Characteristics of the ichthyofaunas of southwestern Australian estuaries,including comparisons with holarctic estuaries and estuaries elsewhere in temperate Australia: A review

Data on the species compositions and the ages, sizes, reproductive biology, habitats and diets of the main species in the ichthyofaunas of seven estuaries in temperate southwestern Australia have been collated. Twenty-two species spawn in these estuaries, of which 21 complete their lifecycles in the estuary. The latter group, which includes several species of atherinids and gobies with short lifecycles, make far greater contributions to the total numbers of fish in the shallows of these estuaries than in those of holarctic estuaries, such as the Severn Estuary in the United Kingdom. This is presumably related in part to far less extreme tidal water movements and the maintenance of relatively high salinities during the dry summers, and thus to more favourable conditions for spawning and larval development. However, since estuaries in southwestern Australia have tended to become closed for periods, there would presumably also have been selection pressures in favour of any members of marine species that were able to spawn in an estuary when that estuary became landlocked. Furthermore, the deep saline waters, under the marked haloclines that form in certain regions during heavy freshwater discharge in winter, act as refugia for certain estuarine species. The contributions of estuarine-spawning species to total fish numbers in the shallows varied markedly from 33 or 34% in two permanently open estuaries to ≥ 95% in an intermittently open estuary, a seasonally closed estuary and a permanently open estuary on the south coast, in which recruitment of the 0 + age class of marine species was poor. The larger estuarine species can live for several years and reach total lengths of ~ 700 mm and some estuarine species move out into deeper waters as they increase in size. Several marine species use southwestern Australian estuaries as nursery areas for protracted periods. However, sudden, marked increases in freshwater discharge in winter and resultant precipitous declines in salinity in the shallows, and in other regions where haloclines are not formed, are frequently accompanied by rapid and pronounced changes in ichthyofaunal composition, partly due to the emigration of certain marine species. In contrast, the ichthyofaunal compositions of macrotidal holarctic estuaries undergo annual, cyclical changes, due largely to the sequential entry of the juveniles of different marine species for short periods. The ichthyofaunal compositions of the narrow entrance channels, wide basins and saline riverine reaches of large, permanently open southwestern Australian estuaries vary, reflecting the marked tendency for some species to be restricted mainly to one or two of these regions. Comparative data indicate that the characteristics determined for ichthyofaunas in southwestern Australian estuaries apply in general to estuaries elsewhere in temperate Australia.     

Estuarine benthic ecology: A European perspective

The study of benthic ecology within European estuaries is discussed. The history of estuarine science, and the definition of an estuary is briefly reviewed. Estuarine benthic ecology is considered with regard to its geographical, economic and political context. It is shown that much recent study has been motivated by attempts to clean up estuaries, which were polluted after decades, or even centuries of pollution. Much has been achieved in the control of pollution, and many estuaries show clear signs of recovery. Particular focus has been on the estuaries entering the North Sea, with the realization that it is not the North Sea that is polluted, but its estuaries. The scientific appraisal of the benthos of estuaries has been particularly assisted by the ecosystem approach, which has both aided our understanding of their function as well as answering some of the questions posed by the impact of mankind. The European Union is identified as a powerful factor in benthic ecology, either through legislation or through the encouragement of collaborative programs, some of which have been particularly important in identifying common factors across a wide geographical spread of estuaries. It is concluded that in general estuaries have got cleaner, but that many problems remain, ranging from eutrophication through to sea-level change. The future of estuarine benthic ecology lies in meeting the new challenges.     

Weak and Habitat-Dependent Effects of Nutrient Pollution on Macrofaunal Communities of Southeast Australian Estuaries

Among the impacts of coastal settlements to estuaries, nutrient pollution is often singled out as a leading cause of modification to the ecological communities of soft sediments. Through sampling of 48 sites, distributed among 16 estuaries of New South Wales, Australia, we tested the hypotheses that (1) anthropogenic nutrient loads would be a better predictor of macrofaunal communities than estuarine geomorphology or local sediment characteristics; and (2) local environmental context, as determined largely by sediment characteristics, would modify the relationship between nutrient loading and community composition. Contrary to the hypothesis, multivariate multiple regression analyses revealed that sediment grain size was the best predictor of macrofaunal assemblage composition. When samples were stratified according to median grain size, relationships between faunal communities and nitrogen loading and latitude emerged, but only among estuaries with sandier sediments. In these estuaries, capitellid and nereid polychaetes and chironomid larvae were the taxa that showed the strongest correlations with nutrient loading. Overall, this study failed to provide evidence of a differential relationship between diffuse nutrient enrichment and benthic macrofauna across a gradient of 7° of latitude and 4°C temperature. Nevertheless, as human population growth continues to place increasing pressure on southeast Australian estuaries, manipulative field studies examining when and where nutrient loading will lead to significant changes in estuarine community structure are needed.     

