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.     

A comparison of the macrobenthic faunas of permanently open and temporarily open/closed South African estuaries

Thirteen estuaries in the Eastern Cape Province, South Africa, were broadly categorised according to size and salinity distribution and were assigned to one of the following categories: permanently open estuaries having a strong salinity gradient between mouth and upper estuary, freshwater-deprived permanently open estuaries, medium-sized temporarily open/closed estuaries, and small, temporarily open/closed estuaries. The macrobenthos collected during surveys was then compared in terms of the following parameters: species composition, salinity, sediment mud content, density of macrobenthic animals, Hill's N0 (species richness), and Hill's N1 (diversity). Mud content was found to be the most important environmental variable responsible for biotic patterns found, and sites were consequently assigned to either a sand zone fauna, or a mud zone fauna. Both types of fauna are present in all estuaries sampled, with upper sites of river dominated estuaries having an additional oligohaline fauna, and freshwater-deprived estuaries providing habitat for many marine species. Small, temporarily open/closed estuaries have the highest macrobenthic density, whereas N0 and N1 are highest in freshwater-deprived permanently open systems. River-dominated permanently open estuaries tend to have lower macrobenthic densities, species richness, and diversities compared to estuaries in the other categories. No seasonal differences in these ecological indices were found within any of the estuarine categories.     

Estuarine Typology and the Structuring of Fish Communities in South Africa

Synopsis Some 190 South African estuaries, covering all biogeographic provinces within the region, were classified into three types based on a combination of mouth condition and estuary size (surface area). The fish communities of the estuary types within each zoogeographic region were described and compared. Multivariate analyses revealed that each estuary type contained somewhat distinct fish communities. In addition, the study identified common patterns in species richness and ichthyofaunal composition. Open estuaries have relatively high species richness; this is a reflection of a permanent or near-permanent connection with the sea which allows access into these estuaries by all marine migrant species within the region. Intermittent connection with the sea limits the recruitment and utilisation of closed estuaries by marine migrant species; this results in reduced species richness in moderate to large closed estuaries. Small closed estuaries exhibit the lowest species richness and this is probably a result of their limited habitat and increased isolation from the sea. The key fishes that utilise estuaries could also be categorised into a number of groups based on their relative importance within each estuary type. Some species are largely restricted to predominantly open systems. Other taxa, while important in predominantly open estuaries, also occur in moderate to large closed systems. Some estuarine-associated species are well represented in all estuary types but exhibit a greater importance in closed estuaries. This study has shown that South African fish communities not only reflect estuarine typology but also respond to these differences in a consistent manner that spans all zoogeographic regions. The prevalence of similar patterns in other parts of the world suggests that estuarine typology is a major driver in the structuring of global estuarine fish communities.     

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.     

Growth,condition, and maturity schedules of an estuarine fish species change in estuaries following increased hypoxia due to climate change

Understanding challenges posed by climate change to estuaries and their faunas remains a high priority for managing these systems and their communities. Freshwater discharge into a range of estuary types in south‐western Australia between 1990 and 2015 is shown to be related to rainfall. This largely accounts for decreases in discharge in this microtidal region being more pronounced on the west coast than south coast, where rainfall decline was less. Results of an oxygen‐balance model imply that, as demonstrated by empirical data for the Swan River Estuary, declines in discharge into a range of estuary types would be accompanied by increases in the extent of hypoxia. In 2013–15, growth and body condition of the teleost Acanthopagrus butcheri varied markedly among three permanently open, one intermittently‐open, one seasonally‐closed and one normally‐closed estuary, with average time taken by females to reach the minimum legal length (MLL) of 250 mm ranging from 3.6 to 17.7 years. It is proposed that, in a given restricted period, these inter‐estuary variations in biological characteristics are related more to differences in factors, such as food resources and density, than to temperature and salinity. The biological characteristics of A. butcheri in the four estuaries, for which there are historical data, changed markedly between 1993–96 and 2013–15. Growth of both sexes, and also body condition in all but the normally‐closed estuary, declined, with females taking between 1.7 and 2.9 times longer to attain the MLL. Irrespective of period, body condition, and growth are positively related. Age at maturity typically increased between periods, but length at maturity declined only in the estuary in which growth was greatest. The plasticity of the biological characteristics of A. butcheri, allied with confinement to its natal estuary and ability to tolerate a wide range of environmental conditions, makes this sparid and comparable species excellent subjects for assessing estuarine “health.”     

