium respectively) and very wet and extremely wet days
concentrations increased in burned Eucalyptus viminalis (annual total precipitation when daily precipitation
leaves during post-fire decomposition, and changes in >95th and 99th percentiles, respectively) (BOM, 2014b).
microbial activity ultimately reduced the amount of bio- Several of these extremes occurred across eastern Aus-
available carbon (P. Love, unpubl. data). In turn, fire- tralia in 2010–2012, breaking the Millennium Drought.
induced changes in the quality and quantity of trophic Severe flooding occurred in central and southern
resources can alter the composition of functional feeding Queensland in 2010–2011, during a period including the
groups (Verkaik et al., 2014; Fig. 4). For example, sub- state’s wettest December on record (NCC BOM, 2011;
stantial inputs of fine suspended sediment after fire in Fig. 1e). Both central and northern Australia experienced
the Cotter River catchment scoured and smothered the wettest dry season on record in 2010 and record
periphyton, reducing the occurrence of herbivorous heavy rainfall and flooding occurred in most states and
scrapers (Peat, Chester & Norris, 2005; T. Nelson, un- territories in 2010, 2011 and 2012 (NCC BOM, 2011,
publ. data). 2012; BOM, 2012a). Heavy rainfall intensity is expected
to increase in Australia in the 21st century, whereas the
number of events may increase, decrease or remain
Knowledge gaps
unchanged depending on region (Alexander & Arblast-
The effects of fire can be immediate and also extend to er, 2009), matching projections elsewhere (IPCC, 2012;
the short term (<1 year), mid term (1–10 years) or long Donat et al., 2013). Such changes in rainfall patterns will
term (>10 years) (Minshall, Brock & Varley, 1989). Least likely affect flood regimes, altering the intensity, dura-
is known about the long-term ecological consequences of tion, frequency and timing of events.

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
Extreme events and riverine ecosystems 2627

Ecological effects Tropical Heavy rainfall

cyclones & floods
The ecological importance of floods in riverine ecosys- (see Fig. 6)
tems is well documented, both within Australia and
internationally. Floods influence ecosystem processes, ↑ Connecvity of ↑ Catchment & Groundwater
affecting within-channel productivity, particulate organic aquac habitats channel erosion recharge
matter retention, decomposition, nutrient transforma-
tions and microbial processes (Baldwin & Mitchell, 2000;
Triggers Movement of Watering of
Hladyz et al., 2012). Floods also affect terrestrial flood- dispersal, large amounts of riparian &
plain biota (Horrocks et al., 2012) and influence surface spawning & organic & floodplain
water and groundwater quality and recharge (Cend on recruitment inorganic maer vegetaon
opportunies (including biota)
et al., 2010) (Fig. 5). Rising waters increase hydrological Droughts
connectivity at various spatial scales, particularly in arid, (pre-flood)
(see Fig. 3)
semi-arid and wet–dry zones (Douglas, Bunn & Davies, ↑ Spread of
2005; Leigh et al., 2010). This facilitates transport of (e.g. invasive) ↑ Risk of hypoxic
materials, such as propagules, nutrients, organic matter species Δ Water blackwater
quality events
and woody debris, both downstream and to adjacent
floodplains and riparian zones (Ballinger & Lake, 2006; ↑ Risk of fish kills
& threat to other
Erskine et al., 2012; Fig. 5). Floods may also cause aquac taxa
siltation and ‘sand slugs’, alter channel morphology, Booms in producon & Δ Community composion
scour substrata, remove in-situ basal resources such as Δ trophic interacons (↓ sensive, ↑ tolerant biota)
macrophytes and detritus and translocate, injure or kill
Fig. 5 Effects of heavy rainfall and floods on rivers, as based on
biota (Lake, 2000; Fig. 5). links among drivers and responses identified in the Australian lit-
Much research has focused on flood-driven hydrologi- erature (see text for detail). Responses may be affected by the inten-
cal connectivity and food webs. Floods drive spatial sub- sity, timing and/or extent of these events. For example, the extent
sidies in food resources and affect the relative of habitat connectivity and amount of channel erosion and scouring
increase as these events increase in intensity and extent; timing of
importance of autochthonous and allochthonous carbon
these events relative to drought will affect the risk of hypoxic
sources (Bunn et al., 2006; Reid et al., 2008). In some blackwater events; timing and intensity of events relative to life-his-
cases, mobile consumers such as fish and birds may tory stages and age of biota may affect the chance of translocation,
transfer resources between habitats connected by inun- injury or death. See Fig. 2, for explanation of arrows, shapes and
dation (Jardine et al., 2012). River–floodplain connections
can also facilitate ‘boom-bust’ cycles in basal productiv-
ity and invertebrate emergence, with flow-on effects to 2012; McCarthy et al., 2014). However, non-native fish
other components of the food web (Boulton & Lloyd, such as Cyprinus carpio were apparently unaffected.
1992; Bunn et al., 2006; Fig. 5). The floods of 2010–2011 Floods can facilitate the spread of non-native and
increased taxonomic richness and abundance of zoo- invasive species. Widespread flooding has probably
plankton in several southern-MDB rivers (Ning et al., facilitated the spread of Phyla canescens, an invasive
2013). In central Australia, post-flood booms in fish pro- riparian plant, across eastern Australia (Murray, Stokes
duction subsequently fuelled high levels of heterotrophic & Van Klinken, 2011; Fig. 5). By contrast, remnant popu-
bacterial production once waters (and fish) retracted to lations of native freshwater mussels are threatened by
isolated waterholes (Burford et al., 2008). extreme floods, particularly where events exacerbate
Floods also alter water quality. Hypoxic blackwater existing impacts of catchment degradation (Jones & By-
events can result from the microbial breakdown of rne, 2010). However, accumulation of large woody deb-
deposited or leached organic matter, although their ris during flooding can facilitate post-flood recovery of
intensity and extent are often exacerbated by pre-flood native riparian vegetation (e.g. in Australia’s semi-arid
drought and human activities (Whitworth, Baldwin & savannah; Pettit & Naiman, 2005) and, in some cases,
Kerr, 2012; Figs 4 & 5). As the Millenium Drought was floods favour native over non-native species (Costelloe
breaking in late 2010, one such event killed many native et al., 2010; Ho, Bond & Thompson, 2013). In south-east-
crustaceans and fish and led to the emergence of native ern Australia, flooding increased species richness of
crayfish from affected waters (King, Tonkin & Lieshcke, native plants in E. camaldulensis stands, with twice as