Taxonomy of estuarine invertebrates in Australia

An Australian estuary is primarily a marine-dominated environment, subjected to major salinity changes only after heavy rains and during flood conditions. In southeastern Australia estuarine biota are similar to those in shallow coastal bays and some of the coastal lagoons, and most taxonomic studies on estuarine fauna have occurred in this region. Several major surveys of estuarine and coastal bays have been undertaken in Australia during the past three decades and these surveys have largely provided the material for taxonomic studies of the major groups of macrobenthos. All these studies have revealed a diverse and abundant benthic macrobenthos. The composition of estuarine invertebrate fauna is compared with open exposed coasts, including fauna found in soft strata and on hard strata. Australia has a limited specialized estuarine biota much of which is restricted to particular habitats, and a highly diverse marine community which thrives in the sheltered protected waters of estuaries and the associated soft sediments. Currently, interactive computer-based keys are being developed which should greatly assist the student and the benthic ecologist to identify the fauna and prevent potential loss of data. Some aspects of the current debate of the level of taxonomic resolution required to answer various ecological questions are discussed. Finally, some comments are suggested as to future directions in which taxonomists and estuarine ecologists should proceed in order to be able to detect changes or losses of estuarine biodiversity and the loss of the associated biological information which may be critical in understanding the functioning of the estuarine ecosystem.     

Testate amoebae analysis in ecological and paleoecological studies of wetlands: past, present and future

Testate amoebae are an abundant and diverse polyphyletic group of shelled protozoa living in aquatic to moist habitats ranging from estuaries to lakes, rivers, wetlands, soils, litter, and moss habitats. Owing to the preservation of shells in sediments, testate amoebae are useful proxy indicators complementary to long-established indicators such as pollen and spores or macrofossils. Their primary use to date has been for inferring past moisture conditions and climate in ombrotrophic peatlands and, to a lesser extent, to infer pH in peatlands and the trophic or nutrient status of lakes. Recent research on these organisms suggests other possible uses in paleoecology and ecology such as sea-level reconstruction in estuarine environments, as indicators of soil or air pollution, and monitoring recovery of peatland. We review the past and present use of testate amoebae, the challenges in current research, and provide some ideas on future research directions.     

Eutrophication in Australian Rivers, Reservoirs and Estuaries – A Southern Hemisphere Perspective on the Science and its Implications

Australian science has made rapid advances in the last decade in understanding eutrophication processes in inland waters and estuaries. The freshwater research on which these advances are based was triggered by well-publicised blooms of cyanobacteria during the 1980s and early 1990s, particularly a 1000 km long bloom on the Darling River. In estuaries the study which greatly enhanced our understanding but simultaneously served to stimulate further research into estuarine eutrophication, the Port Phillip Bay Study, was initially designed to address perceived problems of toxicants in the Bay but provided profound insights into drivers for, and ecosystem responses to, eutrophication. Subsequent estuarine research has largely been stimulated by management questions arising from Australia’s increasing coastal development for residential purposes. The research has shown that some of the beliefs extant at the time of the blooms were incorrect. For example, it is now clear that stratification and light penetration, not nutrient availability, are the triggers for blooms in the impounded rivers of southeastern Australia, although nutrient exhaustion limits the biomass of blooms. Again, nitrogen seems to play as important a role as phosphorus does in controlling the biomass of these freshwater blooms. The research has also shown that aspects of eutrophication, such as nutrient transport, are dominated by different processes in different parts of Australia. Many of the biophysical processes involved in eutrophication have now been quantified sufficiently for models to be developed of such processes as sediment-nutrient release, stratification, turbidity and algal growth in both freshwater and estuarine systems. In some cases the models are reliable enough for the knowledge gained in particular waterbodies to be applied elsewhere. Thus, there is now a firm scientific foundation for managers to rely upon when managing algal blooms. Whilst these findings have already been presented to managers and communities throughout Australia, there is still a considerable way to go before they are absorbed into their modus operandi.     