The ichthyoplankton of a seasonally closed estuary in temperate Australia. Does an extended period of opening influence species composition?

Fish eggs and larvae were collected monthly between September 1987 and April 1989 from sites throughout the main basin and within the saline regions of the two main tributary rivers of Wilson Inlet, a seasonally closed estuary in south-western Australia. Of the eggs, 43.7% belonged to Engraulis australis (Shaw) and the rest to unidentified teleosts. The larval fish assemblage comprised 17 families represented by 25 species. The Gobiidae contained the highest number of species (four) and contributed approximately 57% of all larvae caught. Pseudogobius olorum (Sauvage) and E. australis were the most abundant species, contributing 43.9 and 27.9% to the total larval catch, respectively. The larvae of species which breed within Wilson Inlet dominated the assemblage, both in terms of number of species (64%) and contribution to total catch (99.9%). The numbers of the eight marine species and one freshwater species represented in the ichthyoplankton were very low. Classification and multi-dimensional scaling ordination showed that the composition of the larval fish fauna at the various sites during a period when the estuary remained open to the sea (December 1988-April 1989) was similar to that of the corresponding sites during the same period in the previous year when the estuary had become closed (December 1987-April 1988). This can be attributed to the spatial distribution, time of occurrence and abundance of estuarine-spawned larvae being similar in the two periods and to the rarity of marine-spawned larvae, even in the spring and summer of 1988/1989 when the estuary was open for the whole time when most marine teleosts spawn in south-western Australia. The low occurrence of marine-spawned larvae in Wilson Inlet reflects the fact that tidal water movement within the basin of the system is so small that it is unable to facilitate the transport and dispersion of larvae. The ichthyoplankton of Wilson Inlet resembles that of other poorly-flushed estuaries in that it is low in species richness and dominated by estuarine-spawned larvae.     

Benthic macrofaunal communities in intermittent estuaries during a drought: Comparisons with permanently open estuaries

Drought conditions have prevailed in many areas of NSW since 2002. On the mid-north coast, below-average rainfall resulted in reduced riverine flows and the extended closure of intermittent estuaries within the Solitary Islands Marine Park. Patterns of structure of benthic infaunal communities were evaluated at the height of the drought to determine if they differed between closed, intermittent estuaries and permanently open estuaries within the region. Replicate van Veen grab samples were taken in the upper, mid- and lower reaches of six intermittent and three permanently open estuaries and sieved to retain the macrofauna. A range of physico-chemical measures was also taken at each sampling time. Multivariate analyses of assemblage data revealed a significant difference between the structure of the two estuary types and also among estuaries within each type. Differences between estuary types were attributable to small differences in the abundance of a number of taxa but also to the absence of the amphipod Urohaustorius metungi from most of the intermittent estuaries. In contrast, these small amphipods dominated communities in the lower reaches of the permanently open estuaries. Physico-chemical variables were highly variable among estuaries and were not strongly correlated with assemblage patterns. Correlations with catchment size were the strongest and, as most of the intermittent estuaries in the region are smaller than the permanently open estuaries, this confounds the interpretation of assemblage patterns in this preliminary study. In order to differentiate between the effects of catchment size and entrance status, the same estuaries need to be resurveyed during periods when at least some of the intermittent estuaries are open.     

Are nest sites a limiting resource for speleophilic nesting fishes in sand dominated temporarily open/closed estuaries?

