Neil E. Pettit

Fire in the riparian zone: Characteristics and ecological consequences

We review the current understandings of the frequency, spatial distributions, mechanisms, and ecological consequences of fire in riparian zones. Riparian zones are well known for influencing many ecological processes at local to landscape scales, and fire can have an important ecosystem-scale influence on them. Riparian zones differ from surrounding uplands in their biophysical templates, moisture regimes and disturbance regimes; as a consequence the characteristics and effects of fire are different than in adjacent uplands. Fire impacts on riparian zones vary proportionally with the severity and extent of burning in the catchment and are affected by stream size. Riparian zones can act as a buffer against fire and therefore as a refuge for fire-sensitive species. However, under some circumstances, such as dry pre-fire climatic conditions and the accumulation of dry fuel, riparian areas become corridors for fire movement. Fire incursion into riparian zones creates canopy gaps and drier conditions, which allow subsequent build up of dead wood and establishment of fire adapted species. In concert, this increases fuel loads and the probability of another fire. Secondary effects of riparian fire include altering nutrient fluxes and cycling, increasing sediment loads, and stimulating erosion. We conclude that riparian fires are potentially important in shaping ecological characteristics in many regions, but this is poorly quantified. A better understanding of riparian fire regimes is essential to assess the effects of fire in helping shape the complex ecological characteristics of riparian zones over the longer-term.

Flow–ecology relationships: closing the loop on effective environmental flows

Providing flows for biota and environmental processes is a challenging water management issue. For society the ability and willingness to allocate water to sustain the environment is increasingly competitive due to escalating demand and as a consequence of climate change. In response, an array of environmental flow (E-flow) methods have developed. Our view is that few E-flows have been implemented and even fewer evaluated in a research and management context. Much of our science effort in E-flows has been directed primarily at method development, with less attention being given to monitoring, evaluation and subsequent revision of E-flow strategies. Our objectives are to highlight the lack of connection between current trends in E-flow literature and theory with assessment of the efficacy and practical application of these methods. Specifically, effective E-flows need to be explicit about flow-ecology relationships to adequately determine the amount and timing of water required. We briefly outline the historical development of E-flows and discuss how serial development of methods and techniques has restricted implementation, evaluation and revision. We highlight areas where methods are lacking, such as incorporation of data on flow-ecology relationships into operational use of E-flow methods. We suggest four initial steps that will improve the applicability, implementation and ultimate success of E-flows.

Consumer - resource coupling in wet-dry tropical rivers

1. Despite implications for top-down and bottom-up control and the stability of food webs, understanding the links between consumers and their diets remains difficult, particularly in remote tropical locations where food resources are usually abundant and variable and seasonal hydrology produces alternating patterns of connectivity and isolation. 2. We used a large scale survey of freshwater biota from 67 sites in three catchments (Daly River, Northern Territory; Fitzroy River, Western Australia; and the Mitchell River, Queensland) in Australias wet-dry tropics and analysed stable isotopes of carbon (δ(13) C) to search for broad patterns in resource use by consumers in conjunction with known and measured indices of connectivity, the duration of floodplain inundation, and dietary choices (i.e. stomach contents of fish). 3. Regression analysis of biofilm δ(13) C against consumer δ(13) C, as an indicator of reliance on local food sources (periphyton and detritus), varied depending on taxa and catchment. 4. The carbon isotope ratios of benthic invertebrates were tightly coupled to those of biofilm in all three catchments, suggesting assimilation of local resources by these largely nonmobile taxa. 5. Stable C isotope ratios of fish, however, were less well-linked to those of biofilm and varied by catchment according to hydrological connectivity; the perennially flowing Daly River with a long duration of floodplain inundation showed the least degree of coupling, the seasonally flowing Fitzroy River with an extremely short flood period showed the strongest coupling, and the Mitchell River was intermediate in connectivity, flood duration and consumer-resource coupling. 6. These findings highlight the high mobility of the fish community in these rivers, and how hydrological connectivity between habitats drives patterns of consumer-resource coupling.