A comparative ecology of estuaries in Nigeria

A bar-built estuary, a drowned river valley and two river delta estuaries were compared and contrasted to elucidate the impact of some abiotic factors, notably climate, salinity, and oxygen on the distribution of the aquatic fauna and flora.Salinity was recognizable as the key factor responsible for the population dynamics in these habitats. The mangrove community is characteristically zoned, but development and distribution of the trees is restricted in view of coastal geomorphic and hydrological processes to the western banks of the estuaries.The preponderance of marine fish species in estuaries was confirmed, although the wide salinity tolerance of some prolific-breeding Cichlidae from freshwater was noted. Forty-eight species of bony fishes were recorded in Lagos Lagoon, the Escravos, and Qua Ibo estuaries and classified by salinity tolerance into stenohaline and euryhaline marine species (66%), and freshwater species (34%). The same estuarine ichthyofauna were split into trophic groups: piscivores (46%), zooplankton feeders (10%), meiobenthos feeders (27%), and macrobenthos feeders (17%).     

Opportunistic use of estuarine habitat by juvenile bull trout, <Emphasis Type="Italic">Salvelinus confluentus</Emphasis>, from the Elwha River before,during, and after dam removal

Estuaries are used by anadromous fishes, either as the definitive marine habitat or as transition habitat as they move to fully marine waters, and extent of estuary use may vary with habitat conditions and fish attributes. Bull trout (Salvelinus confluentus) are commonly fluvial or adfluvial, though anadromous populations also exist. However, little is known about estuary use, especially by juveniles of this threatened species. We sampled the estuaries of the Elwha River, where a spawning population exists, and the nearby Salt Creek, where none exists, to reveal seasonal timing of estuarine use by juvenile bull trout, size of those using the estuary, and possible use of the non-natal estuary. We captured juvenile bull trout (all ≥100 mm FL, most <300 mm) in the Elwha River estuary in all months except August, but primarily December through May. None was captured in Salt Creek’s estuary despite comparable sampling effort. We also evaluated how dam removal on the Elwha River influenced bull trout estuarine occupancy by sampling before, during, and after dam removal, because this process enlarged the estuary but also increased turbidity and sediment transport in the lower river. Catches were low before dam removal, increased during and immediately after removal, and returned to low levels in recent years, suggesting that juveniles temporarily sought refuge from conditions associated with dam removal. Our findings indicate juvenile bull trout occupy estuarine habitat opportunistically; this information may aid conservation efforts as anadromous populations occur elsewhere in rivers with estuaries altered by human development.     

Shock,Stress or Signal? Implications of Freshwater Flows for a Top-Level Estuarine Predator

Physicochemical variability in estuarine systems plays an important role in estuarine processes and in the lifecycles of estuarine organisms. In particular, seasonality of freshwater inflow to estuaries may be important in various aspects of fish lifecycles. This study aimed to further understand these relationships by studying the movements of a top-level estuarine predator in response to physicochemical variability in a large, temperate south-east Australian estuary (Shoalhaven River). Mulloway (Argyrosomus japonicus, 47–89 cm total length) were surgically implanted with acoustic transmitters, and their movements and migrations monitored over two years via fixed-position VR2W acoustic receivers configured in a linear array along the length of the estuary. The study period included a high degree of abiotic variability, with multiple pulses (exponentially high flows over a short period of time) in fresh water to the estuary, as well as broader seasonal variation in flow, temperature and conductivity. The relative deviation of fish from their modal location in the estuary was affected primarily by changes in conductivity, and smaller fish (n = 4) tended to deviate much further downstream from their modal position in the estuary than larger fish (n = 8). High-flow events which coincided with warmer temperatures tended to drive mature fish down the estuary and potentially provided a spawning signal to stimulate aggregation of adults near the estuary mouth; however, this relationship requires further investigation. These findings indicate that pulse and press effects of freshwater inflow and associated physicochemical variability play a role in the movements of mulloway, and that seasonality of large freshwater flows may be important in spawning. The possible implications of river regulation and the extraction of freshwater for consumptive uses on estuarine fishes are discussed.     