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
2628 C. Leigh et al.
many natives found in flooded as unflooded areas found in the Australian region, projections for tropical
(Horner et al., 2012). zones in Australia and beyond suggest they may become
less frequent but more intense (CSIRO & BOM, 2012;
IPCC, 2012). The proportion of severe events is therefore
Knowledge gaps
likely to increase.
Floods that are aseasonal, recurrent or interact with
other extreme events (e.g. fire and drought) or land-use
Ecological effects
degradation, can markedly affect river biodiversity.
However, explicit studies of such phenomena are scarce. As with fire, one of the most commonly reported effects
The timing of heavy rainfall and extreme floods relative of tropical cyclones is vegetation damage, with riparian
to key stages of aquatic life cycles may influence recruit- zone vegetation being particularly susceptible (Fig. 6).
ment, dispersal and foraging opportunities (e.g. in fish, In the forests of Magela and Ngarradj Creek catch-
Humphries, King & Koehn, 1999), and shifts in flood fre- ments, Northern Territory, wind gusts during STC
quency and intensity may affect the persistence or Monica created significant treefall and riparian zones
spread of invasive species, for example by providing sustained greater damage than did other areas (Saynor
opportunities for passive or active dispersal within and & Erskine, 2008; Staben & Evans, 2008). In some parts
between river networks. However, more research is of the Ngarradj Creek catchment, the entire riparian
needed to determine the vulnerability and dispersibility rainforest was destroyed. STC Monica occurred at the
of invasive species during floods. Impacts of extreme end of the tropical wet season when riparian zone soils
floods on genetic diversity of threatened populations are were saturated and streams inundated, so these factors
also uncertain, although it has been proposed that more probably exacerbated the damage (Saynor & Erskine,
large-scale floods could increase gene flow, while more 2008; Fig. 6).
intense floods could create population bottlenecks (Wor- Riparian vegetation was also damaged during STC
thington Wilmer et al., 2011). Research on ecological out- Larry (Fig. 1d). Six months after that event, pioneer spe-
comes of environmental flows featuring managed cies in north-east Queensland rainforests were more
flooding may help us understand impacts of future flood severely damaged than other species. They suffered dis-
regimes, but existing data are biased towards particular proportionately more uprooting, particularly along
taxa (e.g. fish and floodplain vegetation) and systems stream edges (Pohlman, Goosem & Turton, 2008). This
(e.g. MDB rivers; Aldous et al., 2011). We require more was associated with sandy soils near streams, producing
rigorous, high-resolution hydrological models (Pittock loose root systems, and bank erosion during post-cyclo-
et al., 2006), and these must be better integrated with nic floods. However, while damage to riparian vegeta-
ecological models to better predict relationships between tion can be substantial, the associated delivery of large
changing rainfall, future flood regimes and ecological woody debris to rivers contributes important habitat
responses. and increases channel complexity (Erskine et al., 2012;
Fig. 6). After STC Monica in the Ngarradj Creek catch-
ment, the number of pieces of large wood per metre of
Tropical cyclones channel more than doubled (Erskine et al., 2012).
Tropical cyclones producing heavy rainfall change
Definitions, observations and projections
water quality in rivers (Fig. 6). Rainfall following Tropi-
Tropical cyclones are low-pressure systems that develop cal Cyclone Celeste in north-east Queensland (January
over warm, tropical marine-waters and are associated 1996) rapidly increased water level and discharge in the
with gale-force winds (sustained at ≥63 km h 1 or in Burdekin River (Alexander et al., 2001). River water
gusts >90 km h 1) (BOM, 2014c). Winds sustained at became highly turbid, with suspended solids concentra-
≥118 km h 1 (or gusts >165 km h 1) qualify events as tions >4000 mg L 1. Conductivity decreased rapidly but
severe tropical cyclones (STCs), also known as hurri- recovered to pre-event conditions within 1–2 weeks. In
canes or typhoons. Well-documented STCs of the last contrast, increased suspended solids concentrations were
50 years in Australia include Tracy, Larry (Fig. 1d), observed up to a year following STC Monica (Evans &
Monica and Yasi (BOM, 2014d). STC Monica was associ- Moliere, 2010). Differences in the effects of Celeste and
ated with maximum recorded winds of 130 km h 1 and Monica on water quality were likely influenced by dif-
rainfall of up to 340 mm in 24 h. Although no clear ferences in their timing relative to the first, flushing
trends in tropical cyclone characteristics have been flows of the wet season (Alexander et al., 2001), their
© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
Extreme events and riverine ecosystems 2629