Dissolved organic carbon biolability decreases along with its modernization in fluvial networks in an ancient landscape

The metabolism of dissolved organic carbon (DOC) along fluvial networks determines what fraction of organic matter is exported to the ocean. Although it is thought fresh rather than older DOC is preferred by bacteria, old DOC can also be highly bioavailable to stream bacterial communities. In strongly seasonal and oligotrophic regions, we argue that groundwater inputs of old DOC may increase the bioavailability of stream organic matter. We sampled 22 streams along a gradient of size (wetted widths from 1 to 60 m) and one groundwater spring in the Kimberley region of northwest Australia to determine how the age and bioavailability of streamwater DOC varied with stream size. Our hypothesis was that stream DOC would become more enriched in 14C (younger) and less bioavailable as streams increased in size and depleted 14C-DOC was metabolized by stream microbial communities. We also used fluorescence characterization of DOC, ultraviolet absorbance at 254nm (SUVA254), δ13C-DOC and lignin phenol yields to assess how these indicators of DOC character influenced the bioavailability and age of stream DOC. Stream evaporation/inflow ratios (E/I, used as a proxy for catchment water residence time), determined from changes in stream δ18O along the gradient of stream size, were positively related to DOC concentration and carbon-normalized lignin yields, while δ13C-DOC became more depleted with increasing E/I. Stream Δ14C-DOC varied from −452.1‰ (groundwater) to 48.9‰ and showed progressive enrichment as streams increased in size and accumulated DOC mainly from terrestrial plant material. Older DOC corresponded to higher bioavailability (R2 = 0.67, P < 0.01), suggesting that old bioavailable DOC, which has escaped from subterranean food webs utilizing 14C-depleted carbon, is common to one of the oldest landscapes on earth. Therefore, rapid biotic uptake of old bioavailable DOC originating in groundwater springs and the accumulation of modern, terrestrially derived DOC work in opposite directions affecting DOC dynamics along fluvial networks. We suggest the metabolism of old DOC along fluvial networks provides a biogeochemical link between non-contemporary carbon fixation and modern river productivity.

Productivity, Disturbance and Ecosystem Size Have No Influence on Food Chain Length in Seasonally Connected Rivers

The food web is one of the oldest and most central organising concepts in ecology and for decades, food chain length has been hypothesised to be controlled by productivity, disturbance, and/or ecosystem size; each of which may be mediated by the functional trophic role of the top predator. We characterised aquatic food webs using carbon and nitrogen stable isotopes from 66 river and floodplain sites across the wet-dry tropics of northern Australia to determine the relative importance of productivity (indicated by nutrient concentrations), disturbance (indicated by hydrological isolation) and ecosystem size, and how they may be affected by food web architecture. We show that variation in food chain length was unrelated to these classic environmental determinants, and unrelated to the trophic role of the top predator. This finding is a striking exception to the literature and is the first published example of food chain length being unaffected by any of these determinants. We suggest the distinctive seasonal hydrology of northern Australia allows the movement of fish predators, linking isolated food webs and potentially creating a regional food web that overrides local effects of productivity, disturbance and ecosystem size. This finding supports ecological theory suggesting that mobile consumers promote more stable food webs. It also illustrates how food webs, and energy transfer, may function in the absence of the human modifications to landscape hydrological connectivity that are ubiquitous in more populated regions.

Variability in flood disturbance and the impact on riparian tree recruitment in two contrasting river systems

The vegetation within the riparian zone performs animportant ecological function for in-stream processes.In Australia, riparian zones are regarded as the mostdegraded natural resource zone due to disturbancessuch as river regulation and livestock grazing. Thisstudy looks at factors influencing vegetation dynamicsof riparian tree species on two contrasting riversystems in Western Australia. The Blackwood River insouth-western Australia is influenced by aMediterranean type climate with regular seasonalwinter flows. The Ord River in north-western Australiais characterized by low winter base flows andepisodic, extreme flows influenced by monsoon rains inthe summer. For both rivers, reproductive phenology ofstudied overstory species is timed to coincide withseasonal river hydrology and rainfall. An evendistribution of size classes of trees on the BlackwoodRiver indicated recruitment into the population iscontinual and related to the regular predictableseasonal river flows and rainfall. In contrast, on theOrd River tree size class distribution was clustered,indicating episodic recruitment. On both rivers treeestablishment is also influenced by elevation abovethe river, microtopography, moisture status and soiltype. In terms of vegetation dynamics riparianvegetation on the Ord River consists of long periodsof transition with short lived stable states incontrast to the Blackwood river where tree populationstructure is characterized by long periods of stablestates with short transitions.

