Marine reserve effects on population density and size structure of commonly and rarely exploited limpets in South Africa

Marine reserves are commonly used to conserve living resources, but their effectiveness where policing is difficult is unclear. We compared monthly population density and size structure data collected over 20 months for two rarely and two commonly exploited intertidal limpets inside and outside reserves in South Africa. Densities were greater inside reserves, but significantly so in very few months. Unexpectedly, no significant reserve effect was detected for the territorial Scutellastra longicosta, shown by surveys to be favoured by harvesters. Reserve effects for the rarely exploited S. granularis probably result from indirect effects of higher barnacle cover in reserves where trampling is reduced, while interviews indicated that Cellana capensis is not targeted but large individuals are sometimes misidentified and hence are harvested outside the reserve. There were few reserve effects on mean or mean maximum size. The results indicate a gradient of exploitation among species and sites. One non-reserve site was more heavily exploited than the other, while one reserve experienced more poaching than the other. The effectiveness of marine reserves thus differed between reserves and among species. Exploited limpets were generally larger and at higher densities in reserves, but these effects were rarely statistically significant, indicating that reserve effects are weak compared to natural variability, probably reflecting ineffective policing.     

Comparative salinity tolerances of four siphonariid limpets in relation to habitat restriction of the rare and endangered Siphonaria compressa

The salinity tolerances of four South African species of limpet in the genus Siphonaria were experimentally tested, in the context that one of them, S. compressa, is South Africa's most endangered marine mollusc and is restricted to two lagoonal localities that normally experience little variation in salinity. Its habitat is additionally restricted in that it occurs only on the eelgrass Zostera capensis. The other three species, S. capensis, S. concinna and S. serrata, are abundant and widespread on temperate rocky shores. We hypothesised that S. compressa would have limited tolerance to low salinities, contributing to its absence from otherwise suitable estuaries that experience regular fluctuations of salinity. The percentage survival of the limpets was recorded in salinities ranging from 2 to 35 over that period. Both salinity and species significantly affected the survival of the limpets over the 96 hours. Survival was highest in salinities close to those of normal seawater, and greatly reduced in the hypo saline waters. S. compressa had the lowest salinity tolerance of the four species, its time-to-50% mortality in hyposaline waters being significantly less than that for the other three species, which differed in a manner that could be correlated with their zonation patterns. Restriction to eelgrass, coupled with low salinity tolerance, probably explains why S. compressa is found at only two localities, both of which normally experience salinities close to that of seawater. Even there, mass mortalities of S. compressa have been recorded in association with die-backs of eelgrass after abnormal freshwater flooding or sedimentation—further emphasising the vulnerability of S. compressa to extinction.     

Reproduction in the intertidal limpet Cellana ornata in southern New Zealand

The reproductive cycle of the abundant intertidal limpet Cellana ornata (Dillwyn 1817) was examined over 19 months, covering two breeding periods, on a rocky intertidal platform in southern New Zealand. A gonad index was calculated, and histological sections were used to count the numbers of mature and immature eggs present. Fecundity was estimated using counts of eggs before and after spawning. To determine any variation between sites, the gonad index of limpets at a second site dominated by boulders was examined over 6 months during the second breeding period. C. ornata had a single spawning period annually during summer, with greatest gonad sizes in January‐February. There were significant differences in peak gonad indices between two breeding seasons on the platform. Within the second breeding period, both the timing and magnitude of peak gonad development differed between sites, with lower reproductive output at the boulder site. Fecundity increased with increasing size, but individuals are probably capable of reproducing after their first year on the shore. The life history of this species is discussed in comparison to other limpets in New Zealand.     

Reproductive biology of horse mackerel Trachurus capensis and Trachurus trecae in Namibia and Angola