Storm surges & on event timing, frequency and intensity (Barron et al.,
Tropical 2011; McGrath et al., 2012; Fig. 6). Past long-term
coastal flooding
cyclones declines in tropical cyclone frequency are associated
(see Fig. 7)
with ‘water storage droughts’ and desertification (Nott,
2011; McGrath et al., 2012). Tropical cyclones in northern
Heavy rainfall & floods
↑ Intensity of Australia also deliver rains to more southerly regions as
(see Fig. 5)
winds &/or they degenerate to extropical lows during their overland
wind gusts trajectories. These rains supply water that produces
Groundwater flows and sustains biota in temporary wetlands of dry-
recharge land and semi-arid river systems (Roshier et al., 2001)
Sandy ↑ Saturaon ↑ Intensity & and in catchments that discharge into the Great Barrier
riparian of riparian extent of stream Reef lagoon (Boer, 2010).
soils soils & floodplain
Knowledge gaps
↑ Damage to
Much of our knowledge of tropical cyclone effects is
catchment & Wetland
especially riparian drawn from vegetation studies. We know little about
replenishment how they affect instream biota (e.g. fish, invertebrates;
although see Mallin et al., 1999) and ecosystem pro-
↑ Erosion
cesses, either directly or via indirect effects on riparian
zones and water quality (Fig. 6). Although we have
some evidence suggesting cyclone (and flood) timing
↑ Delivery of ↓ Shading ↑ Fine sediment, affects ecological responses (e.g. Alexander et al., 2001;
large woody organic maer & Saynor & Erskine, 2008), our understanding of this
debris to streams nutrient input aspect and cyclone–flood interactions is quite limited.
Moreover, the lack of studies combining pre- and
↑ Instream habitat for post-event data limits understanding of impacts and
aquac biota & ↑ Δ Water quality recovery.
channel complexity

