River flow, zooplankton and dominant zooplanktivorous fish dynamics in a warm-temperate South African estuary.

The possible links between river flow, zooplankton abundance and the responses of zooplanktivorous fishes to physico-chemical and food resource changes are assessed. To this end, the seasonal abundance, distribution and diet of the estuarine round-herring Gilchristella aestuaria and Cape silverside Atherina breviceps were studied in the Kariega Estuary. Spatio-temporal differences were determined for selected physico-chemical variables, zooplankton abundance and zooplanktivorous fish abundance and distribution. Results indicated that, following a river flood event in winter (>30 m3  s-1 ), altered physico-chemical conditions occurred throughout the estuary and depressed zooplankton stocks. Abundance of G. aestuaria was highest in spring, with this species dominant in the upper and middle zones of the estuary, while A. breviceps was dominant in summer and preferred the middle and lower zones. The catch per unit of effort of both zooplanktivores also declined significantly following the flooding, thus suggesting that these fishes are reliant on zooplankton as a primary food source for healthy populations. Copepods dominated the stomach contents of both fish species, indicating a potential for strong interspecific competition for food, particularly in the middle reaches. Temporal differences were evident in dietary overlap between the two zooplanktivorous fish species and were correlated with river flow, zooplankton availability and fish distribution. The findings of this study emphasize the close trophic linkages between zooplankton and zooplanktivorous fishes under changing estuarine environmental conditions, particularly river flow and provide important baseline information for similar studies elsewhere in South Africa and the rest of the world.

Biomass and productivity of fishes in estuaries: a South African case study.

Estuaries are well known for their role as nutrient and detrital sinks that stimulate high levels of both primary and secondary production which, in turn, support a large biomass of fishes per unit area. This study reviews available information on coastal fish biomasses (g m-2 wet mass) and productivity (g m-2 wet mass year-1 ) in order to place South African data on these topics into a global perspective. Using biogeographic fish productivity estimates, together with estuarine water area, the approximate annual teleost production in South African estuaries was calculated at 585, 1706 and 13 904 t in the cool temperate, warm temperate and subtropical regions, respectively. Total annual fish production in estuaries on the subcontinent is conservatively estimated at 16 195 t, but this figure is likely to fluctuate widely, depending on recruitment success and annual environmental conditions pertaining to these systems. Approximately 2000 t of fish are estimated to be harvested by fishing activities in South African estuaries each year, which represents c. 12% of annual fish production. Although this figure may appear sustainable, the reality is that there are a few heavily targeted estuary-associated marine species at the top of the food chain that are being overexploited by both anglers and subsistence fishermen. Natural mortalities due to piscivorous fish and bird predation has been estimated at c. 3% of total fish biomass per month in the East Kleinemonde Estuary, but this figure will vary considerably depending on bird abundance and foraging patterns along the coast. In contrast to catches made by the fishermen, piscivorous fishes and birds are targeting mainly juvenile marine fish and small estuarine resident species that are very abundant and generally low down in the food web.

Tidal amplitude and fish abundance in the mouth region of a small estuary.

Using an acoustic underwater camera (Dual Frequency IDentification SONar, DIDSON), the abundance and direction of movement of fishes > 80 mm total length (LT ) in the mouth of a small South African estuary during spring and neap tidal cycles were observed. While the sizes of fishes recorded were consistent across both tide cycles, the number of fishes passing the camera was significantly greater during the smaller neap tides. Schooling behaviour was more pronounced for fishes that were travelling into the estuary compared to fishes swimming towards the ocean.

Modulational instability of co-propagating internal wavetrains under rotation.

Weakly-nonlinear unidirectional long internal waves in a non-rotating frame are well described by the Korteweg-de Vries equation (KdV). Within the KdV framework, all isolated monochromatic wavetrains are stable to modulational instability. However, analysis of a coupled nonlinear Schrödinger equation system (CNLS) has shown that all systems of two co-propagating monochromatic wavetrains in the KdV are modulationally unstable. To take into account the effect of the background rotation of the Earth on long internal waves, this analysis is extended here to derive the CNLS for the rotation-modified KdV, or Ostrovsky, equation. Rotation stabilises wavetrain pairs when the wavelengths of both waves comprising the wavetrains are longer than the linear wave with maximum group velocity. The particular case when the wavetrains have different wavenumbers but the same linear group speed is emphasised.