Harvest control measures for Namibia and Angola are defined using reference points for spawning biomass, which requires knowledge of spawning potential and dynamics of fishery stocks. The reproductive biology of two species of horse mackerel, Trachurus capensis and Trachurus trecae, in these waters was investigated for spawning distribution in time and space. Samples were collected during two scientific cruises conducted in February and July/August 2007 off the coast of Namibia and Angola. A total of 396 fish ovaries was examined to determine the spawning dynamics of the two species. The ovaries were histologically analysed and their oocyte distributions mapped. The length (TL)–weight relationships of the two species differed significantly. The observed development of oocyte recruitment and absence of a hiatus in the oocyte distribution indicated that these species are indeterminate spawners. Analyses of the data from the NatMIRC database (2001–2004) indicated annual changes in gonadosomatic index (GSI) for T. capensis at different latitudes, but these changes were small. GSI also fluctuated between years and areas, but there were no clear spatial or temporal trends, with averages ≤4.0% at all stations. The GSI of both species was not indicative of the histologically determined development of gonad maturity. Oocyte maturation for T. capensis indicated that smaller and younger fish spawn first. Batch fecundity (F) for T. trecae was best described by F = 4 986 TL (cm) – 99 645 (r ² = 0.83). This study provided the first analyses of oocyte development in time and space for the two species in the region.     

Interactions between Zostera capensis,Callianassa kraussi and Upogebia africana: deductions from field surveys in Langebaan Lagoon,South Africa

Callianassid sandprawns and eelgrasses have been postulated to have mutually negative effects on each other. The relationship between Callianassa kraussi and Zostera capensis was investigated in Langebaan Lagoon on the west coast of South Africa by correlative field surveys. Seven facts point to mutually antagonistic interactions between the species: (1) cover of Z. capensis was negatively correlated with density of C. kraussi; (2) Z. capensis was largely confined to the high shore, whereas C. kraussi occurred in unvegetated sandbanks below the eelgrass beds; (3) at sites where Z. capensis was absent, C. kraussi extended farther up the shore than in its presence; (4) C. kraussi reached a larger maximum size in sandbanks than in beds of Z. capensis; (5) immediately below eelgrass beds there was a 'frontier zone' where above-ground Z. capensis was absent but below-ground root-rhizomes appeared to exclude C. kraussi; (6) sediment penetrability was greater in sandbanks than in eelgrass beds; (7) rate of burrowing by C. kraussi was faster in unvegetated sandbanks than in eelgrass beds. Conversely, the density and sizes of the mudprawn Upogebia africana were greater in beds of Z. capensis than in sandflats dominated by C. kraussi. Although this evidence is correlative, it suggests that Z. capensis excludes C. kraussi because it stabilises the sediment and inhibits burrowing, and that C. kraussi prevents establishment of eelgrass beds by destabilising the sediment and suspending material that smothers the eelgrass. In the process, two distinct habitats are created. U. africana may benefit directly from eelgrass because it requires stable sediment for its semi-permanent U-tubes; but also indirectly because Z. capensis excludes C. kraussi, which may disrupt these feeding burrows. At the head of the lagoon where sediments are very fine and compact, U. africana occurs both in and below the eelgrass bed, and coexists with at least low densities of C. kraussi, although Z. capensis and C. kraussi are still negatively correlated.     

Foraging ecology of an endemic shorebird, the African Black Oystercatcher (Haematopus moquini) on the south–east coast of South Africa

We investigated small–medium (1–300 km) scale variation in the foraging ecology of the African Black Oystercatcher during its breeding season, using traditional diet analysis coupled with carbon and nitrogen stable isotope analysis. Fieldwork was conducted between January and March 2006 and 2007, on rocky shores on the south–east coast of South Africa at East London, Kenton and Port Elizabeth. Middens of shelled prey left by adults feeding their chicks were collected from five territories and the abundances of the collected prey on the foraging areas were estimated using quadrats. Blood samples from 45 birds (16 females, 10 males and 19 chicks) and tissues from the predominant prey species on the territory of each breeding pair were collected for isotope analysis. The Manly–Chesson selectivity index revealed that adults feed their chicks preferentially with the limpet Scutellastra cochlear and the Mediterranean mussel Mytilus galloprovincialis, if available. A slight enrichment in the ¹⁵N stable-carbon isotope signature was observed towards the west in both prey and oystercatchers. Differences in isotope signatures between males and females from the same breeding pair indicate sex-related differences in the diet. Both had signatures indicating a mixed diet, but with males exhibiting a signature closer to that of limpets and females closer to that of mussels. In the single case where mussels were rare on the feeding territory, the two members of a pair showed carbon signatures which were identical and very similar to that of limpets. These results indicate dietary partitioning between genders in breeding pairs.     

