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.

An overview of grey mullet (Mugilidae) global occurrence and species-rich ecoregions, with indications of possible past dispersal routes within the family.

This review examines the published literature on the distribution and species richness of the family Mugilidae around six continents as well as their phylogenetic relationships in a time-calibrated tree. Three mugilid species-rich regions were identified globally, namely the Coral Triangle, southern Asia and southern Africa, all of which have between 16 and 18 morphologically recognized species. Two of the species hotspots are tropical, and only southern Africa incorporates temperate waters. The centre of mugilid evolution and then global dispersion appears to be located in the tropical waters of the Indo-Pacific. Speciation within the family was promoted mainly by plate tectonics, eustatic sea-level changes, tropical climate and high habitat diversity within this region. The number of these tropical species that radiated out to the coastal waters of Africa and America was clearly less than the number that remained in the central Indo-Pacific region. Nonetheless, access by tropical mugilid species was greater to Africa than the Americas because of the direct tropical and subtropical coastal connectivity to the former continent. This enabled more tropical mugilids to reach Africa than America, with the latter probably relying on Palaeocene "island hopping" or "rafting" to move eastwards across the Pacific Ocean. In addition, tropical mugilids were able to access warm western and central African coastal waters from the Pacific prior to the closure of the Tethys Sea gap in the eastern Mediterranean and prior to the development of the cool Benguela Current upwelling off the southwest African coast, and from America through at least two transatlantic rafting events.

Contributions to the taxonomy of the mugilid genus Chelon Artedi (Teleostei: Mugilidae), with a major review of the status of C. persicus Senou, Randall & Okiyama, 1995.

New data based on multigene phylogenetic analyses using the COI, 16S, and cytb genes and subsequent molecular species delimitation revealed that the mugilid genus Chelon contains 11 species. Of these, two species, Chelon sp. A and Chelon sp. B, remained unidentified in previous studies. While Chelon sp. B seemingly is a close relative to C. dumerili (Steindachner, 1870), no more detailed information were provided for Chelon sp. A which is present along the east coast of South Africa. Genetic analyses performed in this study, revealed that specimens from the Arabian/Persian Gulf and the Red Sea are nested within one genetic lineage alongside Chelon sp. A. Morphological, morphometric, and meristic analyses of specimens from all three areas demonstrated that they belong to Chelon persicus Senou, Randall & Okyiama, 1995. Chelon persicus was originally described in the genus Chelon but was later placed either in the genus Liza or Planiliza. However, results presented herein confirm its placement in Chelon and its distribution range is extended in the Western Indian Ocean from the type locality (Arabian/Persian Gulf) west to the Red Sea and south to South Africa. Further, the uncertain validity of the recently described Chelon caeruleus Deef, 2018 from the Mediterranean Sea and a previously proposed generic character distinguishing Chelon from Planiliza and Parachelon, the shape of the paired postzygapophyses of the second vertebra, are briefly discussed.

Fish species in estuaries - from partial association to complete dependency.

Estuarine dependency by certain fish species has been clearly demonstrated in a number of studies, but the term has also been used for those species and guilds that are not dependent on estuaries. The origins and development of the term are explored, and definitions for four types of estuarine fish association are provided which may be helpful in facilitating classification and comparisons between fish taxa using estuarine systems in different parts of the world. In addition, a diagrammatic representation of more recent estuary-associated guild categories is presented to provide context and clarity on the estuarine dependency issue, and the fact that a continuum in terms of dependency is perhaps most appropriate for describing fish species relationships with estuaries. ETHICAL STATEMENT: This brief communication is a philosophical discussion and did not involve the capture, use or care of any living or dead fish. Therefore, no permit or animal ethics clearance was required to undertake the literature review.

Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent.

The Waterloo Farm lagerstätte in South Africa provides a uniquely well-preserved record of a Latest Devonian estuarine ecosystem. Ecological evidence from it is reviewed, contextualised, and compared with that available from the analogous Swartvlei estuarine lake, with a particular emphasis on their piscean inhabitants. Although the taxonomic affinities of the estuarine species are temporally very different, the overall patterns of utilisation prove to be remarkably congruent, with similar trophic structures. Significantly, both systems show evidence of widespread use of estuaries as fish nurseries by both resident and marine migrant taxa. Holocene estuaries are almost exclusively utilised by actinopterygians which are overwhelmingly dominated by oviparous species. Complex strategies are utilised by estuarine resident species to avoid exposure of eggs to environmental stresses that characterize these systems. By contrast, many of the groups utilising Devonian estuaries were likely live bearers, potentially allowing them to avoid the challenges faced by oviparous taxa. This may have contributed to dominance of these systems by non-actinoptergians prior to the End Devonian Mass Extinction. The association of early aquatic tetrapods at Waterloo Farm with a fish nursery environment is consistent with findings from North America, Belgium and Russia, and may be implied by the estuarine settings of a number of other Devonian tetrapods. Tetrapods apparently replace their sister group, the elpistostegids, in estuaries with both groups having been postulated to be adaptated to shallow water habitats where they could access small piscean prey. Correlation of tetrapods (and elpistostegids) with fish nursery areas in the Late Devonian lends strong support to this hypothesis, suggesting that adaptations permitting improved access to the abundant juvenile fish within the littoral zone of estuarine lakes and continental water bodies may have been pivotal in the evolution of tetrapods.