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.

Proteomic analysis of Frankliniella occidentalis and differentially expressed proteins in response to tomato spotted wilt virus infection.

Tomato spotted wilt virus (TSWV) is transmitted by Frankliniella occidentalis in a persistent propagative manner. Despite the extensive replication of TSWV in midgut and salivary glands, there is little to no pathogenic effect on F. occidentalis. We hypothesize that the first-instar larva (L1) of F. occidentalis mounts a response to TSWV that protects it from pathogenic effects caused by virus infection and replication in various insect tissues. A partial thrips transcriptome was generated using 454-Titanium sequencing of cDNA generated from F. occidentalis exposed to TSWV. Using these sequences, the L1 thrips proteome that resolved on a two-dimensional gel was characterized. Forty-seven percent of the resolved protein spots were identified using the thrips transcriptome. Real-time quantitative reverse transcriptase PCR (RT-PCR) analysis of virus titer in L1 thrips revealed a significant increase in the normalized abundance of TSWV nucleocapsid RNA from 2 to 21 h after a 3-h acquisition access period on virus-infected plant tissue, indicative of infection and accumulation of virus. We compared the proteomes of infected and noninfected L1s to identify proteins that display differential abundances in response to virus. Using four biological replicates, 26 spots containing 37 proteins were significantly altered in response to TSWV. Gene ontology assignments for 32 of these proteins revealed biological roles associated with the infection cycle of other plant- and animal-infecting viruses and antiviral defense responses. Our findings support the hypothesis that L1 thrips display a complex reaction to TSWV infection and provide new insights toward unraveling the molecular basis of this interaction.

Genome size and ploidy of Thysanoptera.

Flow cytometry was used to study the genome sizes and ploidy levels for four thrips species: Franklinothrips orizabensis Johansen (Thysanoptera: Aeolothripidae), Frankliniella occidentalis Pergande, Frankliniella fusca Hinds, and Thrips tabaci Lindeman (Thysanoptera: Thripidae). F. orizabensis males and females had 1C genome sizes of 426 Mb and 422 Mb, respectively. Male and female F. fusca had 1C genome sizes of 392 Mb and 409 Mb, whereas F. occidentalis males and females had smaller 1C genomes that were 345 Mb and 337 Mb, respectively. Male F. orizabensis, F. occidentalis and F. fusca were haploid and females diploid. Five isofemale lines of T. tabaci, initiated from parthenogenetic, thelytokous females and collected from different locations in North Carolina, were included in this study; no males were available. One isofemale line was diploid with a genome size of 1C = 310 Mb, and the other four had a mean genome size of 1C = 482 Mb, which is consistent with evidence from microsatellite data of diploidy and polyploidy, respectively, in these same five thelytokous lines. This is the first study to produce genome size estimates for thysanopteran species, and report polyploidy in T. tabaci populations.

Systematics of the grey mullets (Teleostei: Mugiliformes: Mugilidae): molecular phylogenetic evidence challenges two centuries of morphology-based taxonomy.

The family Mugilidae comprises mainly coastal marine species that are widely distributed in all tropical, subtropical and temperate seas. Mugilid species are generally considered to be ecologically important and they are a major food resource for human populations in certain parts of the world. The taxonomy and systematics of the Mugilidae are still much debated and based primarily on morphological characters. In this study, we provide the first comprehensive molecular systematic account of the Mugilidae using phylogenetic analyses of nucleotide sequence variation at three mitochondrial loci (16S rRNA, cytochrome oxidase I, and cytochrome b) for 257 individuals from 55 currently recognized species. The study covers all 20 mugilid genera currently recognized as being valid. The family comprises seven major lineages that radiated early on from the ancestor to all current forms. All genera that were represented by two species or more, except Cestraeus, turned out to be paraphyletic or polyphyletic. Thus, the present phylogenetic results generally disagree with the current taxonomy at the genus level and imply that the anatomical characters used for the systematics of the Mugilidae may be poorly informative phylogenetically. The present results should provide a sound basis for a taxonomic revision of the mugilid genera. A proportion of the species with large distribution ranges (including Moolgarda seheli, Mugil cephalus and M. curema) appear to consist of cryptic species, thus warranting further taxonomic and genetic work at the infra-generic level.

