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.     

Fish trophic structure in estuaries,with particular emphasis on estuarine typology and zoogeography

A comparative analysis of the fish trophic structure was undertaken on some 190 South African estuaries spanning three zoogeographic regions and incorporating three broad estuarine types. Fish biomass trophic guild compositions and biomass trophic spectrum profiles were analysed using multivariate statistical techniques and included both inter‐regional (zoogeographic) and intra‐regional (estuarine typology) comparisons. Differences in the fish trophic structure of the various estuary types within each zoogeographic region were observed; these were linked to the relative biomass contribution of the various trophic guilds and also to differences in biomass trophic spectrum profiles of the fishes in each estuary type within each region. In spite of these differences in trophic structure, all estuaries were dominated by detritivores, which suggests that the main food source (detritus) is similar in all biogeographic regions. Preliminary indications are that a similar dependence by estuary‐associated fishes on detritus food sources exists on a global basis but that detailed studies are required in order to confirm this assertion.     

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.     

Life cycles and challenges for fish species that breed in South African estuaries

The literature currently recognizes four guilds of estuarine resident fish species, namely solely estuarine, estuarine and marine, estuarine and freshwater, and estuarine migrant. In this review the life cycles of actual representatives from these four guilds are assessed to determine whether the current definitions, which have never been formally tested, are appropriate to fish species resident in South African estuaries. Detailed information and diagrammatic life cycles are provided for the selected species covered by this review. A potential new estuarine resident guild category is also identified, namely, those taxa that are primarily estuarine but also have subpopulations recorded in both adjacent marine and freshwater habitats. The full range of reproductive characteristics employed by estuary resident species is examined, ranging from live bearers, pouch and nest brooders, to a suite of oviparous taxa that attach their ova to estuarine rocks, shells, and submerged vegetation, all of which assists with larval retention within the estuarine environment. The small size and early reproductive maturity of most estuarine resident species is highlighted, with reduced vulnerability to predation in shallow, sheltered, often turbid estuary waters offering considerable protection during spawning events when compared to the open ocean. In addition, these small fish would not have to move considerable distances at any stage of their life cycle, since egg, larval, juvenile, and adult stages all occur in the same place. The existence of contingent subpopulations within many estuarine resident species is noted, physico-chemical stresses on these species are highlighted, and the eurytopic nature of these small fish taxa emphasized.     

The Influence of Selected Environmental Variables on Fish Assemblage Structure in a Range of Southeast African Estuaries

This study compared the ichthyofaunal assemblages in 10 Eastern Cape Province, South Africa, estuaries to identify whether there were any structural variations between the assemblages and to determine the environmental and physical factors linked to these variations. Species belonging to both the marine migrant and estuarine resident groups, by consistent representation within each estuary grouping, were responsible for the separation of fish assemblage structures between the different types of systems. The data further revealed that fish assemblages in the three main types of estuaries investigated were distinct, linked primarily to estuary mouth status and estuary size.     

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.     

FISH CONSERVATION IN SOUTH AFRICAN ESTUARIES: PRESSURES,PROBLEMS AND PROSPECTS

Summary

A review of the conservation status of fish in the estuarine environment around the South African coastline reveals that some species face serious problems associated either with habitat destruction, and its associated biological, physical and chemical components or exploitation. The 65 species considered fall into three categories; truly estuarine species, which are dependent on estuaries for their entire life; marine species dependent on estuaries during the juvenile phase of their life cycle; and marine species whose juveniles occur mainly in estuaries but are also found at sea. Included in the first two categories are 14 species of fish which are on the South African Red Data list. These comprise one species which is endangered, five which are vulnerable and eight which are rare. All groups are considered in relation to factors in estuaries which are affecting their populations. A conservation strategy is suggested for certain estuarine types or for specific estuaries which could ensure the survival of the entire range of estuarine faunas.     

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.     

