Fine-scale genetic structure, estuarine colonization and incipient speciation in the marine silverside fish Odontesthes argentinensis

The identification of incipient ecological species represents an opportunity to investigate current evolutionary process where adaptive divergence and reproductive isolation are associated. In this study we analysed the genetic structure of marine and estuarine populations of the silverside fish Odontesthes argentinensis using nine microsatellite loci and 396 bp of the mitochondrial DNA (mtDNA) control region. Our main objective was to investigate the relationship among estuarine colonization, divergent selection and speciation in silversides. Significant genetic structure was detected among all marine and estuarine populations. Despite the low phylogeographic structure in mtDNA haplotypes, there was clear signal of local radiations of haplotypes in more ancient populations. Divergence among marine populations was interpreted as a combined result of homing behaviour, isolation by distance and drift. On the other hand, ecological shifts due to the colonization of estuarine habitats seem to have promoted rapid adaptive divergence and reproductive isolation in estuarine populations, which were considered as incipient ecological species. This conclusion is supported by the existence of a set of environmental factors required for successful reproduction of estuarine ecotypes. The pattern of genetic structure indicates that phenotypic and reproductive divergence evolved in the face of potential gene flow between populations. We suggest that the 'divergence-with-gene-flow' model of speciation may account for the diversification of estuarine populations. The approach used can potentially identify 'incipient estuarine species', being relevant to the investigation of the evolutionary relationships of silversides in several coastal regions of the world.     

Pre-adaptive plasticity in atherinids and the estuarine seat of teleost evolution

With the aid of models and selected examples, an analysis of speciation within the family Atherinidae is presented. This is a large family of pelagic fish, many members of which look strikingly similar. They are primarily a marine coastal group, favouring sheltered estuaries and lagoons, although in some regions they have invaded fresh waters.
Many atherinids show a high degree of intraspecific morphological variability which can be linked to their estuarine habitat. It is argued that estuarine and lagoonal environments are physically highly variable and this has acted to select for generalist genotypes able to adjust their morphology, physiology and behaviour to a wide range of conditions. Such plasticity pre-adapts atherinids to invade and rapidly speciate in fresh waters containing vacant niches. Examples of such invasions and speciation are presented, and the rate of evolution discussed.
The selective forces acting on atherinids should be common to other fish groups, suggesting that many freshwater fish may have estuarine ancestors. The influence of estuaries in evolution may be out of all proportion to their size.     

Phylogenetic relationships of Atherina hepsetus and Atherina boyeri (Pisces: Atherinidae) populations from Greece, based on mtDNA sequences

The genetic divergence and the phylogenetic relationships of six Atherina boyeri (freshwater and marine origin) and five Atherina hepsetus populations from Greece were investigated using partial sequence analysis of 12s rRNA, 16s rRNA and control region mtDNA segments. Three different well divergent groups were revealed; the first one includes A. boyeri populations living in the sea, the second includes A. boyeri populations living in the lakes and lagoons whereas the third one includes all A. hepsetus populations. Fifty-seven different haplotypes were detected among the populations studied. In all three mtDNA segments examined, sequence analysis revealed the existence of fixed haplotypic differences discriminating A. boyeri populations inhabiting the lagoon and the lakes from both the coastal A. boyeri and the A. hepsetus populations. The genetic divergence values estimated between coastal (marine) A. boyeri populations and those living in the lagoon and the lakes are of the same order of magnitude as those observed among coastal A. boyeri and A. hepsetus populations. The results obtained by different phylogenetic methods were identical. The deep sequence divergence with the fixed different haplotypes observed suggests the occurrence of a cryptic or sibling species within A. boyeri complex.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 151–161.     

Enzymatic survey of four populations of Atherina boyeri based on electrophoresis and the occurrence of a microsporidiosis

Biochemical genetics of a Atherinid species, Atherina boyeri were undertaken to determine variability. Among the 17 loci studied, 7 were polymorphic, of which three were retained for a more detailed comparison of populations inhabiting the French Mediterranean coastal lagoons. A microsporidiosis is present in these fishes and is limited to certain lagoons. Comparison of allelic frequencies at the polymorphic loci between fish populations of infected and healthy lagoons yielded no significant differences. Several hypotheses are discussed and attempt to agree parasite distribution with genetical results.     