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.

Effects of altered estuarine submerged macrophyte bed cover on the omnivorous Cape stumpnose Rhabdosargus holubi.

The ecological importance of submerged macrophyte beds to fishes within estuaries was investigated through the example of the ubiquitous Cape stumpnose Rhabdosargus holubi, an omnivorous, vegetation and estuary-dependent species, using stable-isotope techniques and long-term abundance (catch-per-unit-effort) data from the East Kleinemonde Estuary, South Africa. Outputs from a Bayesian mixing model using δ(13) C and δ(15) N signatures indicated that the submerged macrophytes Ruppia cirrhosa and Potamogeton pectinatus were not a primary source of nutrition for R. holubi, confirming previous work that revealed that macrophytes are consumed but not digested. Long-term seine netting data showed reduced abundance of R. holubi during a prolonged period of macrophyte senescence, suggesting that submerged macrophyte habitats provide shelter that reduces mortality (predation risk) and a food-rich foraging area.

Analysis of expressed sequence tags from Maize mosaic rhabdovirus-infected gut tissues of Peregrinus maidis reveals the presence of key components of insect innate immunity.

The corn planthopper, Peregrinus maidis, causes direct feeding damage to plants and transmits Maize mosaic rhabdovirus (MMV) in a persistent-propagative manner. MMV must cross several insect tissue layers for successful transmission to occur, and the gut serves as an important barrier for rhabdovirus transmission. In order to facilitate the identification of proteins that may interact with MMV either by facilitating acquisition or responding to virus infection, we generated and analysed the gut transcriptome of P. maidis. From two normalized cDNA libraries, we generated a P. maidis gut transcriptome composed of 20,771 expressed sequence tags (ESTs). Assembly of the sequences yielded 1860 contigs and 14,032 singletons, and biological roles were assigned to 5793 (36%). Comparison of P. maidis ESTs with other insect amino acid sequences revealed that P. maidis shares greatest sequence similarity with another hemipteran, the brown planthopper Nilaparvata lugens. We identified 202 P. maidis transcripts with putative homology to proteins associated with insect innate immunity, including those implicated in the Toll, Imd, JAK/STAT, Jnk and the small-interfering RNA-mediated pathways. Sequence comparisons between our P. maidis gut EST collection and the currently available National Center for Biotechnology Information EST database collection for Ni. lugens revealed that a pathogen recognition receptor in the Imd pathway, peptidoglycan recognition protein-long class (PGRP-LC), is present in these two members of the family Delphacidae; however, these recognition receptors are lacking in the model hemipteran Acyrthosiphon pisum. In addition, we identified sequences in the P. maidis gut transcriptome that share significant amino acid sequence similarities with the rhabdovirus receptor molecule, acetylcholine receptor (AChR), found in other hosts. This EST analysis sheds new light on immune response pathways in hemipteran guts that will be useful for further dissecting innate defence response pathways to rhabdovirus infection.

Wheat curl mite resistance: interactions of mite feeding with wheat streak mosaic virus infection.