Ichthyofaunal assemblages in estuaries: A South African case study

This review places the life-history styles of fishes associated with South African estuaries in a global context and presents a classification system incorporating all the major life-history categories for estuary-associated fish species around the world. In addition, it documents the early life histories of the major fish groups in South African estuaries, with particular emphasis on the differing modes of estuarine utilization by marine, estuarine and freshwater taxa.This review details factors influencing the ichthyofaunal community structure in South African estuaries. The availability of fish for recruitment into an estuary depends primarily upon the distributional range of euryhaline marine and estuarine species, with tropical and temperate taxa showing marked abundance trends. Within a particular biogeographic region, however, estuarine type and prevailing salinity regime have a major influence on the detailed ichthyofaunal structure that develops. There is an increasing preponderance of marine fish taxa when moving from a freshwater-dominated towards a seawater-dominated type of system, and a decline in species diversity between subtropical estuaries in the north-east and cool temperate systems in the south-west. Similar declines in fish species diversity between tropical and temperate estuaries in other parts of the world are highlighted.Fish assemblages in estuaries adjust constantly in response to changing seasons, salinities, turbidities, etc. Despite persistent fluctuations in both the biotic and abiotic environment, the basic ichthyofaunal structure appears to have an underlying stability and to be predictable in terms of the response of individual species to specific conditions. This stability seems to be governed by factors such as the dominance of eurytopic taxa within estuarine assemblages and the robust nature of food webs within these systems. The predictability arises from factors such as the seasonality associated with estuarine spawning cycles and juvenile fish recruitment patterns. These patterns, together with a well-documented resilience to a wide range of physico-chemical and biotic perturbations, appear to be an underlying feature of fish assemblages in estuaries around the world.In contrast to marine fish species, estuary-associated taxa have received little conservation attention. Apart from the designation of protected areas, the main direct means of conserving estuary-associated fish stocks include habitat conservation and controls over fishing methods, effort, efficiency and seasonality. Of these, the conservation of fish habitats, the most important, because healthy aquatic environments invariably support healthy fish populations. The use of estuarine sanctuaries for fish conservation is briefly reviewed, as well as the legislation governing the USA National Estuarine Research Reserve System (NERRS) and the Australian Marine and Estuarine Protected Area (MEPA) system. It is concluded that South Africa requires an expansion of the existing Estuarine Protected Area (EPA) network, as well as the upgrading of selected 'estuarine reserves' where fishing is permitted, into 'estuarine sanctuaries' where no exploitation of biological resources is allowed.     

Fish communities in temporarily open/closed estuaries from the warm- and cool-temperate regions of South Africa: A review

The majority of estuaries along the coastline of southern Africa are termed temporarily open/closed estuaries (TOCEs) and are closed off from the sea for varying periods by a sandbar which forms at the mouth. It is therefore important to understand the processes occurring within TOCEs and their importance to fishes in order to make sound management recommendations. Estuaries along the coast of South Africa and their associated fish assemblages are biogeographically distinct and occur in either a subtropical, warm-temperate or cool-temperate zone. There are 125 TOCEs found within the cool-temperate and warm-temperate zones. Most fish species found in TOCEs are the juveniles of marine taxa that breed at sea. Permanently open estuaries generally have a higher diversity of species than TOCEs, but TOCEs still provide important nursery areas for many marine species and numerically often have a higher proportion of estuarine resident species. Important taxa in terms of abundance and biomass in warm-temperate TOCEs include the sparids Rhabdosargus holubi and Lithognathus lithognathus, several mugilid species, estuarine residents (particularly Gilchristella aestuaria and Atherina breviceps) and the freshwater cichlid Oreochromis mossambicus. The diversity of fishes in cool-temperate TOCEs is low when compared with warm-temperate systems and Liza richardsonii tends to dominate catches by number and mass in most systems. Several species recorded in TOCEs show clear longitudinal distribution trends. For example Atherina breviceps is generally more abundant in the lower reaches of estuaries. Mouth state, particularly the frequency, timing and duration of mouth opening plays a key role in determining species richness, composition, diversity and abundance in TOCEs. Mouth state is directly linked to freshwater input. Reduced river inflow leads to prolonged mouth closure and shorter open phases, which inhibits immigration and emigration of marine fish species between estuaries and the sea. Understanding of the effects of various processes occurring within these systems, particularly variation in freshwater input, on the biota of these important systems facilitates the development of informed management recommendations.     