Seasonally and spatially referenced estimates of recreational shore-angling effort,catch composition,catch rates and total catch in the Goukamma Marine Protected Area,South Africa

A roving creel survey of the recreational shore fishery along the 16.4-km coastline in the Goukamma Marine Protected Area on the south coast of South Africa was conducted from 2009 to 2011. Some 838 patrols were stratified equally among months, areas and years, but intentionally biased towards weekends. Angler densities at Buffalo Bay and Groenvlei were 0.59 and 0.28 anglers km?1, respectively. Weekend densities were double to quadruple weekday densities and fishing during winter was more popular than during summer. Area, habitat and distance to access points explained variation in angler densities. Shannon–Wiener diversity in catches declined from 2.18 in an earlier (1993–2002) survey to 1.79. Although the order of species abundance in the catches remained largely unchanged, blacktail Diplodus capensis dominance increased to 57.3% by number, at the expense of galjoen Dichistius capensis. Habitat explained 27% of the variance in catch composition. The catch per unit effort (CPUE) for the top nine species ranged from 0.19 to 6.35 fish 100-h?1. The CPUE of all species, except spotted grunter Pomadasys commersonnii, declined. Blacktail and galjoen CPUE declined by 17% and 77%, respectively. The total catch estimate was 2 986 fish y?1. Transgressions of size limits were common. The results suggest that the fishery is overexploited and that catch rates are declining.     

Influence of oceanographic fronts and low oxygen on the distribution of ichthyoplankton in the Benguela and southern Angola currents

The study focuses on ichthyoplankton populations in the southern Angola Current, the Angola-Benguela Front and the coastal upwelling area of the northern and central Benguela Current. The horizontal and vertical distributions of eggs and larvae of sardine Sardinops sagax, anchovy Engraulis encrasicolus and horse mackerel Trachurus trachurus capensis were examined in relation to distribution patterns of temperature, salinity and dissolved oxygen. Samples were collected during February–March 2002 using a Bongo net and a multiple opening/closing net in five depth layers (0–30m, 30–60m, 60–100m, 100–140m and 140–220m). Eggs and larvae were restricted mainly to the shelf areas in the Angola-Benguela Font and southern Angola Current waters. Their vertical distributions in the Angola-Benguela Front and northern Benguela upwelling region showed a sharp decline in abundance at about 60m depth. This decline appeared to be correlated to the distribution of dissolved oxygen in the water column It is hypothesised that low oxygen concentrations have a strong impact on the development and survival of the early life-cycle stages of fish, and that recruitment of sardine and other pelagic species relies more on the upward extension of the oxygen minimum layer than was previously thought.     

Biological Evaluation of Marine ­Protected Area: Evidence of Crowding Effect on a Protected Population of Queen Conch in the Caribbean

Abstract This study provides a first evaluation of the biological impact of a marine fishery reserve on the stock of queen conch (Strombus gigas) in the Turks and Caicos Islands. The density and the shell length of the population living in the reserve are compared with those of the individuals living in the surrounding fished areas. The results show that the adult density is six times higher in the reserve than in the fished areas. The shell length analysis shows that both adults and juveniles are significantly smaller in the reserve than in the fished area. This unexpected result suggests the existence of a crowding effect (i. e. a high density‐induced reduction in growth rate) within the ­reserve. It is hypothesised that this crowding effect is due to the superimposition of two factors leading to very high density values in the reserve: (a) the reduced fishing mortality following the creation of the reserve, (b) the existence of natural barriers that impede the emigration of adults outside the reserve. These results are then discussed in relation to current considerations on marine fisheries reserves.     

Depth-stratified density estimates and distribution of the Cape hake Merluccius capensis and M. paradoxus off Namibia deduced from survey data, 1990–1999

In this study demersal survey data for the period 1990–1999 are used to investigate the average distribution of the Cape hake Merluccius capensis and M. paradoxus off Namibia in terms of density and mean length. Further, biomass estimates are compared on the basis of depth and density stratification. The main distribution of M. capensis was north of 27°S and that of M. paradoxus south of 24°S. M. paradoxus was deeper than M. capensis. For both species, average length increased with depth. M. paradoxus expanded its range to the north through the 1990s as its population size (off South Africa and Namibia) increased. In Namibian waters, small M. paradoxus were found only south of 25°S. Mean length of M. capensis increased north of 21°S, largely as a result of decreased numbers of small fish in shallower water. Abundance estimates stratified by depth were no different from those post-stratified on similar densities.     