Microphytobenthos diversity and community structure across different micro-estuaries and micro-outlets: Effects of environmental variables on community structure.

This study forms the first basic assessment of microphytobenthos (MPB) dynamics in micro-estuaries and micro-outlets in southern Africa. It examines MPB community responses to environmental variables and further investigates MPB composition qualitatively across different micro-estuaries and micro-outlets over four seasons in a warm temperate region of the subcontinent. Combinations of multivariate analyses were used to explore similarities and differences in MPB communities between systems. Human-induced catchment changes between microsystems ranged from no alteration (rating 0; mostly micro-outlets) to extreme modification (rating 5; mostly micro-estuaries). Two hundred and sixty-seven MPB taxa were identified within all the microsystems, with 247 and 230 MPB taxa being observed in the micro-estuaries and micro-outlets, respectively. The MPB communities differed slightly in terms of microsystem types and seasons, but no significant differences were observed. Multivariate analyses (i.e. Boosted Regression Trees, Canonical Correspondence Analysis) showed that water column variables were significant and important in structuring MPB communities, with soluble reactive phosphorus, sediment pH, turbidity, ammonium and temperature being documented as key drivers. The MPB community composition clearly reflected the influence of catchment anthropogenic activities on species composition and structure. Moderately modified catchments resulted in MPB community structure variation among water bodies in relationship to land use and salinity gradients. The study found that; (i) by virtue of their size, microsystems and their catchments are likely to be particularly vulnerable to anthropogenic pressures when compared to systems of larger size; (ii) a typical impacted state may reflect reduced environmental heterogeneity which, compared to larger systems, may be achieved over much shorter time periods (following a particular event) or under much less intensive impacts; and (iii) the response in terms of MPB structure may predictably reflect a concomitant change from a complex community dynamic (structure and spatio-temporal attributes) to one that approaches a homogeneous structure (poor spatial zonation, strong taxonomic dominance, low species diversity).

A review of the influence of biogeography, riverine linkages, and marine connectivity on fish assemblages in evolving lagoons and lakes of coastal southern Africa.

The Holocene evolution of eight South African coastal lakes and lagoons is examined and related to changes in fish composition over that period. Historical and current connectivity with riverine and marine environments are the primary determinants of present-day fish assemblages in these systems. A small and remarkably consistent group of relict estuarine species have persisted in these coastal lakes and lagoons. The loss or reduction of connectivity with the sea has impacted on the diversity of marine fishes in all eight study systems, with no marine fishes occurring in those water bodies where connectivity has been completely broken (e.g. Sibaya, Groenvlei). In systems that have retained tenuous linkages with the sea (e.g., Verlorenvlei, Wilderness lakes), elements of the marine fish assemblage have persisted, especially the presence of facultative catadromous species. Freshwater fish diversity in coastal lakes and lagoons is a function of historical and present biogeography and salinity. From a freshwater biogeography perspective, the inflowing rivers of the four temperate systems reviewed here contain three or fewer native freshwater fishes, while the subtropical lakes that are fed by river systems contain up to 40 freshwater fish species. Thus, the significantly higher fish species diversity in subtropical versus temperate coastal lakes and lagoons comes as no surprise. Fish species diversity has been increased further in some systems (e.g., Groenvlei) by alien fish introductions. However, the impacts of fish introductions and translocations have not been studied in the coastal lakes and lagoons of South Africa. In these closed systems, it is probable that predation impacts on small estuarine fishes are significant. The recent alien fish introductions is an example of the growing threats to these systems during the Anthropocene, a period when human activities have had significant negative impacts and show potential to match the changes recorded during the entire Holocene.

Grey mullet (Mugilidae) as possible indicators of global warming in South African estuaries and coastal waters.

The grey mullet usually occur in large numbers and biomass in the estuaries of all three South African biogeographic regions, thus making it an ideal family to use in terms of possibly acting as an environmental indicator of global warming. In this analysis the relative estuarine abundance of the dominant three groups of mugilids, namely tropical, warm-water and cool-water endemics, were related to sea surface coastal temperatures. The study suggests a strong link between temperature and the distribution and abundance of the three mullet groups within estuaries and indicates the potential of this family to act as an indicator for future climate change within these systems and adjacent coastal waters.

Does boat traffic cause displacement of fish in estuaries?

Estuaries are increasingly under threat from a variety of human impacts. Recreational and commercial boat traffic in urban areas may represent a significant disturbance to fish populations and have particularly adverse effects in spatially restricted systems such as estuaries. We examined the effects of passing boats on the abundance of different sized fish within the main navigation channel of an estuary using high resolution sonar (DIDSON). Both the smallest (100-300 mm) and largest (>501 mm) size classes had no change in their abundance following the passage of boats. However, a decrease in abundance of mid-sized fish (301-500 mm) occurred following the passage of boats. This displacement may be attributed to a number of factors including noise, bubbles and the rapidly approaching object of the boat itself. In highly urbanised estuarine systems, regular displacement by boat traffic has the potential to have major negative population level effects on fish assemblages.