Species richness of benthic macrofauna in Danish estuaries and coastal areas

Aim Cross‐system comparisons of species richness of benthic macrofauna and environmental factors were made of estuaries and adjacent sea areas in order to reveal possible regulating factors of estuarine biodiversity. Location Denmark. Methods Annual species abundance and biomass data from four years, from unvegetated soft sea/estuary floors were used from 26 grids, of which 15 were situated in estuaries. Bottom water data for oxygen concentration, salinity, and temperature were obtained from stations in or close to, the grids. Data on nutrient loading, water residence time and morphology were obtained for whole estuaries. Species richness from the grids, standardized to the same sample size, were related to environmental variables using linear regression. Results Species richness was unrelated to oxygen deficiency and productivity, but positively related to salinity. However, an equally high degree of explanation (R² ～ 0.70) was obtained using a model where richness was positively related to saltwater flux, which was computed from estuary volume and residence time and corrected for freshwater flux. The relationship was present for the dominating groups with pelagic dispersal, Annelida and Mollusca, but not for Crustacea, where recruitment is mainly by benthic pathways. Saltwater flux was strongly positively correlated with salinity, illustrating the high importance of flux from adjacent sea areas for water renewal in Danish estuaries. Similarity of species composition was greater for sites within than between saltwater current pathways, and the increases of richness with saltwater flux and salinity in the estuaries was largely due to marine species occurring in the open sea areas. Conclusion Danish estuaries are largely open systems where a part of estuarine species richness is sustained by dispersal from a species pool in adjacent seas. Results are consistent with the barrier prediction of island biogeography theory, but not the island size prediction.     

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.     

Juvenile salmon in estuaries: comparisons between North American Atlantic and Pacific salmon populations

All anadromous fishes, including juvenile salmon, encounter estuarine habitats as they transition from riverine to marine environments. We compare the estuarine use between juvenile Atlantic salmon (Salmo salar) in the Penobscot River estuary and Pacific salmon (Oncorhynchus spp.) in the Columbia River estuary. Both estuaries have been degraded by anthropogenic activities. Atlantic and Pacific salmon populations in both basins rely heavily on hatchery inputs for persistence. Pacific salmon, as a group, represent a continuum of estuarine use, from species that move through rapidly to those that make extensive use of estuarine habitats. While Atlantic salmon estuarine use is predominantly similar to rapidly moving Pacific salmon, they can exhibit nearly the entire range of Pacific salmon estuarine use. Both slow and rapidly migrating Atlantic and Pacific salmon actively feed in estuarine environments, consuming insect and invertebrate prey. Interactions between juvenile salmon and estuarine fish communities are poorly understood in both estuaries, although they experience similar avian and marine mammal predators. Estuaries are clearly important for Atlantic and Pacific salmon, yet our understanding of this use is currently insufficient to make informed judgments about habitat quality or overall estuary health. This review of salmonid migration through and residency within estuaries identifies actions that could hasten restoration of both Atlantic and Pacific salmon populations.     

Do fish larvae have advantages over adults and other components for assessing estuarine ecological quality?

Using a novel approach to the assessment of ecological quality status of estuarine ecosystems, this study hypothesizes that compared to adult fishes and other components, the younger fish stages will be more sensitive and act as an early warning and will reflect more effectively the ecological status of estuaries. Larval stages of fishes were used to assess the ecological quality status (EQS) of four NW Portuguese estuaries, with different types and magnitudes of human pressures. The larval fish assemblages, together with water column characteristics and pollution indicators (faecal contamination and nutrient load) were sampled in the Lima, Cávado, Ave and Douro estuaries, during spring and autumn 2009. The four estuaries were classified in terms of human pressures by a global pressure index that identified the Cávado estuary as the least impacted estuary, followed by the Ave and Lima, both classified as moderately impacted system, while the Douro was classified as a highly impacted system. The Ave emerged as the most polluted system, carrying the highest nutrient load and sewage contamination. Larval fish assemblages included estuarine species, marine migrants, marine stragglers and the larger estuaries had higher species richness. Compared to adult fishes, three multimetric fish-based indices classified the Cávado, Ave and Douro estuaries with a lower ecological status when fish larvae were used. Similarly, the EQS assessed by macroinvertebrates were equal or higher when compared with fish larvae results. The EQS assessed by fish larvae was negatively correlated with sewage contamination and nitrogen nutrients, but did not reflect other anthropogenic pressures expressed by the global pressure index, which was only detected by adult fish. Fish larvae assessments were able to detect short-time events of hydrological manipulations observed in the Cávado estuary, as well as a seasonal decrease of water quality especially evident in the Ave estuary. The indices used denoted some limitations to the use of fish larvae data, thus emphasising the need for new indices to test the observed tendency for lower EQS given by fish larvae. The advantages and disadvantages of using fish larvae as more sensitive and accurate bioindicators of ecosystem integrity is also discussed as a means of providing strategically important information for improved estuarine management.     