Spawning strategies and transport of early stages of the two Cape hake species,Merluccius paradoxus and M. capensis,in the southern Benguela upwelling system

Seasonal and short-term variability of environmental parameters influence the spawning strategies of fish species. In this study, the spawning strategies and the transport of early stages of the two Cape hake species off South Africa were investigated. Distribution of eggs and larvae of Merluccius paradoxus and M. capensis was analysed in order to derive more detailed and species-specific information on spawning season, spawning location, and transport of early stages. Samples were collected during three pilot surveys between January and October 2007 and during an extensive survey in September/October 2008 in the southern Benguela upwelling system off South Africa. Eggs and larvae of M. paradoxus were found in greater numbers than those of M. capensis during all surveys. Highest abundances were found from September to October, indicating one spawning peak for M. paradoxus during late austral winter to spring. The western Agulhas Bank was identified as the primary spawning ground, and smaller spawning events occurred on the West Coast. Larvae of both species were mainly distributed in subsurface waters between 25 and 100 m. More than 50% of all larvae caught had a total length between 3 and 4 mm and size increased significantly with decreasing latitude. Merluccius capensis were found closer inshore than M. paradoxus, indicating that early stages of the two species followed separate drift routes. We assume that this distribution pattern most likely evolved from differences in spawning location and phenology. The spawning strategies of M. paradoxus and M. capensis are well adapted to a time-frame of optimal transport conditions favourable for larval survival in the highly variable environment of the southern Benguela upwelling system, but the peak spawning of the two species is separated in time and space.     

Occurrence and distribution of Cape hakes Merluccius capensis Cast. and M. paradoxus Franca in the Cape of Good Hope area

A brief history of research into, and the fishery for, Cape hake is given. From twelve monthly surveys in 1972/73, abundance of the two species (Merluccius capensis and M. paradoxus) at depth was found to vary, larger individuals of both species being found deeper than the smaller ones. Off the Cape of Good Hope, M. capensis was never found deeper than 440 m, and there was significant species overlap between 150 and 440 m. M. paradoxus was numerically dominant except at the shallowest station sampled (150 m). Little overlap of mature specimens of each species was recorded. The research data were applied to commercial trawl statistics, revealing that M. capensis accounted for only 12,9 per cent by mass and 4,8 per cent by number of the commercially caught hake in the years 1955–1974. Such low percentages justify considering the Cape hake fishery as a single-species entity for the purposes of stock assessment and management.     

Marine invasive aliens on South African shores: implications for community structure and tropillc functioning

Up to 12 marine mollusc and three crab species are thought to have been deliberately or accidentally introduced to South Africa. Of these, only the Mediterranean mussel Mytilus galloprovincialis and the European shore crab Carcinus meanas have become invasive. M. galloprovincialis, probably introduced in the late 1970s, is already the dominant mussel throughout the Cape west coast. As a result, indigenous intertidal mussels Aulacomya ater have been largely displaced, mussel standing stock has increased and the upshore limit of mussel beds has become considerably elevated. Implications include competitive exclusion of large limpets but enhanced recruitment of juvenile limpets, increased habitat availability for mussel infauna, reduction of habitat for algal infauna and enhanced food availability for predators, particularly terrestrial species. C. maenas was first recorded from Table Bay Docks in 1983 and currently ranges from Camps Bay to Saldanha Bay. Although it is a voracious predator, it appears unable to colonize wave-swept shores, so is unlikely to displace indigenous crabs, or to severely impact prey species on the open coast. Valuable conservation areas and mariculture sites in sheltered lagoonal areas are, however, threatened by it.     

Environmental effects on the reproduction and fecundity of the introduced calcareous sponge Paraleucilla magna in Rio de Janeiro,Brazil