Storm surges and coastal flooding

Δ Aquac
↑ Diversity of fish & community Definitions, observations and projections
macroinvertebrates composion
Coastal flooding results from high water levels pro-
duced by sea-level fluctuations during short-term,
Fig. 6 Effects of tropical cyclones on rivers, as based on links localised storm-surge and wave events, and longer term
among drivers and responses identified in the Australian literature sea-level rise (Eliot, 2012). The primary cause of surge-
(see text for detail). Responses may be affected by the intensity,
related coastal flooding is the landfall of low-pressure
timing and/or frequency of these events. For example, saturated
soils are more likely during wet-season events; events before the systems with strong winds (CSIRO & BOM, 2007).
wet season’s first flushing flows increase inputs of sediment, More intense events occur when storm surge coincides
organic matter and nutrients to streams and the time taken to with high tides, and events become more frequent as
return to pre-event conditions; the timing and frequency of events
sea level rises (S anchez-Arcilla et al., 2008; Fig. 7).
affect the sporadicity of groundwater recharge. See Fig. 2, for
explanation of arrows, shapes and symbols. Surge-related coastal flooding can occur anywhere
topography allows (CSIRO & BOM, 2007; BOM, 2012b).
However, projected changes in storm-surge and inun-
intensity (e.g. erosive forces on soils and sediments) and dation impacts vary regionally and can depend on the
the amount of damage caused to riparian vegetation method by which storm-tide intervals are estimated
(Fig. 6). (CSIRO & BOM, 2007). It is very likely that mean sea-
Recharge of water stores in northern Australia and level rise will continue to contribute to global upward
aquifers in central Australia relies heavily on tropical trends in extreme coastal high water levels (IPCC,
cyclones, and thus recharge can be sporadic depending 2012).
© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
2630 C. Leigh et al.
term shifts in fish community composition were
Tropical cyclones (see Fig. 6)
Storm surges & observed following saltwater surge into freshwater
coastal flooding reaches of rivers near Hurricane Katrina’s landfall zone
Sea-level Heavy rainfall (Schaefer et al., 2006), amphibian species in freshwater
rise & floods coastal wetlands showed resistance to water chemistry
High des (post-surge) changes (Gunzburger et al., 2010). Recovery of freshwa-
(see Fig. 5) ter fauna following saltwater flooding is likely influ-
enced by dispersal characteristics, with active dispersal
Intrusion of
(e.g. swimming) contributing to rapid recovery (Watana-
saline water into Abrupt Δ salinity Sediment be et al., 2014).
groundwater & in lowland river redistribuon
↑ salinity in secons (fresh to Understanding of storm-surge effects in rivers could
& ↑ erosion
groundwater- saline to fresh) be informed by knowledge of species salinity tolerances,
dependent and by studies of tsunamis (e.g. Watanabe et al., 2014) or
rivers salinisation from progressive saltwater intrusion and
Δ Water sea-level rise. However, ramp disturbances such as pro-
quality gressive salinisation may have different impacts from
pulsed events like storm surges, where recovery to pre-
Death or ↓ abundance
of saline-intolerant species, disturbance (freshwater) conditions can occur rapidly.
&/or Δ aquac community On the other hand, storm surges can rework sediments
composion and channels and may even cause mass mortalities
requiring long recovery periods (Cahoon, 2006). Intru-
Fig. 7 Effects of storm surges and coastal flooding on rivers, as sion of saline waters into ground water during surges
based on links among drivers and responses identified in the Aus- may also prolong recovery of aquatic biota in groundwa-
tralian literature (see text for detail). See Fig. 2, for explanation of ter-fed rivers (Fig. 7). Furthermore, surge-induced flood-
arrows, shapes and symbols.
ing may be followed by highly turbid floods (of inland
origin) associated with the same cyclone or storm (e.g.
Gong & Shen, 2009), causing abrupt changes from fresh-
Ecological effects
water to saline water to turbid freshwater conditions.
Storm surges alter habitats by pushing sea water into
estuaries, brackish lagoons and wetlands, which in turn
Future directions and concluding remarks
can redistribute sediment, contribute to erosion (Cahoon,
2006; Howes et al., 2010) and force saline water into fresh- Extreme events are locally and globally important, with
water river sections (CSIRO & Bureau of Meteorology impacts often disproportionate to their duration (Smith,
(BOM), 2007; Kang & King, 2012; Fig. 7). Surge-related 2011). The ecological effects of extremes can be difficult
effects of increased salinity and geomorphological to determine from opportunistic studies, and more
changes on riverine biota (e.g. aquatic macroinverte- research is needed to develop a sound mechanistic
brates) are, however, understudied (Kang & King, 2012), understanding of responses to inform prediction of
and most research has been performed outside Australia, future events, where they occur singly or in combina-
primarily on aquatic macrophytes (e.g. Frazer et al., 2006). tion. Among event types, research on floods, droughts
and fires is more extensive than for heat waves, tropical
cyclones and storm surges. While research on all events
Knowledge gaps
is needed, these latter events remain a particular priority
Effects of storm surge and coastal flooding on riverine for future research.
ecosystems are poorly known in Australia and globally. As the frequency of extreme events increases, the his-
Studies from North America suggest aquatic vegetation tory of past events will become increasingly important in
in coastal fresh waters is susceptible to salinity changes shaping responses to subsequent extremes (IPCC, 2012).
and erosion from storm surge (e.g. Frazer et al., 2006). Compound events will be particularly deleterious, as has
However, these studies have focussed on wetlands, and been observed where heavy rains and flooding follow fire
their applicability to rivers is unclear (Fig. 7). Responses or drought. The intensification and coincidence of
to coastal flooding may also differ among and within extreme events may elicit regime shifts, novel interactions
taxa, but again, knowledge is incomplete. While short- and/or abrupt transitions in species distributions and
© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
Table 1 Hypotheses and potential approaches to explore the impact of extreme climatic events on freshwater habitat and across levels of biological organisation