Connectivity between estuaries and marine environment: Integrating metrics to assess estuarine nursery function

Natural or anthropogenic induced variations in estuaries and the dynamics of marine fish populations potentially promote differences in connectivity between estuaries and marine areas, i.e. in their importance as nursery grounds. Within this context, an integrated assessment of the differential nursery function of the main estuaries along the Portuguese coast for commercial fish species common sole Solea solea, Senegalese sole Solea senegalensis, flounder Platichthys flesus and sea bass Dicentrarchus labrax was performed through several indicators based on available data. Contribution of individual estuaries to marine subpopulations was measured with potential metrics (juvenile density, habitat quantity, juvenile number and habitat quality within estuaries) and effective metrics (estuarine source of young adults in marine environment measured via otolith elemental fingerprints). The relationship between the two types of metrics was also assessed. Estuaries identified as important nursery and/or effective juvenile habitat (EJH) differed with species and no single estuary was best for all, highlighting species-specific regulation of nursery function. Multiple species assessment of nursery and EJH function differed among estuaries. Management and conservation of estuaries should focus on sites with higher contributions to adult subpopulations of multiple species. The importance of defining precise scientific and management objectives was emphasized by the different rankings of estuaries obtained with nursery or EJH criteria. Potential and effective contribution of estuaries were not significantly correlated, but in a quantitative analysis juvenile densities and number of juveniles seem related with effective contribution in some species. An agreement between potential and effective contributions of estuaries is concurrent with the acknowledged minor role of juvenile stage processes in regulation of recruitment to adult subpopulations.     

The distribution of macroinvertebrates and fishes in Tasmanian estuaries

The distributions of 390 taxa of benthic macroinvertebrates collected in forty-eight estuaries and 101 fish species collected in seventy-five Tasmanian estuaries were related to geographical and environmental variables. Distribution patterns for the two taxonomic groups were largely congruent at both between and within-estuary scales. Faunal composition and the number of species collected at a site were primarily related to site salinity, the biomass of seagrass and tidal range. At the broader estuary scale, the distributions of macroinvertebrate and fish assemblages were primarily correlated with the presence of an entrance bar. Species richness varied with geographical location for both macrofauna and fishes, with highest numbers of species occurring in the Furneaux Group, north-eastern Tasmania and south-eastern Tasmania. These patterns primarily reflected differences in estuary type between regions rather than concentrations of locally endemic species. Although the majority of species collected during the study were marine vagrants, they constituted a very low proportion of total animal densities within estuaries. Only four species considered exotic to Tasmania were identifed. Nearly all species recorded from Tasmanian estuaries occurred widely within the state and have also been recorded in south-eastern Australia. Only 1% of estuarine fish species and < 5% of invertebrate species were considered endemic to the state. The generally wide ranges of species around Tasmania were complicated by (i) the absence of most species from the west coast (ii) a small (< 10%) component of species that occurred only in the north-east and Furneaux Group (eastern Bass Strait), and (iii) a few species (< 5%) restricted to other regions. The low number of species recorded from estuaries along the western Tasmanian coast reflected extremely low faunal biomass in that area. This depression in biomass on the west coast was attributed to unusually low concentrations of dissolved nutrients in rivers and dark tannin-stained waters which greatly restricted algal photosynthesis and primary production.     

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.     

Estuarine fish communities respond to climate variability over both river and ocean basins

