Spatio-temporal patterns in the diversity of demersal fish communities off the south coast of South Africa

The diversity of ecological communities has been the focus of many studies. Because biodiversity provides several indicators used in an Ecosystem Approach to Fisheries (EAF) to track changes in fish communities, we investigated the spatial and temporal patterns in the diversity of some demersal fish communities subjected to varying fishing pressure. Depth and catch rate were the most important predictors in explaining changes in diversity followed by longitude and survey year. Diversity, as measured by the various indices except for taxonomic distinctness (∆*), initially declined with increasing depth to about a depth of 80 m, then increased to about 150 m after which it declined. Taxonomic distinctness index (∆*) showed an increase in the taxonomic heterogeneity of the demersal community below the 300-m isobath. Diversity remained relatively constant with increase in longitude to around 24°E (which has the lowest diversity) after which it increased. The assessment of the temporal trend in diversity indicates that survey year has a significant effect on all diversity indices except for ∆*. Diversity increased and dominance declined with time. This may be result of a decline in the abundance of dominant species or an increase in the abundance less dominant species, or a combination of both effects. Multivariate analysis of the set of diversity indices showed three groups of indices: those reflecting species richness (S, Margalef’s d), those measuring mainly taxonomic relatedness (∆*), and those balancing the richness and evenness components of diversity (J′, H′, λ, ∆, Hill’s N1, and Hill’s N2). The relationship between evenness, catch rate, and size was also investigated. Size classes with highest evenness were found to have lowest catch rate and vice versa. This highlights the need to consider the size and trophic level of species when linking diversity to the functioning of ecosystems.     

Comparative persistence of marine fish larvae from pelagic versus demersal eggs off southwestern Nova Scotia,Canada

Two groupings of larval fish were repeatedly identified by principal component analyses of larval densities from four broad-scale surveys during the spring and summer of 1985–1987 off southwestern Nova Scotia, Canada. Larvae originating from pelagic eggs (four species within Gadidae and Pleuronectidae) constituted one group, which were uniformly distributed over the sampling area with densities not correlated with bathymetry, although nearly all spawning occurs on the shallow western cap of Browns Bank, 100 km offshore. Larvae from demersal eggs (five species within Pholidae, Stichaeidae, Cottidae, Agonidae) constituted the second group, which dominated the shallow-water environments both inshore and on Browns Bank. Lower patchiness indices were evident amongst larvae from pelagic eggs in small and large sampling-gear collections (average 3.4 and 3.1, respectively) compared to fish hatching from demersal eggs (average 5.1 and 4.6). Fine-scale nearshore surveys over a 5 wk period in 1987 also showed that larvae of demersal eggs had a less variable distribution along an inshoreoffshore transect. Larvae from demersal eggs appear spatially persistent through the release of well-developed larvae from non-drifting eggs. These conclusions are consistent with other studies over a range of spatial scales in temperate and tropical environments, demonstrating that single-species models of larval dispersal are inadequate to account for the distributional patterns of larval fish in general.     

Diets of the demersal fishes on the shelf off Iwate,northern Japan

Diets of demersal fishes were determined on the shelf (ca. 130 m deep) off Iwate, Japan. Samples were taken in three different types of habitat, an artificial reef (AR) site, a natural reef (NR) site, and sandymud bottom (SB) site, from May 1987 to September 1991, mostly every two months. A total of 67 prey items were recognized in the stomachs of 45 predator fish species. The most important preys were the pelagic fishes Sardinops melanostictus and Engraulis japonicus, which comprised 37% wet wt of the overall stomach contents. The percentage of pelagic fishes was highest at AR site, where fish density was highest. The dominant ten species could be divided into five feeding types. The pelagic fish feeders Physiculus maximowiczi and Gadus macrocephalus fed mainly on S. melanostictus. The dietary breadth of P. maximowiczi was wide, while that of Gadus macrocephalus was narrow. The pelagic crustacean feeder Theragra chalcogramma mostly consumed Themisto japonica and euphausiids and showed the least dietary overlap with other fishes. Benthic fish feeders were Hemitripterus villosus and Liparis tanakai. The benthic crustacean feeders Alcichthys alcicornis and Hexagrammos otakii consumed benthic crustaceans as well as pelagic and benthic fishes and showed the largest dietary breadth. The benthic invertebrate feeders Gymnocanthus intermedius, Dexistes rikuzenius and Tanakius kitaharai fed mainly on polychaetes and benthic crustaceans. But Gymnocanthus intermedius consumed a significant proportion of pelagic fishes. Ontogenetic dietary shift was recognized for these fishes. Pelagic fishes were consumed more intensively by larger individuals, especially true of A. alcicornis, Theragra chalcogramma and Gadus macrocephalus. Predominancy of the two most adundant species, P. maximowiczi and A. alcicornis, may be supported by their wide dietary breadth and the significant proportion of pelagic fish in their diets. Interspecific dietary overlap was low in most cases suggesting that food resources were well partitioned, although some high overlap was observed among the pelagic fish feeders, A. alcicornis, and Gymnocanthus intermedius, and among the benthic invertebrate feeders. Interspecific competition seemed more likely in the benthic invertebrate feeders than in the pelagic fish feeders partly because of superabundance of the pelagic prey S. melanostictus.     