Four centuries of cumulative impacts on a Finnish river and its estuary: an ecosystem health-approach

Changes in forest cover and draining of wetlands for agriculture appear to have caused changes in the aquatic ecosystem of the River Kyrönjoki by the 16th century. In the 19th century, a decline of salmonid fish populations was widely observed as a further sign of degradation. During the latter half of the present century intensified use of naturally acidic soils has resulted in increased acidic run-off. Deterioration of water quality has extended to the estuary, where it has caused large fish-kills and affected the reproductive success of coastal species. Degradation of the coastal ecosystem, first observed in the decline of salmonids and later as a more general decline of other coastal fish populations, can be linked to spatially restricted events. The loss of key river and estuarine habitats exerted an effect over the reproduction and abundance of species migrating throughout the system. This effect contributed to observed temporal and spatial discontinuities in the degradation history. Monitoring changes in critical habitats may prove to be an early indicator of changes in the health of estuarine and coastal aquatic ecosystems.     

Vicariance and dispersal effects on phylogeographic structure and speciation in a widespread estuarine invertebrate

Vicariance and dispersal can strongly influence population genetic structure and allopatric speciation, but their importance in the origin of marine biodiversity is unresolved. In transitional estuarine environments, habitat discreteness and dispersal barriers could enhance divergence and provide insight to evolutionary mechanisms underlying marine and freshwater biodiversity. We examined this by assessing phylogeographic structure in the widespread amphipod Gammarus tigrinus across 13 estuaries spanning its northwest Atlantic range from Quebec to Florida. Mitochondrial cytochrome c oxidase I and nuclear internal transcribed spacer 1 phylogenies supported deep genetic structure consistent with Pliocene separation and cryptic northern and southern species. This break occurred across the Virginian-Carolinian coastal biogeographic zone, where an oceanographic discontinuity may restrict gene flow. Ten estuarine populations of the northern species occurred in four distinct clades, supportive of Pleistocene separation. Glaciation effects on genetic structure of estuarine populations are largely unknown, but analysis of molecular variance (AMOVA) supported a phylogeographic break among clades in formerly glaciated versus nonglaciated areas across Cape Cod, Massachusetts. This finding was concordant with patterns in other coastal species, though there was no significant relationship between latitude and genetic diversity. This supports Pleistocene vicariance events and divergence of clades in different northern glacial refugia. AMOVA results and private haplotypes in most populations support an allopatric distribution across estuaries. Clade mixture zones are consistent with historical colonization and human-mediated transfer. An isolation-by-distance model of divergence was detected after we excluded a suspected invasive haplotype in the St. Lawrence estuary. The occurrence of cryptic species and divergent population structure support limited dispersal, dispersed habitat distribution, and historical factors as important determinants of estuarine speciation and diversification.     

Male reproductive strategy and reserve allocation in sand smelt from brackish lagoons of southern France

In male sand smelt Atherina boyeri , from three coastal brackish lagoons of southern France, the duration of the reproductive season differed according to fish size: larger fish exhibited an extended reproductive season from February to August while smaller males reproduced only from April to June-July. Smaller and larger fish showed differences in their seasonal pattern of reserves. Through the cold season, energy content was exhausted for over-winter maintenance and reproductive needs, and was restored just after fish ceased reproducing. In smaller fish, however, reserves were more rapidly restored than in larger fish, and exhibited a decrease in autumn. This result suggested that small fish invested their energy into growth and winter survival rather than reproduction while older fish invested in reproduction. These results are discussed in terms of life history Strategies.     