Southern African temporarily open/closed estuaries are often characterized by sand dominated mouth regions, a feature in contrast to many permanently open estuaries of the region which regularly have substantial rocky areas. Here we explore how habitat characteristics may reduce nesting opportunities and potentially explain population trends for a common estuarine fish species, the speckled sand goby Psammogobius knysnaensis, in many temporarily open/closed estuaries in South Africa. The present study is the first to document that P. knysnaensis is a speleophilic nester, utilizing the underside of rocks and other hard substrata for egg attachment. Furthermore, our results suggest that ideal nesting habitat for the speckled sand goby is potentially a limiting resource in sand-dominated temporarily open/closed estuaries. We postulate that this is a factor contributing to the relatively low numbers of adult P. knysnaensis found in sandy temporarily open/closed estuaries of the region, although the role of marine connectivity, food availability, predation pressure and other environmental factors in this regard still remain largely unassessed.     

The distribution of the freshwater cichlid Sarotherodon mossambicus in estuarine systems

Synopsis The distribution of Sarotherodon mossambicus in south east African estuaries is reviewed. It occurs commonly in closed estuaries and coastal lakes but is absent from open tidal estuaries. When a closed estuary opens to the sea the species usually retreats to the upper reaches. It is euryhaline and eurythermal and can feed on a wide variety of foods but experiments indicate that S. mossambicus avoids areas where current speeds exceed 370 m h^–1. It is concluded that the distribution of S. mossambicus in estuarine systems is governed by an interplay of the following factors: salinity stability, water currents, suitable breeding sites, presence of marginal vegetation, marine competitors and marine piscivorous fish.     

Recruitment and growth of juvenile yellowfin bream, Acanthopagrus australis Günther (Sparidae), in an Australian intermittently open estuary

Recruitment and growth of juvenile yellowfin bream, Acanthopagrus australis Günther (Sparidae), an economically important species, were examined in an intermittently open Australian estuary for 13 months. Fish had a mean fork length of 77.9 (± SE 9.5) mm after 1 year. Growth rates were faster than those found in previous studies in permanently open estuaries and appeared to be dependent upon the water temperature, which ranged from 0.04 mm/day in winter to 0.67 mm/day during summer. However, periods of slower growth may also reflect the emigration of larger fish from the cohort when the estuary was open and which coincided with lower water temperatures. Growth and length-at-age of fish using length-frequency data may be estimated more accurately in intermittently open estuaries because the time of recruitment can be determined, especially when the estuary opening period is short.     

Reproductive biology and larval morphology of the marine plotosid Cnidoglanis macrocephalus (Teleostei) in a seasonally closed Australian estuary

Monthly trends shown by gonadosomatic indices, the prevalence of the different gonadal stages, and the size distribution of the oocytes, indicate that the large marine and commercially important plotosid Cnidoglanis macrocephalus spawns in Wilson Inlet between October and January. The conclusion that spawning occurs within this seasonally closed estuary was confirmed by the presence of males in large nests and by the capture of newly-hatched, yolk sac larvae from one of those nests. The fact that C. macrocephalus, which is also widely distributed in coastal marine waters throughout much of southern Australia, can spawn within Wilson Inlet would be of particular value to this species in those periods when closure of the estuary would preclude a seawards spawning migration. Sexual maturity is size dependent, with spawning rarely occurring before fish have reached a total length of 425 mm. Sexual maturity was attained by a few fish at the end of their second year, by several at the end of their third year and by most, if not all fish, at the end of their fourth year. Comparisons with data for the more northern and permanently open Swan Estuary indicate that C. macrocephalus also spawns within that system and that the spawning time of this species is related to water temperature. The adult male guards the larvae under its pelvic fins in burrows. The larvae increased in total length from 29 mm just after hatching to 43 mm in the 17–18 days after capture, during which time their yolk sac was resorbed. Details are given of the morphology, morphometrics, meristics and pigmentation of larval C. macrocephalus. In comparison with the larvae of three other plotosid genera, the larva of C. macrocephalus is far larger in size and more developed at hatching and takes a shorter time to transform into a juvenile.     