Seasonality in macroinvertebrate community composition across a neglected ecological boundary, the freshwater-estuarine transition zone

The boundary between freshwater and estuarine ecosystems at the extreme head of estuaries has received little attention from benthic ecologists. This short communication addresses this research caveat, presenting preliminary data on the spatial and temporal (seasonal) variation in the composition of benthic macroinvertebrate communities across the transition zone at the head of a small British estuary. Cluster analysis and multidimensional scaling ordination demonstrated that a distinct community occurred at a point just preceding the recognised transition between fresh and brackish waters (salinity <0.2PSU). This transition community contained a mixture of freshwater and estuarine taxa and, despite seasonal migrations at the head of the estuary, community structure remained significantly distinct from that at adjacent sites throughout the year. These preliminary data suggest that the community dynamics across the transition zone show complex patterns and may offer an important research opportunity in aquatic ecology.     

Transport of molluscan larvae through a shallow estuary

Dispersal of invertebrate larvae is determined by larval swimmingbehavior, the length of planktonic development and the hydrodynamicregime. Larvae of estuarine invertebrates must refrain fromexport or invade an estuary after development in the ocean.This study investigates retention patterns of estuarine molluscsby measuring time series of larval abundance in relation tohydrodynamic processes. Previous investigations of larval dynamicshave generally focused on larger estuarine systems that areoften stratified and have relatively long hydraulic residencetimes. The estuary studied in this investigation supports densepopulations of infaunal clams yet has a water depth to tidalamplitude ratio near unity. To access processes affecting larvalretention, the circulation patterns of the estuary were measuredwith time series of salinity, temperature, pressure and horizontalvelocity. Transport rates of larvae between ocean and estuary,and within the estuary proper, were calculated from velocityand larval concentration time series. The daily residence timeof the estuary was determined for the summer spawning period.The results demonstrate that molluscan larvae were routinelytransported between the estuary and nearshore zone in tidalflows. Based on the magnitude of the horizontal current velocities,passive transport of larvae predominates during most of thetidal cycle in the estuary. Residence time calculations suggestthat the ability of larvae to remain in the estuary throughlarval development is unlikely, and there was no evidence ofselective retention of mature bivalve larvae in the estuary.Rather, larvae are exported rapidly from the estuary and undergodevelopment in the coastal ocean. Mesoscale physical processesin the coastal ocean probably control variation in the deliveryof larvae back to estuarine systems. Recruitment to this andsimilar estuaries must therefore be dependent on invasion.     

Geographical and typological changes in fish guilds of South African estuaries

A comparative analysis of fish estuary association guilds was undertaken on some 190 South African estuaries. This pioneering study spanned three zoogeographic regions and included three broad estuarine types. The guild compositions of the estuaries were compared based on an importance value, incorporating taxonomic composition, numerical abundance and relative biomass. Multivariate analyses included both inter‐regional (zoogeographic) and intra‐regional (estuarine typology) comparisons. The major estuary‐associated guilds (estuarine species and marine migrant species) were important in all estuary types within all biogeographic regions. Significant differences both between regions and between estuary types within regions, however, were recorded. Cool–temperate estuaries were generally dominated by migratory species (estuarine migrants and marine migrant opportunists) while the importance of species dependent on estuaries (estuarine residents and estuarine‐dependent marine migrants) was higher in warm–temperate and subtropical regions. The significance of estuarine nursery areas, particularly in regions where estuaries are few in number, is highlighted. In terms of typology, migratory species assumed a greater importance in predominantly open systems, while freshwater and estuarine‐resident species were more important in predominantly closed systems. Predominantly closed estuaries, however, were also important for marine migrant species, which further highlights the significance of these systems as nursery areas for fishes.     

Ecosystem services transcend boundaries: estuaries provide resource subsidies and influence functional diversity in coastal benthic communities

Background

Estuaries are highly productive ecosystems that can export organic matter to coastal seas (the ‘outwelling hypothesis’). However the role of this food resource subsidy on coastal ecosystem functioning has not been examined.