Life strategies of mykiss <Emphasis Type="Italic">Parasalmo mykiss</Emphasis> and trout <Emphasis Type="Italic">Salmo trutta</Emphasis> introduced in water bodies of Chili

Materials of preliminary investigations of the life strategy of mykiss Parasalmo mykiss and trout Salmo trutta introduced to water bodies of Chili are considered. Analysis of the structure of scales demonstrated that, in mykiss and trout, the estuarine and riverine-estuarine types of life strategy prevail. This seems to be related to good feeding conditions in fjords. The hypothesis is suggested that, in the area of Chilean fjords, the successful introduction to rivers and artificial rearing in cages in brackish water resulted in the formation of a peculiar and highly dynamic population system. Elements of this system are from both fish of natural self-reproducing populations and fish grown in cages. There is a continuous exchange with genetic information between them as the fish that have escaped from cages attain maturity, arrive to rivers for spawning, and reproduce together with wild fish.     

The biology of Boyer's sand smelt, Atherina boyeri Risso in the Bardawil Lagoon on the Mediterranean coast of Sinai

The present study describes some aspects of the life history of Boyer's sand smelt Atherina boyeri in the hypersaline Bardawil Lagoon on the Mediterranean coast of the Sinai peninsula, Egypt. Monthly samples were collected with a small-mesh experimental beach-seine during the years 1973–1974. Age determinations were based on otoliths readings and length frequency distributions which indicate that the Bardawil Lagoon population consists mostly of 0-age group. The largest recorded fish was 63 mm in standard length (S.L.). Length-weight relationship has been calculated as: W = 13.7 × 10⁻⁶× L ^2.93 where W = weight (g) and L =S.L. (mm). Atherina boyeri mature at the length of 34 mm. Spawning takes place from March to September. The average number of eggs in the ovary of a ripe female at the beginning of the spawning season was found to be 522. Females of all sizes were more numerous than males. In immature fish, less than 34 mm long, females constituted 53% of the population. In larger fish the number of females was higher; in fish over 34 mm long females constituted 64% and in fish over 51 mm—96%. Atherina boyeri in the Bardawil Lagoon feeds on both zoobenthos and zooplankton, mainly amphipods and copepods. Polychaetes, mysidaceans, insects and fish were also represented. Atherina boyeri is the host of endoparasitic Trematoda, Acanthocephala and Nematoda. The examined fish were found to have metacercaria in their mesenteries and liver.     

Habitat predicts reproductive superfetation and body shape in the livebearing fish Poeciliopsis turrubarensis

Superfetation, the ability of females to simultaneously carry more than one brood at different developmental stages, is an unusual reproductive strategy that has independently evolved several times in the livebearing fish family Poeciliidae. Why this strategy has evolved remains uncertain. One hypothesis is that superfetation is a response to selective pressures that constrain the physical space within a female in which her offspring can develop. This hypothesis is reasonable, because superfetation should reduce the total volume needed to house developing embryos – that is, fewer large, fully developed embryos will be held by a superfetating female (with several broods at different developmental stages) than a non‐superfetating female (where all embryos reach a fully developed stage at the same time). In this study, we explore this ‘morphological constraint’ hypothesis of superfetation by examining the livebearing fish, Poeciliopsis turrubarensis. We found that populations vary markedly in degree of superfetation, with individuals carrying from two to four distinct broods across different geographic areas. These populations also occupy a range of habitat types: some populations occur in slow moving coastal rivers near the ocean, while other populations occur far inland in fast moving waters that drain steep mountain environments. In comparing populations from these two types of environments, we find a strong association between stream habitat type and the degree of superfetation within populations. Fish from inland populations have higher levels of superfetation than their coastal counterparts. In addition, geometric morphometric analysis revealed that inland populations are also more fusiform than fish from coastal locations. Combined, these two lines of evidence support the ‘morphological constraint’ hypothesis, and suggest that the life history strategy of superfetation could be driven by environmental pressures that favor a more streamlined phenotype.     