The benthic macrofauna along the estuarine gradient of the Schelde estuary

The intertidal benthic macrofauna of the Schelde estuary (The Netherlands and Belgium) was sampled in late autumn of 1990 at 50 stations along the whole salinity gradient (between Vlissingen and Dendermonde), including the freshwater tidal part. All stations were situated in sheltered areas with a relatively muddy sediment. Species richness, diversity and total biomass of the benthic macrofauna decreased along the salinity gradient from Vlissingen to Dendermonde, while total density showed no clear trend. Especially the oligohaline and freshwater tidal part of the Schelde estuary was characterized by a very impoverished benthic community, composed only of Oligochaeta. No other species (freshwater, marine or brackish) was observed in this part of the estuary. The marine part had a more diverse macrozoobenthos structure than that of the brackish part. Species found only in the marine zone areCerastoderma edule, Tharyx marioni, Eteone longa, Nephtys hombergii andCapitella capitata. In the brackish part of the estuary,Corophium volutator was a typical, dominant species. However, a lot of the dominant species were common in both the marine and brackish part of the Schelde estuary (e.g. Heteromastus filiformis, Pygospio elegans, Nereis diversicolor, Macoma balthica). The observed gradient in species composition and dominance is compared with some other European estuaries. The marine and brackish part of the Schelde estuary is quite similar to other european estuaries. The freshwater tidal part, however, was more impoverished.     

Comparisons between the roles played by estuaries in the life cycles of fishes in temperate Western Australia and Southern Africa

Synopsis The rivers of south-western Australia and south-eastern Africa lie at similar latitudes, open into the Indian Ocean and frequently have estuaries that are periodically closed off from the sea by sand bars at their mouths. The present study has compared the species, genera and families represented in the fish assemblages of estuaries in south-western Australia and temperate southern Africa, i.e. below 31°S, and the way in which these estuaries are used by fish. The Clupeidae, Mugilidae, Atherinidae and Gobiidae were important families in both regions. However, the Terapontidae and Tetraodontidae, and the tropical families Apogonidae and Gerreidae, were represented by large numbers of individuals only in the estuaries of south-western Australia. Although 45 out of a total of 112 families and 32 of 233 genera occurred in both south-western Australia and temperate southern Africa, only 15 of the 326 species were found in both regions. The contributions made by the number of marine species which regularly enter estuaries in large numbers (marine estuarine-opportunists) to the total number of species recorded in the estuaries of south-western Australia and temperate southern Africa were similar (13.4 and 12.2% respectively) and the same was also true of species capable of completing their life cycles in estuaries (8.8 and 8.2%). The number of fresh water and diadromous species recorded in both regions was small. By contrast, the species of marine stragglers contributed approximately 70% to the total number of species in both regions. The adaptations of marine estuarine-opportunists and estuarine spawners to life in estuaries, and particularly to the effects of the closure of estuary mouths, is discussed. Although only one marine species was restricted to estuaries at any particular interval of its life cycle in south-western Australia, the juveniles of a number of marine species were confined to estuaries in temperate southern Africa. It is suggested that this difference can be attributed to the presence of a greater area and quality of alternative nursery habitats in the inshore marine environments in south-western Australia than in southern Africa.     

Characteristics of the ichthyofauna of a temperate microtidal estuary with a reverse salinity gradient,including inter‐decadal comparisons

