Body size drives allochthony in food webs of tropical rivers.

Food web subsidies from external sources ("allochthony") can support rich biological diversity and high secondary and tertiary production in aquatic systems, even those with low rates of primary production. However, animals vary in their degree of dependence on these subsidies. We examined dietary sources for aquatic animals restricted to refugial habitats (waterholes) during the dry season in Australia's wet-dry tropics, and show that allochthony is strongly size dependent. While small-bodied fishes and invertebrates derived a large proportion of their diet from autochthonous sources within the waterhole (phytoplankton, periphyton, or macrophytes), larger animals, including predatory fishes and crocodiles, demonstrated allochthony from seasonally inundated floodplains, coastal zones or the surrounding savanna. Autochthony declined roughly 10% for each order of magnitude increase in body size. The largest animals in the food web, estuarine crocodiles (Crocodylus porosus), derived ~80% of their diet from allochthonous sources. Allochthony enables crocodiles and large predatory fish to achieve high biomass, countering empirically derived expectations for negative density vs. body size relationships. These results highlight the strong degree of connectivity that exists between rivers and their floodplains in systems largely unaffected by river regulation or dams and levees, and how large iconic predators could be disproportionately affected by these human activities.

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.

Developing plant functional groups to identify changes in functional composition and diversity in a dryland river experiencing artificially sustained flows.

Land use and climate changes are driving significant shifts in the magnitude and persistence of dryland stream surface flows. The impact of these shifts on ecological functioning is largely unknown, particularly where streams have become wetter rather than drier. This study investigated relationships between hydrologic regime (including surface water persistence, differences in groundwater depth and altered flooding dynamics) with plant traits and riverine vegetation functional composition. Our study system was a previously ephemeral creek in semi-arid northwest Australia that has received groundwater discharge from nearby mining operations for >15 years; surface flows are now persistent for ∼27 km downstream of the discharge point. We aimed to (i) identify plant functional groups (FGs) associated with the creek and adjacent floodplain; and (ii) assess their distribution across hydrological gradients to predict shifts in ecological functioning in response to changing flow regimes. Seven FGs were identified using hierarchical clustering of 40 woody perennial plant species based on morphometric, phenological and physiologic traits. We then investigated how FG abundance (projective foliar cover), functional composition, and functional and taxonomic richness varied along a 14 km gradient from persistent to ephemeral flows, varying groundwater depths, and distances from the stream channel. Dominant FGs were (i) drought avoidant mesic trees that are fluvial stress tolerant, or (ii) drought tolerant xeric tall shrubs that are fluvial stress intolerant. The drought avoidant mesic tree FG was associated with shallow groundwater but exhibited lower cover in riparian areas closer to the discharge (persistent surface flows). However, there were more FGs and higher species richness closer to the discharge point, particularly on the floodplain. Our findings demonstrate that quantifying FG distribution and diversity is a significant step in both assessing the impacts of mine water discharge on riverine ecosystems and for planning for post-mining restoration.

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 Australia's 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.

Fish mediate high food web connectivity in the lower reaches of a tropical floodplain river.

High levels of hydrological connectivity during seasonal flooding provide significant opportunities for movements of fish between rivers and their floodplains, estuaries and the sea, possibly mediating food web subsidies among habitats. To determine the degree of utilisation of food sources from different habitats in a tropical river with a short floodplain inundation duration (~2 months), stable isotope ratios in fishes and their available food were measured from three habitats (inundated floodplain, dry season freshwater, coastal marine) in the lower reaches of the Mitchell River, Queensland (Australia). Floodplain food sources constituted the majority of the diet of large-bodied fishes (barramundi Lates calcarifer, catfish Neoarius graeffei) captured on the floodplain in the wet season and for gonadal tissues of a common herbivorous fish (gizzard shad Nematalosa come), the latter suggesting that critical reproductive phases are fuelled by floodplain production. Floodplain food sources also subsidised barramundi from the recreational fishery in adjacent coastal and estuarine areas, and the broader fish community from a freshwater lagoon. These findings highlight the importance of the floodplain in supporting the production of large fishes in spite of the episodic nature and relatively short duration of inundation compared to large river floodplains of humid tropical regions. They also illustrate the high degree of food web connectivity mediated by mobile fish in this system in the absence of human modification, and point to the potential consequences of water resource development that may reduce or eliminate hydrological connectivity between the river and its floodplain.

Riparian condition influences spider community structure and the contribution of aquatic carbon subsidies to terrestrial habitats.