Events and future

projections Hypotheses* Approaches*

Heat waves and hot Habitat Measurement of habitat characteristics to identify

days will increase in Increased importance of cool-water refuges for heat-sensitive taxa thermal refugia
number and duration Species and populations Experiments applying high temperature extremes
More frequent exceedance of upper thermal tolerance limits of biota of different durations and frequencies,
Reduced resilience (recovery) of heat-sensitive populations downscaling climate models to generate realistic
Local extinction or contraction in distributions of heat-sensitive taxa temperature extremes and regimes
Communities Modelling to predict changes in distributions of
Loss of favourable habitat and species and immigration of new species will alter community heat-sensitive taxa, particularly for known
structure vulnerable taxa
Organic matter processing may alter due to local extinctions and behavioural effects of heat stress on
primary consumers
Droughts will increase Habitat and events Experiments applying different hydrological (dis)
in intensity, extent Increased extent of hydrological disconnection (terrestrialisation); habitat loss, including drought connectivity treatments and drying disturbances
and duration refugia; increased importance of remaining refugia of different intensities over multiple
Increased incidence of droughts interacting with other extreme events (compound events) spatiotemporal scales
Species and populations Experiments applying ‘drought disturbances’ in

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
Loss or contraction in distributions of vulnerable taxa (including flow-dependant taxa) combination with other extreme event treatments
Communities Experiments on dry-stream organic matter
Changes in richness and community composition resulting from changes in habitat and/or local breakdown, pre-conditioning and consumption or
species extinctions, including loss of flow-dependent taxa from stream networks and changes in use by biota
vegetation type, and increased likelihood of novel community trajectories Modelling to predict changes in availability and
Functioning configuration of drought refugia and their
Accumulation of organic matter on dry streambeds; increased occurrence of hypoxic blackwater accessibility to vulnerable taxa
events; altered organic matter processing rates in streams; increased fuel load for and risk of fire;
potentially driving shifts in behaviour (e.g. foraging) and composition of terrestrial biota
Extended periods of reduced nutrient input into downstream (e.g. lowland) reaches
Changes in food-web structure and dynamics via loss of rare taxa and predators
Fires will increase in Habitat and events Experiments manipulating the quantity and quality
frequency, the risk of Accumulation of fine sediment on streambeds and reduction in stream habitat size and quality of organic matter in streams
fire will increase, and Increased exposure of stream channels to sun from decrease in canopy cover, increasing stream Experiments manipulating sediment and ash
annual fire seasons temperatures and exacerbating effects of heat waves and hot days inputs under both dry- and wet-stream conditions
will start earlier and Increased incidence of fire–flood interactions (compound events) Modelling to predict extent of sediment
last longer Species and populations accumulation and transport in streams, including
Loss or contraction in distributions of heat-sensitive taxa and those sensitive to fine sediment regions with vulnerable taxa, communities and/
accumulation or ecosystems
Changes in richness and community composition resulting from changes in habitat, local species
extinctions and/or responses to changes in organic matter quality and quality (see Functioning)
Changes in vegetation type and structure in riparian zones
Extreme events and riverine ecosystems

Increased quantity of refractory carbon (charcoal) and reduced quality of organic matter in streams
and resultant changes to food webs
Table 1 (Continued)