A study of the population biology of Athevina boyeri Risso, 1810 in Aberthaw Lagoon, on the Bristol Channel, in South Wales

The biology of the sixth population of A. boyeri Risso, 1810 to be recorded outside of the Mediterranean basin is described. The 2 year life cycle was similar to that of other populations of the species that have been investigated. A maximum standard length of 93 mm was recorded for 2-year-old fish. Sexual maturity was attained by females at a length of 39 mm. A temporal lag in ovarian development was noted between the two spawning age groups, with the older age group spawning first. Details are provided of batch spawning in A. boyeriand the difficulties of assessing fecundity in batch spawners is discussed. Discrepancies between direct and indirect methods of assessing batch size indicated that the eggs around the largest mode may not represent the next batch to be spawned. Evidence is presented to suggest that in A. boyeri these eggs represent a ' store' from which batches of eggs are subsequently released.     

Life history strategies of four small-size fishes in the Suez Canal, Egypt

Of 35 species collected from the shores of the Suez Canal and its lakes, five were abundant year round. Sihouettea aegyptia and Aphanius dispar are of Red Sea (warm water) origin, whereas Pomatoschistus marmoratus, Aphanius fasciatus and Atherina boyeri are Mediterranean species. S. aegyptia and A. dispar are distributed throughout the Suez system whereas A. fasciatus is restricted to the northern canal. P. marmoratus and A. boyeri have not spread southward beyond the Bitter Lakes. Salinity is the main limiting factor for the distribution of Mediterranean species. S. aegyptia and A. dispar are dominant in the Bitter Lakes (salinity 44‰), whereas P. marmoratus and A. boyeri are abundant in Timsah Lake (salinity 7·8–41·6‰). There was considerable interannual and monthly variation in the relative abundance of each species. The populations were dominated by a single age group, and life spans were no more than 2 years. Rapid growth was evident during the first spring in P. marmoratus and A. boyeri and during the first summer, early autumn in S. aegyptia and A. dispar . The relative abundance of each food item in the diet varied with fish size and season. S. aegyptia, P. marmoratus and A. dispar fed mostly on harpacticoid copepods, polychaetes, demersal eggs, diatoms and blue-green algae whereas A. boyeri fed mostly on planktonic copepods. The warm water species are summer spawners, whereas the temperate species are autumn-winter spawners.     

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.     

The icthyofauna of the Mhlathuze coastal lakes: some preliminary results

The fish faunas of the four Mhlathuze coastal lakes and the lower river comprise a diverse assortment of over fifty marine, estuarine and freshwater species. Three freshwater species are endemic to KwaZulu-Natal and nine estuarine species are endemic to southern Africa. Five species are of conservation significance. Species numbers in Lakes Mzingazi and Cubhu are similar historically and both lakes served as secondary nursery habitats for estuarine associated fishes. This role has been impacted by the construction of weirs at their outlets which prevent successful recruitment of estuarine species, especially during drought years when lake water levels are low. The fish faunas of Lakes Nsezi and Mangeza are depauperate and lack marine or estuarine components. In order that these systems fulfil their potential function as secondary nursery habitats to many estuarine fish species, minimum lake water levels must be set to ensure sufficient outflows at proposed fish ladders during critical spawning and migration times.     

Morphological variation in British Atherinids, and the status of Atherina presbyter Guvier (Pisces: Atherinidae)

A series of morphometric and meristic analyses conducted on specimens of sand smelt, Atherina , from populations around the British Isles, and including classic A. boyeri and A. presbyter forms, has shown that the characteristics variously used in the past to distinguish these two species are invalid. Multivariate analyses showed no significant splitting of the material into two groups which might correlate to these two species; classic A. boyeri and the very large Atlantic coast A. presbyter individuals represent the tails of a continuum of form. From these and published data, it is concluded that the A. boyeri morphology varies under the influence of conditions of temperature and salinity during embryo development, and the isolation of populations maintains comparative morphological distinctions resulting from local selection and random genetic drift.     

Coastal and estuarine ecological risk assessment: the need for a more formal approach to stressor identification

Application of ecological risk assessment to coastal and estuarine systems is accelerating although it initially lagged behind applications to land and freshwaters. Broader spatial and temporal scales, and multiple stressor integration are appropriately being considered more frequently in all risk assessment activities. This expansion and integration is essential for coastal risk assessment. Because coastal assessments must deal with co-occurrence of several candidate stressors manifesting within broad spatial and temporal scales, wider use of formal methods for assessing causal linkages is needed. Simple Bayesian inference techniques are discussed here to demonstrate their utility in quantifying the belief warranted by available information. The applicability of Bayesian techniques is illustrated with two examples, possible causes of fish kills on the Mid-Atlantic US coast and possible causes of hepatic lesions in fish of Puget Sound (Washington, US).     