The majority of plant viruses are dependent on arthropod vectors for spread between plants. Wheat streak mosaic virus (family Potyviridae, genus Tritimovirus, WSMV) is transmitted by the wheat curl mite, Aceria tosichella Keifer, and this virus and vector cause extensive yield losses in most major wheat (Triticum aestivum L.)-growing regions of the world. Many cultivars in use are susceptible to this vector-virus complex, and yield losses of 10-99% have been documented. wheat curl mite resistance genes have been identified in goat grass, Aegilops tauschii (Coss) Schmal., and transferred to hexaploid wheat, but very few varieties contain effectively wheat curl mite resistance, due to virulent wheat curl mite populations. However, wheat curl mite resistance remains an effective strategy to reduce losses due to WSMV. The goal of our project was to identify the most effective, reproducible, and rapid method for assessing wheat curl mite resistance. We also wanted to determine whether mite resistance is affected by WSMV infection, because the pathogen and pest commonly occur together. Single and group wheat curl mite infestations produced similar amounts of leaf rolling and folding on wheat curl mite-susceptible wheat varieties that were independent of initial wheat curl mite infestation. This finding will allow accurate, efficient, large-scale screening of wheat germplasm for wheat curl mite resistance by infesting plants with sections of wheat leaf tissue containing mixed stages of wheat curl mite. The wheat curl mite-resistant breeding line 'OK05312' displayed antibiosis (reduced wheat curl mite population development). The effect of WSMV infection on wheat curl mite reproduction was genotype-dependent. Mite populations increased on infected wheat curl mite- and WSMV-susceptible plants compared with uninfected plants, but WSMV infection had no significant effect on wheat curl mite populations on resistant plants. OK05312 is a strong source of wheat curl mite resistance for wheat breeding programs.

An assessment of the size structure, distribution and behaviour of fish populations within a temporarily closed estuary using dual frequency identification sonar (DIDSON).

The fish populations within the East Kleinemonde Estuary, South Africa, were studied using Dual frequency IDentification SONar (DIDSON) during March 2010 which allowed for direct comparisons of the abundance, distribution and behaviour of a wide size range of fishes along the length of the system. Small schooling fishes <100 mm in total length (L(T) ) were most abundant while larger size classes 100-300 and 300-500 mm were less abundant, but evenly distributed longitudinally within the estuary. Fishes <100 mm were most abundant within the mouth region which may reflect the feeding ecology of species that fall into these size classes and the effects of higher turbidities further up the estuary. Large fishes >500 mm were similarly most abundant near the mouth, and fishes of this size being mostly piscivorous, probably showed these distribution patterns due to the location of smaller prey fishes. Differential behaviour was observed among the longitudinal sections, with small fishes forming schools in areas of low turbidity and particularly in the presence of large predatory fishes. Results from this study present a snap-shot of the composition, distribution and behaviour of the ichthyofauna within a small temporarily closed estuary, and also show that abiotic factors and interactions of various size classes can influence their distribution. There is significant potential for DIDSON in future ecological research within these systems.

Analysis of expressed sequence tags for Frankliniella occidentalis, the western flower thrips.

Thrips are members of the insect order Thysanoptera and Frankliniella occidentalis (the western flower thrips) is the most economically important pest within this order. F. occidentalis is both a direct pest of crops and an efficient vector of plant viruses, including Tomato spotted wilt virus (TSWV). Despite the world-wide importance of thrips in agriculture, there is little knowledge of the F. occidentalis genome or gene functions at this time. A normalized cDNA library was constructed from first instar thrips and 13 839 expressed sequence tags (ESTs) were obtained. Our EST data assembled into 894 contigs and 11 806 singletons (12 700 nonredundant sequences). We found that 31% of these sequences had significant similarity (E< or = 10(-10)) to protein sequences in the National Center for Biotechnology Information nonredundant (nr) protein database, and 25% were functionally annotated using Blast 2GO. We identified 74 sequences with putative homology to proteins associated with insect innate immunity. Sixteen sequences had significant similarity to proteins associated with small RNA-mediated gene silencing pathways (RNA interference; RNAi), including the antiviral pathway (short interfering RNA-mediated pathway). Our EST collection provides new sequence resources for characterizing gene functions in F. occidentalis and other thrips species with regards to vital biological processes, studying the mechanism of interactions with the viruses harboured and transmitted by the vector, and identifying new insect gene-centred targets for plant disease and insect control.

The status of fish conservation in South African estuaries.