Data on the fish fauna of the Leschenault Estuary on the lower west coast of Australia were collected and used as a model to elucidate the characteristics of permanently open estuaries with a reverse salinity gradient, which undergo seasonal changes similar to many other estuaries with Mediterranean climate. Focus was placed on determining (1) the relationships of the number of species, density, life cycle category and species composition of fishes with region (within estuary), season and year and salinity, (2) whether species are partitioned along the lengths of such systems and (3) the extent and significance of any inter‐decadal changes in species composition. The analyses and interpretation involved using multi‐factorial permutational multivariate analysis of variance (PERMANOVA) and analysis of similarity (ANOSIM) designs, and three new or recently published visualization tools, i.e. modified non‐metric multidimensional scaling (nMDS) plots, coherent species curves and segmented bubble plots. The base, lower, upper and apex regions of the Leschenault Estuary, along which the salinity increased in each season except in winter when most rainfall occurs, were sampled seasonally for the 2 years between winter 2008 and autumn 2010. Estuarine residents contributed twice as many individuals, but less than half the number of species as marine taxa. While the numbers of marine species and estuarine residents declined between the base or lower and apex regions, the individuals of marine species dominated the catches in the base region and estuarine residents in the other three regions. Ichthyofaunal composition in each region underwent conspicuous annual cyclical changes, due to time‐staggered differences in recruitment among species, and changed sequentially along the estuary, both paralleling salinity trends. Different groups of species characterized the fauna in the different regions and seasons, thereby partitioning resources among species. The ichthyofauna of the apex region, in which salinities reached 54 and temperatures 36° C, recorded the highest maximum density and, in terms of abundance, was dominated (90%) by three atherinid species, emphasizing the ability of this family to tolerate extreme conditions. Comparisons between the data for 2008–2010 and 1994 demonstrate that the spotted hardyhead Craterocephalus mugiloides and the common hardyhead Atherinomorus vaigiensis had colonized and become abundant in the Leschenault Estuary in the intervening period. This represents a southwards extension of the distribution of these essentially tropical species during a period of increasing coastal water temperatures as a result of climate change. The abundance of weed‐associated species, e.g. the western gobbleguts Ostorhinchus rueppellii and the soldier Gymnapistes marmoratus, increased, whereas that of the longfinned goby Favonigobius lateralis decreased, probably reflecting increases in eutrophication and siltation, respectively.     

Life-history styles of fishes in South African estuaries

Synopsis Fish utilizing South African estuaries may be divided into two major groups according to the location of their spawning sites. The marine group comprises large species which spawn at sea, enter estuaries mainly as juveniles, and return to the sea prior to attaining sexual maturity. The estuarine group is dominated by small species which have the ability to complete their life cycle within the estuarine environment. They tend to produce relatively few, demersal eggs, or exhibit parental care, which facilitates the retention of eggs and young within the estuary, whereas the marine group release large numbers of small pelagic eggs during spawning and exhibit no parental care. This is contrary to the theory that estuaries (unpredictable environments) should favour altricial life-history styles and the marine inshore zone (a more predictable environment) should favour precocial styles. However, if the total ichthyofauna of South African estuaries is considered, then altricial species predominate. The fact that both altricial and precocial traits are well represented within the overall estuarine fish community suggests that the various taxa have adapted their life-history styles, in different ways, to ensure the utilization of abundant food resources available within these fluctuating systems. A detailed comparison of the life-history styles of the estuarine teleostGilchristella aestuaria and the marine fishMugil cephalus is used to illustrate the contrasting manner in which these two species have succeeded in exploiting South African estuaries.     

Changes in rocky reef fish assemblages throughout an estuary with a restricted inlet

Rocky reef habitat is common in many estuaries, yet its role as a habitat for fishes is poorly understood. There is also limited understanding of how access of coastal species into estuaries and habitat quality can affect the distribution of rocky reef fishes within estuaries. This study used baited remote underwater video stations to determine spatial patterns in fish assemblages associated with rocky reef habitat throughout a barrier estuary with a permanently open but restricted inlet. Estuarine rocky reefs provided habitat for a diverse assemblage of fishes, many of which were large juveniles and subadults. In the absence of a pronounced salinity or temperature gradient, a clear transition in fish assemblages occurred from coastal waters, through the inlet channel, to the central estuary, and into the inner estuary. The inlet channel, notably its narrowness and length, limits tidal input into this estuary, which acts as a significant impediment to the dispersal of many coastal fishes, and insufficient habitat excludes many coastal rocky reef species from the inner estuary. This study highlights the need to recognise estuarine rocky reefs as providing habitat for diverse fish assemblages and the role inlets play in restricting access of coastal species.     