Events and future

projections Hypotheses* Approaches*

Heavy rainfall (and Habitat and events Experiments applying ‘flood disturbances’
floods) will increase Increased frequency, intensity and inundation extent and novel timing of floods (hydrological successively and/or in combination with other
in number and connection events) extreme event treatments
intensity Increased occurrence of successive floods and floods occurring in combination with other extreme Experiments applying different hydrological
C. Leigh et al.

events (compound events) connectivity treatments over multiple spatial

Increased occurrences of channel erosion scales
Species and populations Modelling to predict changes in recruitment
Increased frequency of ‘habitat-connecting’ events will facilitate spread of species, including success and distributions of vulnerable taxa and
invasives, and the transport of other materials (organic and inorganic) invasives
Novel timing of events may trigger spawning cues in some species possibly leading to unsuccessful
recruitment, for example if spawning occurs when seasonal conditions (e.g. temperatures) are not
suitable for development
Changes in richness and community composition resulting from changes in habitat, local species
extinctions and/or invasions
Increased transport of organic and inorganic materials within and between catchments
Tropical cyclones will Habitat Experiments applying ‘wind treatments’ of
increase in intensity Increased destruction of riparian vegetation (treefall and other wind damage) and increased delivery different strength and duration under high-flow
of large woody debris to streams and control-flow conditions
Increased sheltering habitat for fish and macroinvertebrates and increased substrate for biofilm Experiments and surveys to establish how loss and
growth (food for consumers) recovery of riparian vegetation, and changes in
Increased intensity of water-quality changes following events (e.g. extreme increases in turbidity), turbidity, alter food webs and aquatic community
with prolonged recovery composition
Species and populations Modelling to predict extent and duration of
Creation of (temporarily) unsuitable habitat for turbidity-sensitive aquatic biota, potentially causing turbidity and other water-quality changes during
local extinctions and after events of different intensity
Substantial damage to or loss of wind-sensitive taxa in riparian zones and increased risk of damage
to wind-tolerant taxa
Changes in richness and community composition resulting from changes in habitat and/or local
Altered food-web dynamics resulting from changes in riparian vegetation and organic matter inputs
Storm surges will Habitat Experiments applying abrupt water-quality
increase in frequency Increased frequency of abrupt changes in salinity in freshwater sections of rivers and increased changes
and coastal flooding likelihood of events affecting sections further upstream than previously known Modelling to predict extent and persistence of
will increase in extent Increased temporal and spatial variability in channel morphology saltwater inundation, particularly in regions with
Species and populations vulnerable taxa, communities and/or ecosystems
Loss or contraction in distributions of salinity-sensitive taxa
Terrestrial and aquatic communities of affected rivers become more similar to those typically found
in estuarine habitats

*Response variables in relation to these hypotheses and corresponding approaches may include but are not restricted to aerial and aquatic stages of aquatic macroinvertebrates; fish;
terrestrial and aquatic plants; microbial communities; food webs; leaf-litter breakdown rates and organic matter processing dynamics; water-quality parameters; availability, quality

© 2014 John Wiley & Sons Ltd, Freshwater Biology, 60, 2620–2638
and position of refugia.
Extreme events and riverine ecosystems 2633
ecosystem functioning (IPCC, 2014). Successive events of the combined effects of multiple stressors (e.g. Lunt, Jan-
the same type may alter the organisation of food webs sen & Binns, 2012) will help fill knowledge gaps and
and disrupt the successional development of biological better enable us to predict responses to the events and
communities (e.g. Ledger et al., 2012, 2013) or initiate novel regimes of the future.
novel development trajectories (Turner et al., 1998).
Understanding of compound events and long-term Acknowledgments
effects of extreme events could be enhanced by exploit-
ing long-term data collected by monitoring programs for We thank Pettina Love, Tom Nelson and Anthea Flo-
other purposes such as bioassessment but which capture rance for information on fires from their Honours
pre- and post-event information (e.g. Rose et al., 2008). research at the University of Canberra; Steve Turton,
Standardisation and maintenance of monitoring pro- Campbell Clarke and Tom Nelson for photographs; and
grams across regions, along with question-driven pro- Andrew Boulton, Sarah Boulter, Dave Strayer and two
grams and adaptive designs would also improve the anonymous reviewers for their comments and edits that
utility of long-term datasets to elucidate mechanisms improved the manuscript.
underlying ecological responses to and recovery from
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