Methodology/Principal Findings

We investigated the influence of estuarine primary production as a resource subsidy and the influence of estuaries on biodiversity and ecosystem functioning in coastal mollusk-dominated sediment communities. Stable isotope values (δ¹³C, δ¹⁵N) demonstrated that estuarine primary production was exported to the adjacent coast and contributed to secondary production up to 4 km from the estuary mouth. Further, isotope signatures of suspension feeding bivalves on the adjacent coast (Dosinia subrosea) closely mirrored the isotope values of the dominant bivalves inside the estuaries (Austrovenus stutchburyi), indicating utilization of similar organic matter sources. However, the food subsidies varied between estuaries; with estuarine suspended particulate organic matter (SPOM) dominant at Tairua estuary, while seagrass and fringing vegetation detritus was proportionately more important at Whangapoua estuary, with lesser contributions of estuarine SPOM. Distance from the estuary mouth and the size and density of large bivalves (Dosinia spp.) had a significant influence on the composition of biological traits in the coastal macrobenthic communities, signaling the potential influence of these spatial subsidies on ecosystem functioning.

Conclusions/Significance

Our study demonstrated that the locations where ecosystem services like productivity are generated are not necessarily where the services are utilized. Further, we identified indirect positive effects of the nutrient subsidies on biodiversity (the estuarine subsidies influenced the bivalves, which in turn affected the diversity and functional trait composition of the coastal sediment macrofaunal communities). These findings highlight the importance of integrative ecosystem-based management that maintains the connectivity of estuarine and coastal ecosystems.     

Role of meiofauna in estuarine soft-bottom habitats*

Meiofauna are ubiquitous in estuaries worldwide averaging 10⁶ m^?2. Abundance and species composition are controlled primarily by three physical factors: sediment particle size, temperature and salinity. While meiofauna are integral parts of estuarine food webs, the evidence that they are biologically controlled is equivocal. Top down (predation) control clearly does not regulate meiofaunal assemblages. Meiofauna reproduce so rapidly and are so abundant that predators cannot significantly reduce population size. Food quantity (bottom up control) also does not appear to limit meiofaunal abundance; there is little data on the effect of food quality. In estuarine sediments meiofauna: (i) facilitate biomineralization of organic matter and enhance nutrient regeneration; (ii) serve as food for a variety of higher trophic levels; and (iii) exhibit high sensitivity to anthropogenic inputs, making them excellent sentinels of estuarine pollution. Generally mineralization of organic matter is enhanced and bacterial production stimulated in the presence of meiofauna. Tannins from mangrove detritus in northern Queensland appear to inhibit meiofaunal abundance and therefore the role of meiofauna in breakdown of the leaves. Meiofauna, particularly copepods, are known foods for a variety of predators especially juvenile fish and the meiofaunal copepods are high in the essential fatty acids required by fish. The copepod’s fatty acid composition is like that of the microphytobenthos they eat; bacterial eaters (nematodes?) do not have the essential fatty acids necessary for fish. Most contaminants in estuaries reside in sediments, and meiofauna are intimately associated with sediments over their entire life-cycle, thus they are increasingly being used as pollution sentinels. Australian estuarine meiofauna research has been concentrated in Queensland, the Hunter River estuarine system in New South Wales, and Victoria’s coastal lagoons. Studies in northern Queensland have primarily concentrated on the role of nematodes in mineralization of organic matter, whereas those from southern Queensland have concentrated on the role of meiofauna as food for fish and as bacterial grazers. The New South Wales studies have concentrated on the Hunter River estuary and on the structure and function of marine nematode communities. In Victoria, several fish have been shown to eat meiofauna. The Australian world of meiofaunal research has hardly been touched; there are innumerable opportunities for meiofaunal studies. In contaminated estuarine sediments reduced trophic coupling between meiofauna and juvenile fish is a basic ecological question of habitat suitability, but also a question with relevance to management of estuarine resources. Because meiofauna have short lifecycles, the effects of a contaminant on the entire life-history can be assessed within a relatively short time. The use of modern molecular biology techniques to assess genetic diversity of meiofauna in contaminated vs uncontaminated sediments is a promising avenue for future research. Much of the important meiofaunal functions take place in very muddy substrata; thus, it is imperative to retain mudflats in estuaries.     