Zooplankton in different types of estuaries (using Curonian and Vistula estuaries as an example)

Based upon the results of long-term monitoring of zooplankton in the Curonian and Vistula estuaries carried out by the author, specific features of the structure and functioning of the zooplankton communities are studied. It is shown that the zooplankton communities in the open (Vistula Bay) and closed (Curonina Bay) types of estuaries differ in the species structures, numbers, biomasses, and functioning. The similarity of the zooplankton communities of these estuaries has to do with the commonness of their trophic structure. The zooplankton community in the Vistula estuary is more affected by the gradient of abiotic factors than the Curonian bay community. The mechanism providing the stability of the zooplankton communities in the open-type estuaries is the energy flow through the omnivorous species and the increase in the number of facultative predators at the terminal links of the trophic chain. It is shown that the Vistula estuary zooplankton community has a higher specific productivity rate and a higher rate of its circulation and use.     

Dietary compositions of the sparid <Emphasis Type="Italic">Acanthopagrus</Emphasis><Emphasis Type="Italic"> butcheri</Emphasis> in three normally closed and variably hypersaline estuaries differ markedly

We have compared the species composition and diversity of the diets of black bream, Acanthopagrus butcheri, in three normally closed estuaries on the central south coast of Western Australia, which vary markedly in the extents to which they become hypersaline during dry periods. Although black bream was caught seasonally in Stokes Inlet during this 3-year study, it was obtained from the Hamersley and Culham inlets only during the earlier seasons because salinities in those two estuaries subsequently rose to levels that caused massive mortalities of this sparid. Although a wide range of taxa, including macrophytes, polychaetes, molluscs, crustaceans, insects and teleosts, were ingested by A. butcheri in each estuary, the frequencies of ingestion and volumetric dietary contributions of these taxa varied greatly among the fish in these three estuaries. Thus, for example, in comparison with other estuaries, relatively greater contributions were made to the diet by polychaetes and crustaceans in Stokes Inlet, by macrophytes in Hamersley Inlet, and by insects (mainly chironomid larvae) in Culham Inlet. The relatively greater contribution of teleosts to the diets of black bream in the Hamersley and Culham inlets than in Stokes Inlet, and also differences in the main teleost species ingested in the first two estuaries, are consistent with differences in the densities of fish overall and of the main fish species in those estuaries. The diversity of the diet was far greater in Stokes Inlet than in the other two far more variably saline estuaries, presumably reflecting a greater diversity of food. The dietary compositions of black bream in upstream pools in the tributary of Culham Inlet, which offer refuge when salinities increase markedly in the main body of the estuary, differ from those in those downstream regions, further emphasising the opportunistic nature of the feeding behaviour of black bream. The dietary compositions of black bream underwent size-related changes, but the taxa contributing most to those changes varied greatly among estuaries. Size-related changes would be particularly beneficial in reducing intraspecific competition for food in the two estuaries that vary greatly in salinity and would thus be likely to contain a less diverse range of prey.     

A COMPARISON OF THE MOLLUSCAN COMMUNITIES ON INTERTIDAL SAND FLATS IN OYSTER HARBOUR AND PEEL INLET, WESTERN AUSTRALIA

Intertidal molluscs on sandflats in Oyster Harbour and PeelInlet, Western Australia, were compared using transects. Twenty-fourspecies of molluscs were recorded in Oyster Harbour and only7 in Peel Inlet. Two groupings were foundin Oyster Harbour:a midlittoral component and infralittoral fringe. There wasno vertical community structure in Peel Inlet. Molluscs retainedon a 1 mm sieve had about the same mean density and biomassin the two areas but molluscs retained on a 2 mm sieve had amean density of 203/m² and a biomass of 132 g/m² in Oyster Harbourand were absent in Peel Inlet. Several possible reasons forthe absence of large molluscs in Peel Inlet are discussed. PeelInlet is on the west coast in a faunal overlap zone where thenumber of species is reduced. The mean temperature in Peel Inletis 2-3°C higher than in Oyster Harbour and salinity variationsare much more pronounced. (Received 20 November 1979;