Estuary-dependent fish species are defined as those taxa whose populations would be adversely affected by the loss of estuarine habitats. Of the 155 species regularly recorded in South African estuaries, only 32 (21%) are completely dependent on these systems, but this figure increases to 103 species (66%) if partially dependent taxa are included in the analysis. The conservation of fishes in estuaries on the subcontinent is threatened by a number of factors, including habitat degradation, disruption of essential ecological processes, hydrological manipulations, environmental pollution, overexploitation due to fishing activities and, more recently, climate change and the effects of introduced aquatic animals. Although major threats to fishes are usually linked to environmental degradation, there is increasing evidence that the stocks of certain fish species are overexploited or collapsed. Fish conservation and fisheries management does not depend on the implementation of a single action, but rather the co-ordination of a detailed plan, often in a multidisciplinary context. Some examples of innovative means of contributing to estuarine fish conservation in a South African context include the determination and implementation of the ecological freshwater requirements for estuaries, the zoning of estuaries for different uses and the recognition that the maintenance of ecological processes are vital to aquatic ecosystem health. Apart from the designation of protected areas, the main direct means of conserving fish species and stocks include habitat conservation, controls over fishing methods, effort, efficiency and seasonality, pollution control and the prevention of artificial manipulation of estuary mouths. Since becoming a democracy in 1994, environmental legislation, policy and institutional arrangements in South Africa have undergone some major changes, which, if fully implemented, will be very positive for fish conservation in estuaries on the subcontinent.

Generation of nonlinear internal waves by flow over topography: Rotational effects.

We use the forced Ostrovsky equation to investigate the generation of internal waves excited by a constant background current flowing over localized topography in the presence of background rotation. As is now well known in the absence of background rotation, the evolution scenarios fall into three cases, namely subcritical, transcritical, and supercritical. Here an analysis of the linearized response divides the waves into steady and unsteady waves. In all three cases, steady waves occur downstream but no steady waves can occur upstream, while unsteady waves can arise upstream only when there is a negative minimum of the group velocity. The regions occupied by the steady and unsteady waves are determined by their respective group velocities. When the background current is increased, the wave number of the steady waves decreases. In addition, the concavity (canyon or sill), the topographic width, and the relative strength of the rotation play an important role in the generation mechanism. Nonlinear effects modulate the wave amplitude and lead to the emergence of coherent wave packets. All these findings are confirmed by numerical simulations.

Micro-estuaries and micro-outlets as incipient estuarine systems - Does size and coastal connectivity count?

Micro-estuaries and micro-outlets are very small coastal systems that share some biotic and abiotic characteristics with larger temporarily closed estuaries, but differ in that the former have small localized catchments and limited connectivity with the marine environment. This multidisciplinary study reviews the first comprehensive investigation into the ecological structure of a subset of southern African micro-estuaries and micro-outlets, elucidating the biotic and abiotic components that characterize these systems. Furthermore, the microsystems (which have an open water area of less than 5 ha and a depth of less than 1 m) are compared with larger estuarine systems in the region, with the former representing one end of a continuum ranging from micro-outlets to large estuarine lakes and bays. In terms of the abiotic regime, the micro-outlets were dominated by oligohaline or freshwater conditions, while the micro-estuaries were mostly mesohaline. Relatively few freshwater-associated microalgae, zooplankton, macrozoobenthos and fish taxa were present in the micro-outlets, while a richer combination of freshwater and estuary-associated taxa dominated the micro-estuaries. Due to their small size and limited periods of abiotic stability, microsystems only serve as intermediate nursery areas for 0+ juveniles of estuary-associated marine fish which, after less than a year, migrate to nearby larger estuarine systems in order to complete their juvenile life stages. When compared to temporarily closed estuaries, predominantly open estuaries, estuarine lakes and estuarine bays, microsystems are depauperate in terms of typical estuarine and marine biota. However, further research is recommended on microsystems in southern Africa and globally, in order to define in greater detail their physico-chemical and biological characteristics.

A soluble form of the Tomato spotted wilt virus (TSWV) glycoprotein G(N) (G(N)-S) inhibits transmission of TSWV by Frankliniella occidentalis.