The influence of freshwater flows on two estuarine resident fish species show differential sensitivity to the impacts of drought,flood and climate change

Estuaries are particularly susceptible to climate change and drought resulting in atypical changes to freshwater flows. How such changes in flow impact on the ecology of estuarine fishes may depend on how a species moves in response to changing flow conditions. Acoustic telemetry was used to interpret fine-scale movements of two co-inhabiting estuarine fish species, black bream, Acanthopagrus butcheri and estuary perch, Macquaria colonorum in relation to freshwater flows, season and moon phase. We found black bream to be highly mobile, regularly travelling the length of the estuary and into the neighbouring estuaries. In contrast, estuary perch had particular home ranges and made occasional, upstream or downstream movements. Possibly influenced by freshwater flows, estuary perch moved at greater rates in the Tambo compared to fish in the Mitchell. Black bream resided in the upper estuary during winter and spring and the lower estuary during summer and autumn, whereas estuary perch remained in the upper estuary throughout the year, with occasional downstream movements in winter and spring. This study revealed 1) significantly large increases in freshwater flows result in mass downstream movements in both species, 2) fish moved upstream during full moons and 3) there are contrasting spatio-temporal patterns in movement between species. The results from this study highlight that estuarine fishes are likely to show differential sensitivity to the impacts of drought and climate change and illustrate how acoustic telemetry methods can be used to determine the environmental needs of fishes and help efforts to conserve and manage estuaries worldwide.     

δ15N of estuarine fishes as a quantitative indicator of urbanization

Nitrogen stable isotope values (δ¹⁵N) have commonly been used as a qualitative indicator of catchment urbanization in estuaries, but no quantitative relationship has to date been established between δ¹⁵N and degree of urbanization. We sampled five species of common estuarine fish (Mugil cephalus, Acanthopagrus australis, Sillago ciliata, Gerres subfasciatus and Herklotsichthys castelnaui) of different trophic levels from six estuaries in Southeast Queensland, Australia, to test if a quantitative relationship exists between fish δ¹⁵N and urbanization in the catchment. Degree of urbanization (urban %) in the catchment adjacent to the tidal estuary was measured using polygons constructed in Google Earth images, and verified using data from the detailed Queensland Land Use Mapping Project (QLUMP). Significant linear relationships exist between fish δ¹⁵N and urban % in most cases irrespective of the section of the estuary (upper, middle and lower) where the fish were caught, with no difference in slope and elevation between lines established for different sections. Inclusion of additional fish δ¹⁵N data from other Australian estuaries in a combined multiple regression model adding trophic level as a predictor generated a significant relationship predicting increase in fish δ¹⁵N values with increasing urban % and trophic level. There is therefore a generic quantitative relationship between estuarine fish δ¹⁵N values and degree of urbanization applicable to different fish species, geographic location or feeding habit. While the generality of this relationship needs to be tested further in other geographic locations, our findings potentially improve the applicability of fish δ¹⁵N values as a quantitative indicator of urban influence in estuaries.     

Pieter Hendrik Nienhuis: Aquatic Ecologist and Environmental Scientist

Prof. Dr. Pieter Hendrik (Piet) Nienhuis worked for almost 40 years in all aspects of aquatic ecology and environmental science and retired on 31 October 2003. He can be characterised as a distinguished scientist, shaped in an applied estuarine and aquatic research ambience of the former Delta Institute for Hydrobiological Research (DIHO) in Yerseke in the Netherlands. His appointment as a full professor at the Radboud University Nijmegen offered him a challenging step from monodisciplinarity in ecology, via multidisciplinarity in the application of ecological knowledge in river science to interdisciplinarity in environmental science and management. This paper describes his education, teaching activities, research, scientific publications, science management, and significance for various scientific disciplines. He made important contributions to biosystematics of angiosperms and algae, the ecology of seagrasses, nutrient cycling and eutrophication in estuarine ecosystems, and the integrated modelling of the ecological functioning of estuaries. Subsequently, he paid much attention to environmental problems in river basins, ecological rehabilitation and sustainable development. His work influenced the view of ecologists, aquatic scientists and water managers in the Netherlands as well as abroad, in particular regarding the drawbacks of compartmentalization of the estuaries and the importance of connectivity and morphodynamics in river systems. In hindsight, it appears as a logical line that he gradually moved from estuarine ecological research that became increasingly driven by societal and environmental problems to the field of environmental science and management.