The calcareous sponge Paraleucilla magna (Porifera, Calcarea) has been the subject of several studies in the last decade. It was first described along the Brazilian coast, where it is considered cryptogenic, and was subsequently found in the Mediterranean, where it is considered invasive. The wide artificial distribution of this species allows us to compare different aspects of the biology of an introduced species in different locations. Here, we analysed the effects of selected environmental parameters on the reproductive dynamics of P. magna in Rio de Janeiro (Brazil) over 18 months and compared our results with those obtained for the same species in the Mediterranean Sea. Specimens were collected monthly and analysed through histological methods. The density of reproductive elements in each month was calculated, and the effects of environmental parameters (photoperiod, precipitation, temperature, phytoplankton and bacterioplankton) were analysed using a regression tree analysis. Paraleucilla magna was reproductive throughout the study period. The densities of the reproductive elements (oocytes, embryos and larvae) showed no seasonality, and this species presented one of the highest reproductive efforts documented to date in the phylum Porifera (99.0 oocytes · mm^?3; 89.0 embryos · mm^?3; 319.0 larvae · mm^?3). The main environmental parameters related to the reproduction of P. magna were temperature, photoperiod and bacterioplankton. Temperature was the main driver associated with the densities of oocytes and embryos, while bacterioplankton was the main driver of larvae (positive relationships). In Rio de Janeiro, larvae were present and continuously released. This strategy is different from that observed in the Mediterranean, where a larger larval output was observed but only during the summer months. Our results show that P. magna is a species with a strong invasive potential, considering its high and continuous reproductive effort. This high fecundity stimulated by high temperatures may be a key factor contributing to the growth of P. magna populations and its invasion of new areas.     

Macroalgal associations of motile epifaunal invertebrate communities on coral reefs

Small grazing motile epifaunal invertebrates play an important ecosystem role on coral reefs, influencing both the abundance and composition of macroalgal communities and acting as a key food source for a range of predatory fishes. The first aim of this study was to investigate the associations between motile epifaunal communities and four common macroalgal species (Lobophora variegata, Dictyota divaricata, Microdictyon marinum and Halimeda opuntia) on fore‐reef environments in the Exuma Cays (Bahamas, wider Caribbean). Secondly, we investigated the implications of the well documented rise of Caribbean macroalgal cover on invertebrate densities by surveying sites inside and outside the Exuma Cays Land and Sea Park (ECLSP), where increases in parrotfish grazing intensity inside the marine reserves have led to reductions in macroalgal cover. Therefore, surveys compared similar reefs with significantly different macrolagal cover. Comparisons between macroalgal species revealed a four to fivefold difference in motile epifaunal densities per unit volume of macroalgae. Post‐hoc tests revealed that this difference was significant only for Lobophora, with no difference observed among the other species. As macroalgae provide both a refuge from predation and a food source for grazing epifauna, the higher densities of epifauna observed in Lobophora may be attributed to either refuge from visual predators through morphological features (high cover of overlapping blades close to the substrate) or lack of palatability for parrotfish grazing, providing a more stable refuge. Our results revealed no significant differences in diversity, density or community structure of motile epifauna per unit volume of macroalgae between sites inside and outside the ECLSP. Since canopy height and invertivore biomass did not vary systematically across reserve boundaries, this suggests that algal cover does not affect the density of epifaunal invertebrates. However, areal cover was consistently higher for all macroalgal species at sites outside the ECLSP than those inside the reserve. Therefore, when scaled by aerial cover of macroalgae, total abundance of epifauna was twofold higher outside the ECLSP. We suggest that the increasing abundance of macroalgae on Caribbean reefs may be having dramatic effects on epifaunal invertebrate populations and potentially their ecological functions.     

Top predators in the Benguela ecosystem — implications of their trophic position

Many top predators in the Benguela ecosystem feed on prey species targeted by commercial fisheries. Their roles as indicators of the state of exploited prey resources, as competitors with commercial fisheries for resources, and as susceptible to impact from commercial fishing on those resources are briefly considered. Trends in the occurrence of anchovy Engraulis capensis and pilchard Sardinops ocellatus in the diet of Cape gannets Morus capensis off South Africa's west coast are related significantly to survey estimates of the abundance of these fish species, and they provide useful confirmation of those estimates. In the 1980s, anchovy decreased in the diet of Cape gannets, but pilchard increased. In both the northern and southern portions of the Benguela system, groundfish were thought to eat most (66–73%) of the total quantity of cephalopods and vertebrates consumed by predators and man in the 1980s. South African fur seals Arctocephalus pusillus pusillus, predatory pelagic fish and man removed roughly equal amounts, with squids, seabirds and cetaceans having a smaller impact. In the 1980s, man and seals removed about two million tons live mass more than in 1930. Indices of the rate of natural mortality of anchovy and pilchard attributable to Cape gannets are not related to biomass of the prey species. That for anchovy was high in 1989 when a poor anchovy year-class was formed. Decreased abundance of anchovy led to poor breeding by Cape cormorants Phalacrocorax capensis in 1989 and 1990. A model linking the Cape cormorant population with anchovy is used to explore the impact of possible exploitation strategies for anchovy on Cape cormorants.     