Tomato spotted wilt virus (TSWV) is an economically important virus that is transmitted in a persistent propagative manner by its thrips vector, Frankliniella occidentalis. Previously, we found that a soluble form of the envelope glycoprotein G(N) (G(N)-S) specifically bound thrips midguts and reduced the amount of detectable virus inside midgut tissues. The aim of this research was to (i) determine if G(N)-S alters TSWV transmission by thrips and, if so, (ii) determine the duration of this effect. In one study, insects were given an acquisition access period (AAP) with G(N)-S mixed with purified virus and individual insects were assayed for transmission. We found that G(N)-S reduced the percent of transmitting adults by eightfold. In a second study, thrips were given an AAP on G(N)-S protein and then placed on TSWV-infected plant material. Individual insects were assayed for transmission over three time intervals of 2 to 3, 4 to 5, and 6 to 7 days post-adult eclosion. We observed a significant reduction in virus transmission that persisted to the same degree throughout the time course. Real-time reverse transcription polymerase chain reaction analysis of virus titer in individual insects revealed that the proportion of thrips infected with virus was reduced threefold when insects were preexposed to the G(N)-S protein as compared to no exposure to protein, and nontransmitters were not infected with virus. These results demonstrate that thrips transmission of a tospovirus can be reduced by exogenous viral glycoprotein.

Natural nutrient enrichment and algal responses in near pristine micro-estuaries and micro-outlets.

Naturally-occurring pristine estuarine ecosystems are rare in modern environments due to anthropogenic encroachment. There are more than 100 outlets around the South African coast arising from streams flowing from small catchments close to the sea. Eight near natural systems were sampled seasonally over the period of a year to acquire baseline information on water quality and chlorophyll a status across a variety of algal guilds (benthic microalgae, phytoplankton and macroalgal cover). Albeit on a much smaller-scale, these systems represent natural surrogates of larger temporarily open/closed estuaries (TOCEs). Inorganic nutrients (ammonium, total oxidized nitrogen and soluble reactive phosphate), phytoplankton and microphytobenthos chlorophyll a, as well as macroagal percentage cover, were measured using standard methods. Algae showed a seasonal trend, with blooms of both micro- and macro-algae occurring during summer, with a dieback recorded in autumn. During summer, only one system had a phytoplankton peak in chlorophyll a above 20µgL-1, while the microphytobenthos concentrations in three of the systems were above 100mgm-2. Summer blooms of green filamentous macroalgae occurred in all four micro-outlets and in one micro-estuary. Using a linear mixed-effects modelling approach, significant drivers for algal growth related to temperature, nutrient conditions, light availability and water residence time, all of which are known to stimulate primary production. The results show that enrichment from natural sources display similar responses from primary producers to mesotrophic and/or eutrophic water bodies, with the exception that they revert to a natural state rather than continue into a degraded state as is the case in artificially enriched systems. This importantly demonstrates how larger temporarily/open closed estuaries, most of which are anthropogenically degraded, might have functioned under a former more balanced state. Some of these larger systems now respond to nutrient enrichment by exhibiting permanent cultural eutrophication.

Whitham modulation theory for the Ostrovsky equation.

This paper derives the Whitham modulation equations for the Ostrovsky equation. The equations are then used to analyse localized cnoidal wavepacket solutions of the Ostrovsky equation in the weak rotation limit. The analysis is split into two main parameter regimes: the Ostrovsky equation with normal dispersion relevant to typical oceanic parameters and the Ostrovsky equation with anomalous dispersion relevant to strongly sheared oceanic flows and other physical systems. For anomalous dispersion a new steady, symmetric cnoidal wavepacket solution is presented. The new wavepacket can be represented as a solution of the modulation equations and an analytical solution for the outer solution of the wavepacket is given. For normal dispersion the modulation equations are used to describe the unsteady finite-amplitude wavepacket solutions produced from the rotation-induced decay of a Korteweg-de Vries solitary wave. Again, an analytical solution for the outer solution can be given. The centre of the wavepacket closely approximates a train of solitary waves with the results suggesting that the unsteady wavepacket is a localized, modulated cnoidal wavetrain. The formation of wavepackets from solitary wave initial conditions is considered, contrasting the rapid formation of the packets in anomalous dispersion with the slower formation of unsteady packets under normal dispersion.