Estuaries are dynamic environments at the land–sea interface that are strongly affected by interannual climate variability. Ocean–atmosphere processes propagate into estuaries from the sea, and atmospheric processes over land propagate into estuaries from watersheds. We examined the effects of these two separate climate‐driven processes on pelagic and demersal fish community structure along the salinity gradient in the San Francisco Estuary, California, USA. A 33‐year data set (1980–2012) on pelagic and demersal fishes spanning the freshwater to marine regions of the estuary suggested the existence of five estuarine salinity fish guilds: limnetic (salinity = 0–1), oligohaline (salinity = 1–12), mesohaline (salinity = 6–19), polyhaline (salinity = 19–28), and euhaline (salinity = 29–32). Climatic effects propagating from the adjacent Pacific Ocean, indexed by the North Pacific Gyre Oscillation (NPGO), affected demersal and pelagic fish community structure in the euhaline and polyhaline guilds. Climatic effects propagating over land, indexed as freshwater outflow from the watershed (OUT), affected demersal and pelagic fish community structure in the oligohaline, mesohaline, polyhaline, and euhaline guilds. The effects of OUT propagated further down the estuary salinity gradient than the effects of NPGO that propagated up the estuary salinity gradient, exemplifying the role of variable freshwater outflow as an important driver of biotic communities in river‐dominated estuaries. These results illustrate how unique sources of climate variability interact to drive biotic communities and, therefore, that climate change is likely to be an important driver in shaping the future trajectory of biotic communities in estuaries and other transitional habitats.     

Estuarine recruitment of a marine goby reconstructed with an isotopic clock

Information on movement patterns of marine fishes between estuarine populations and stocks at sea is fundamental to understanding their population dynamics, life history tactics and behavior. Furthermore, understanding estuarine habitat use by marine fishes is crucial for their effective conservation and integrated estuarine management. Although large numbers of young marine fish make use of temperate estuaries in highly predictable abundance patterns, very little is known about how estuarine populations interact with the populations at sea. Recruitment of sand goby Pomatoschistus minutus (Pallas, 1770) into the low salinity zone of the Scheldt estuary (Belgium) was reconstructed over an entire year by means of an isotopic clock. These results were combined with a growth model to yield age and length at immigration. Sand gobies entered the upper Scheldt estuary almost continuously from May onwards, except in July when they appeared to avoid the estuary due to warm summer temperatures. About 70% of the fish caught in the upper estuary resided there for less than 1 month, which indicates a strong temporal overlap of immigration and emigration. This complex migration pattern suggests that estuarine residence is caused by trade-offs made at the individual level, whereby migration is probably triggered by temperature. The high turnover of individuals in the estuarine population leads us to question the functional role of the estuary for marine fishes. Sand gobies entering the upper estuary had a wide range of ages and body sizes, although they were at least 2 months old and had a minimum standard length of approximately 20 mm. This study shows that the use of an isotopic clock strongly complements catch data and is useful to describe the connectivity between populations.     

Why are there so few freshwater fish species in most estuaries?

The freshwater fish assemblage in most estuaries is not as species rich as the marine assemblage in the same systems. Coupled with this differential richness is an apparent inability by most freshwater fish species to penetrate estuarine zones that are mesohaline (salinity: 5·0–17·9), polyhaline (salinity: 18·0–29·9) or euhaline (salinity: 30·0–39·9). The reason why mesohaline waters are avoided by most freshwater fishes is difficult to explain from a physiological perspective as many of these species would be isosmotic within this salinity range. Perhaps, a key to the poor penetration of estuarine waters by freshwater taxa is an inability to develop chloride cells in gill filament epithelia, as well as a lack of other osmoregulatory adaptations present in euryhaline fishes. Only a few freshwater fish species, especially some of those belonging to the family Cichlidae, have become fully euryhaline and have successfully occupied a wide range of estuaries, sometimes even dominating in hyperhaline systems (salinity 40+). Indeed, this review found that there are few fish species that can be termed holohaline (i.e. capable of occupying waters with a salinity range of 0–100+) and, of these taxa, there is a disproportionally high number of freshwater species (e.g. Cyprinodon variegatus, Oreochromis mossambicus and Sarotherodon melanotheron). Factors such as increased competition for food and higher predation rates by piscivorous fishes and birds may also play an important role in the low species richness and abundance of freshwater taxa in estuaries. Added to this is the relatively low species richness of freshwater fishes in river catchments when compared with the normally higher diversity of marine fish species for potential estuarine colonization from the adjacent coastal waters. The almost complete absence of freshwater fish larvae from the estuarine ichthyoplankton further reinforces the poor representation of this guild within these systems. An explanation as to why more freshwater fish species have not become euryhaline and occupied a wide range of estuaries similar to their marine counterparts is probably due to a combination of the above described factors, with physiological restrictions pertaining to limited salinity tolerances probably playing the most important role.