Effects of salinisation on riparian plant communities in experimental catchments on the Collie River, Western Australia

Although the salinisation of streams has long been recognised as one of Western Australias most serious environmental and resource problems, there is very little published information on the effects of salinisation on riparian flora and fauna. We studied riparian vegetation in three experimental catchments on the Collie River in Western Australia. The catchments are situated within a 5-km area of state forest and are geologically and botanically similar, but differ in the extent of clearing, groundwater levels and stream salinity. In each catchment, transects were taken perpendicular to the direction of streamflow, and 4-m2 quadrats taken along each transect. Within each quadrat, soil salinity was measured, all plants were identified to species level and percentage cover estimated. The catchments differed significantly in soil salinity, with salinity being greatest in the most extensively cleared catchment and increasing towards the floor of the valley. Plant-species richness, species diversity and species composition were significantly related to soil salinity, both among catchments and among quadrats within the most extensively cleared catchment. Plant-species richness and diversity decreased with increasing soil salinity, an effect that may be partly due to a decline in perennial herb and shrub species. This may have an impact on other components of the riparian ecosystem.

Does flood rhythm drive ecosystem responses in tropical riverscapes

Biotic communities are shaped by adaptations from generations of exposure to selective pressures by recurrent and often infrequent events. In large rivers, floods can act as significant agents of change, causing considerable physical and biotic disturbance while often enhancing productivity and diversity. We show that the relative balance between these seemingly divergent outcomes can be explained by the rhythmicity, or predictability of the timing and magnitude, of flood events. By analyzing biological data for large rivers that span a gradient of rhythmicity in the Neotropics and tropical Australia, we find that systems with rhythmic annual floods have higher-fish species richness, more stable avian populations, and elevated rates of riparian forest production compared with those with arrhythmic flood pulses. Intensification of the hydrological cycle driven by climate change, coupled with reductions in runoff due to water extractions for human use and altered discharge from impoundments, is expected to alter the hydrologic rhythmicity of floodplain rivers with significant consequences for both biodiversity and productivity.

POSTFIRE RESPONSE OF FLOOD‐REGENERATING RIPARIAN VEGETATION IN A SEMI‐ARID LANDSCAPE

Piles of large wood (LW) deposited by major floods in river corridors can interact with naturally occurring wildfires from uplands to impact the regeneration of riparian vegetation. This study examines the spatial and short-term temporal response of riparian vegetation and soil nutrients to fire along the Sabie River, South Africa, with special emphasis on the effects of burned LW piles. At the study site there were 112 species of plants recorded with 28% of species restricted to the burned plots. As expected, vegetation cover was significantly lower in burned plots as compared with the unburned plots 12 months postfire. There was a significant influence of LW on species richness with fewer species recorded in the LW plots. For both fire and LW treatments, plant cover showed a significant change over three years. After an initial increase from 12 to 24 months (postfire) there was a decline in plant cover after 36 months. Species community composition was distinctly different between burned and unburned plots 12 months postfire, and the presence of LW affected species composition for burned plots but not for unburned ones. Time series ordination of LW plots highlighted the changes in species composition over the three years of sampling. Of trees with accumulations of LW within 5 m of their base, 48% had been killed by fire as compared to only 4% with no LW accumulations in close proximity. Soil-available P was significantly higher in the burned plots and even higher with burned LW while there were no effects on soil total N. There was also a significant positive trend between available P in soils and plant vegetation cover. Soil-exchangeable K was also significantly higher and total C significantly lower in the burned and LW plots. Burned plots also had significantly higher soil electrical conductivity (EC) and soil pH. The patchy nature of the studied fire, whose complexity is exacerbated by the distribution of flood deposited LW, acted to create a mosaic of alternate successional states as the riparian community recovers from flooding and the subsequent fire. We suspect that the resultant heterogeneity will increase ecosystem resilience by providing flexibility in the form of more options for a system response to subsequent disturbances.

Seasonal changes in water quality and macrophytes and the impact of cattle on tropical floodplain waterholes

The present study indicates the critical role of hydrologic connectivity in floodplain waterholes in the wet–dry tropics of northern Australia. These waterbodies provide dry-season refugia for plants and animals, are a hotspot of productivity, and are a critical part in the subsistence economy of many remote Aboriginal communities. We examined seasonal changes in water quality and aquatic plant cover of floodplain waterholes, and related changes to variation of waterhole depth and visitation by livestock. The waterholes showed declining water quality through the dry season, which was exacerbated by more frequent cattle usage as conditions became progressively drier, which also increased turbidity and nutrient concentrations. Aquatic macrophyte biomass was highest in the early dry season, and declined as the dry season progressed. Remaining macrophytes were flushed out by the first wet-season flows, although they quickly re-establish later during the wet season. Waterholes of greater depth were more resistant to the effects of cattle disturbance, and seasonal flushing of the waterholes with wet-season flooding homogenised the water quality and increased plant cover of previously disparate waterholes. Therefore, maintaining high levels of connectivity between the river and its floodplain is vital for the persistence of these waterholes.