Degradation of riparian zones can alter aquatic and terrestrial communities of flora and fauna and disrupt their role in assimilating and mobilising carbon between the two ecosystems. Riparian spiders that predate on emergent aquatic invertebrates can contribute to carbon flux and the structure of aquatic and riparian food webs. The impact of riparian degradation on spiders in temperate rivers of Australia and their role in this broader ecosystem function is poorly understood. We surveyed the riparian zone of four rivers of south-western Australia in areas of natural intact vegetation and degraded agricultural land to explore whether riparian spider abundance, and diversity may be affected by changes to riparian condition. We also assessed the impact of the riparian condition on carbon fluxes between aquatic and terrestrial environments, using stable isotope analysis. We found overall abundance of riparian spiders was higher in degraded agricultural sites compared to natural intact sites and the structure of spider assemblages was different. Orb-weaver spiders (Araneidae and Tetragnathidae) were found to be more abundant in agricultural areas where canopy cover and understory are sparse as a result of livestock grazing and trampling. The contribution of carbon from aquatic invertebrates in a natural intact site was 48.5% for Orb-weavers and 41.6% for Cursorial Hunter spiders but reduced to 19.6% and 39.9% respectively in a degraded agricultural site. These results suggest that the position of spiders in riparian food webs and the amount of aquatic subsidy may change according to the condition and complexity of the riparian zone.

Multi-scale characterisation of stream nutrient and carbon dynamics in sandy near coastal catchments of south-western Australia.

Managers tasked with repairing degraded stream ecosystems require restoration strategies that are tailored to local and regional characteristics. Emerging evidence suggests that local reach-scale approaches may be as effective, if not more so, than catchment-scale actions in highly permeable coastal landscapes, particularly if there is hydraulic connectivity to shallow groundwater and where recharge is strongly seasonal. This study assessed the relative influence of catchment-scale land use and reach-scale vegetation structure on the distribution of carbon and nutrient concentrations of streams within urban and agricultural catchments of the Perth region of south-western Australia. We used linear mixed-effects models to evaluate the extent to which phosphorus, nitrogen and carbon concentrations in different stream zones (streamwater, and fluvial and parafluvial sediments) were explained by catchment and reach-scale attributes and moderated by high versus low-flow periods, i.e., in wet versus dry months. We found that reach-scale vegetation (woody plant cover, annual plant cover) was a better predictor of nutrient concentrations than catchment-scale land use, particularly total imperviousness, a common measure of urbanisation. Flow was also important, with carbon and nutrient concentrations better described by reach- or catchment-scale attributes during the low flow period. The extent to which individual catchment and reach attributes influenced the distribution of nutrients in different stream zones was complex. However, our results suggest that planting woody vegetation can reduce nitrogen concentrations in surface water and fluvial sediments. Reducing the abundance of weedy annual species and restoring woody perennial species may further reduce phosphorus concentrations in surface water. We conclude that local riparian restoration can be a cost-effective strategy for managing excess nutrients and carbon in flat and permeable urban landscapes, particularly during low flow periods.

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.

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.

Flood-deposited wood debris and its contribution to heterogeneity and regeneration in a semi-arid riparian landscape.

We investigated whether large woody debris (LWD) piles create nodes of environmental resources that contribute to the recovery of riparian vegetation and that also augment the heterogeneity and resilience of the riverine system. River and riparian systems are typified by a large degree of heterogeneity and complex interactions between abiotic and biotic elements. Disturbance such as floods re-distribute the resources, such as LWD, and thereby add greater complexity to the system. We examined this issue on a semi-arid savanna river where approximately a 100-year return interval flood in 2000 uprooted vegetation and deposited substantial LWD. We investigated the micro-environment within the newly established LWD piles and compared this with conditions at adjacent reference sites containing no LWD. We found soil nutrient concentrations to be significantly higher in LWD piles compared with the reference plots (total N +19%, available P +51%, and total C +36%). Environmental variables within LWD piles and reference sites varied with landscape position in the river-riparian landscape and with LWD pile characteristics. Observed differences were generally between piles located in the terrestrial and riparian areas as compared to piles located on the macro-channel floor. After 3 years the number and cover of woody species were significantly higher when associated with LWD piles, regardless of landscape position or pile type. We conclude that LWD piles formed after large floods act as resource nodes by accumulating fine sediments and by retaining soil nutrients and soil moisture. The subsequent influence of LWD deposition on riparian heterogeneity is discerned at several spatial scales including within and between LWD piles, across landscape positions and between channel types. LWD piles substantially influence the initial developmental of riparian vegetation as the system regenerates following large destructive floods.