Impact of climate and hydrodynamics on sole larval immigration towards the Tagus estuary, Portugal

Spawning grounds of the soles, Solea solea and Solea senegalensis, are distant from the estuarine nurseries where juveniles concentrate. Recruitment of these species is highly dependent on the success of the larval migration towards the inshore nursery grounds. Unfavourable climate and hydrodynamic circulation may lead to high mortality rates at this stage. The relation between river drainage, NAO index and the North-South wind component intensity over the three months prior to the end of the estuarine colonization and the densities of S. solea and S. senegalensis in the nursery grounds were investigated for both species based on a discontinuous historical dataset (from 1988 to 2006) for the Tagus estuary. Multiple linear regression models were developed for sole density and environmental data (separately for each species). Results showed that river drainage is positively correlated with juveniles’ densities of both species, possibly due to the existence of chemical cues used by larvae for movement orientation. NAO index and the North-South wind component intensity relations with soles densities were non-significant. It was concluded that the high complexity of the Portuguese upwelling system makes it hard to detect causal relations of the environmental variables tested. The importance of river flow for coastal ecosystems was stressed. Since climate change scenarios predict a strong decrease in rain fall over the Portuguese river basins, as well as a concentrated period of heavy rain in winter, it was hypothesised that future river drainage decrease over much of the year may lead to lower recruitment success for soles, especially for S. senegalensis.     

Population structure and growth of polydorid polychaetes that infest cultured abalone Haliotis midae

Polydorid polychaetes can infest cultured abalone thereby reducing productivity. In order to effectively control these pests, their reproductive biology must be understood. The population dynamics and reproduction of polydorids infesting abalone Haliotis midae from two farms in South Africa is described using a length-based, age-structured model. Shells were infested mainly by introduced Boccardia proboscidea. Polydora hoplura and Dipolydora capensis were also present but in numbers too few to identify factors influencing infestation. At both farms, B. proboscidea lived for a minimum of 12 months. Growth rate, size at maturity, maximum size, infestation intensity, recruitment, percentage of the population brooding and mortality appear to be affected by abalone feeding regime and water temperature, and these factors need to be considered in controlling infestation. Brooders and recruits were present throughout the year, but increased significantly during mid- to late winter/early spring when water temperature and day length increased. Treatment measures should therefore be implemented throughout the year but with increased effort when water temperature increases.     

Kermadec Island giant limpet occurring fossil in New Zealand,and relict distributions in the tropics

A large fossil limpet from the Isolated Hill Limestone (Oligocene) of Mason River, Marlborough, is described as Patella (Scutellastra) kermadecensis aurorae n. subsp., a subspecies of the living Kermadec giant limpet. Thus this limpet is not a primary endemic species at the Kermadec Islands, but a surviving relict population of a species formerly more widespread during the Tertiary. Such relict populations in the Indo‐West Pacific imply that the tropics were not immune from the effects of Pleistocene climatic fluctuations.     

Prevalence of Kudoa thyrsites (Myxozoa) in South African sardine Sardinops sagax,kingklip Genypterus capensis,and Cape hakes Merluccius capensis and M. paradoxus

Prevalence of infection with the myxozoan parasite Kudoa thyrsites is not well documented in sardine Sardinops sagax, the Cape hakes Merluccius capensis and M. paradoxus, and kingklip Genypterus capensis. Quantitative polymerase chain reaction assays were used to determine the prevalence of K. thyrsites in wild-caught S. sagax (n?=?296), Merluccius spp. (n?=?162) and G. capensis (n?=?70), harvested in 2015. Sardinops sagax were harvested from the east, south and west coasts of South Africa, whereas M. capensis, M. paradoxus and G. capensis were taken from the west coast only. Relationships between K. thyrsites infection prevalence and fish capture locality, season, sex and size were determined with Chi-square tests. Prevalence was highest in S. sagax (91%), followed by M. capensis and M. paradoxus (87%) and G. capensis (40%). Prevalence was independent of sex of the host for all fish species. In the case of S. sagax only, the prevalence of infection was independent of locality of capture but depended on season, with prevalence lowest during spring and highest during autumn. No significant relationship (p?>?0.05) was found between infection prevalence and host size for any of the fish species.