Habitat quality of estuarine nursery grounds: Integrating non-biological indicators and multilevel biological responses in Solea senegalensis

Estuaries are highly valuable ecosystems that provide various goods and services to society, such as food provision and supporting nursery habitats for various aquatic species. Estuarine habitat quality assessment is thus critical in managing both ecological and economic value. In this work, various biological and non-biological indicators of habitat quality in estuarine nursery areas were determined, encompassing local environmental conditions, chemical contamination, anthropogenic pressures, juvenile Solea senegalensis condition, biomarkers response to contamination and juvenile density. The various indicators provided an integrated view on habitat quality and their responses were broadly concordant. Nursery quality assessment based on anthropogenic pressure indicators and fish biomarker responses were very similar, signaling nursery areas with higher anthropogenic pressure in Tejo and Ria de Aveiro estuaries. Yet, favorable environmental conditions across all sites could have contributed to lessen the potential hazardous biological effects of exposure to anthropogenic stressors, resulting in soles’ fairly good condition and generally high juvenile density. Nevertheless, a mismatch between high juvenile density and high estuarine contribution to adult coastal populations was observed in areas with higher anthropogenic pressures. Although a causal relationship cannot be established, the results emphasize the need to fully understand how the estuarine period spent in estuaries and local processes determine the quantity and quality of juveniles exported to marine adult populations, which is critical to achieve the full potential of the fish production service of estuaries and coastal stock replenishment.     

A review on the early life history and ecology of Japanese sea bass and implication for recruitment

Recruitment in marine fishes is regulated largely by the demographic changes that occur during the early life stages; therefore, a thorough understanding of early life stages is essential for predicting recruitment variability in fishes. Japanese sea bass (JSB), Lateolabrax japonicus, is a coastal marine fish distributed in East Asian coastal waters, and is regarded as highly important for commercial and recreational fisheries, for marine and brackish water aquaculture as well as for stock enhancement. JSB is a typical estuarine dependent temperate fish, which spawns in shelf areas and coastal embayments and the larvae and juveniles are dispersed and transported into shallow nearshore habitats and estuaries where they spend the early life. In this paper, we provide insight into the early life history and ecology of JSB through a revision of the available information and using the data we obtained from a relatively long-term research. We review and discuss the distribution and habitat use, food and feeding, age and growth, mortality and recruitment of larval and juvenile JSB in coastal waters around Japan. We extend our discussions in all available dimensions: habitat-specific, ontogenetic, and spatio-temporal, and highlight the importance of nursery habitats. We also discuss the implications of early life history for recruitment of JSB as well as the possible effects of climate change. At the end, we point out potential areas for future research.     

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.     

Salinity tolerance of non-native Asian swamp eels (Teleostei: Synbranchidae) in Florida,USA: comparison of three populations and implications for dispersal

Three populations of non-native Asian swamp eels are established in peninsular Florida (USA), and comprise two different genetic lineages. To assess potential for these fish to penetrate estuarine habitats or use coastal waters as dispersal routes, we determined their salinity tolerances. Swamp eels from the three Florida populations were tested by gradual (chronic) salinity increases; additionally, individuals from the Miami population were tested by abrupt (acute) salinity increases. Results showed significant tolerance by all populations to mesohaline waters: Mean survival time at 14 ppt was 63 days. The Homestead population, a genetically distinct lineage, exhibited greater tolerance to higher salinity than Tampa and Miami populations. Acute experiments indicated that swamp eels were capable of tolerating abrupt shifts from 0 to 16 ppt, with little mortality over 10 days. The broad salinity tolerance demonstrated by these experiments provides evidence that swamp eels are physiologically capable of infiltrating estuarine environments and using coastal waters to invade new freshwater systems.