Growth of demersal fish species of the Mexican Pacific Ocean

The growth of 24 species of demersal fish caught off the Sinaloa-Nayarit coast and in the Gulf of Tehuantepec (Mexican Pacific Ocean) was determined by means of length-frequency analysis. Fishes constitute an important bycatch of the penaeid shrimp fishery, and 235 species were identified in catches made on research cruises in both areas in 1989 to 1992. Growth rates (cm yr^-1) ranged from 0.13 cm for lutjanids and serranids to 0.95 cm for the carangid Selene peruviana; the other species displayed growth rates within this range. Growth varied seasonally, with minimum growth in spring, and is probably related to seasonal changes in the waters of the area. By means of length-frequency analysis, we determined von Bertalanffy growth parameters for the primary species of this multispecies fishery. Such data is fundamental for the future commercial exploitation of these fishes.     

Effects of periodic hypoxia on distribution of demersal fish and crustaceans

Effects of periodic hypoxia (O₂ < 2 mg l^–1) on distribution of three demersal fish species, spot (Leiostomus xanthurus), hogchoker (Trinectes maculatus) and croaker (Micropogonias undulatus), and of two crustacean species, mantis shrimp (Squilla empusa) and blue crab (Callinectes sapidus), were investigated in the lower York River, Chesapeake Bay, USA. Trawl collections were made in four depth strata (5 to 10, 10 to 14, 14 to 20 and > 20 m) during normoxia and hypoxia from 26 June to 20 October 1989. Three periods with hypoxia in the bottom water (below 10 m depth) occurred in mid-July, early August and early September, each with a duration of 6 to 14 d. The demersal fish and crustaceans studied were all affected by hypoxia, and a general migration from deeper to shallower water took place during July and August. However, when oxygen conditions improved after a hypoxic event all species, exceptS. empusa, returned to the deeper areas. The degree of vertical migration was related to levels of oxygen concentration and varied for the different species.M. undulatus was the most sensitive species to low oxygen, followed byL. xanthurus andC. sapidus. T. maculatus andS. empusa were more tolerant and survived in 14 to 25% oxygen saturation by increasing ventilation rate and, forS. empusa, by also increasing blood pigment (haemocyanin) concentration. Periodic hypoxia driven by the spring-neap tidal cycle may represent a natural phenomenon with which the fishes and crustaceans are in a delicate balance. Areas experiencing periodic short-lived hypoxia may be good nursery grounds for fisheries species, and there is no indication that the habitat value in the study area of lower York River is lessened. However, if eutrophication lengthens the time of hypoxia or brings the system closer to anoxia the system may change and become characteristically stressed. The migratory and physiological responses of these species to hypoxia are good indicators of the severity of oxygen stress and could be used as part of an early warning monitoring system for changes in environmental quality.Contribution of the Virginia Institute of Marine Science     

The effect of predation on artificial reef juvenile demersal fish species

There is a concern that artificial reefs (AR) may act purely as fishing aggregation devices. Predators attracted to ARs can influence the distribution and abundance of prey fish species. Determining the role of predators in AR is important in advancing the understanding of community interactions. This paper documents the effects of predation on fish assemblages of AR located near a coastal lagoon fish nursery. The Dicentrarchus labrax is a very opportunistic species preying on juveniles (0⁺ and 1⁺ age classes) of several demersal fish species on the ARs. Reef prey and sea bass abundance were negatively correlated. The mean numbers of prey per sea bass stomach increased with the increase of reef fish prey abundance, suggesting that predation has a significant influence, resulting in a decrease in prey abundance. Prey mortality (4–48%) of demersal reef fish associated species depends on bass density. Prey selection was related both with prey abundance and vulnerability. Results showed that D. labrax predation on AR-fish associated species can increase prey natural mortality. However, the role of bass predation on the ecological functioning of exploited ARs is not clear. There may be increases in local fishing yields due either to an increase in predator biomass through aggregation of sea bass attracted to ARs or to greater production. In contrast, predation on juveniles of economically important reef fish preys, especially the most frequent and abundant (Boops boops), can contribute to a decrease in recruitment to the fishery. Our results indicate that inter-specific interactions (predator–prey) are important in terms of conservation and management, as well as for the evaluation of the long-term effects of reef deployment. Thus, it is necessary to consider ecological interactions, such as predation, prior to the development and deployment of artificial habitats as a tool for rehabilitation.     

Vertical migration and feeding of Euphausia lucens at two 72 h stations in the southern Benguela upwelling region

Diel vertical migration of Euphausia lucens was studied over 72 h at an inshore and at a midshelf station in the southern Benguela upwelling region during February 1991, using stratified net tows and by employing gut contents (chlorophyll/copepods) as tracers of historic position. Soon after population arrival in the surface layers of the inshore station (abundant chlorophyll), E. lucens became dispersed and redistributed over depth. This was accompanied by a rapid downward transfer of surface-collected chlorophyll. A continuous exchange of individuals between deep and shallow water was subsequently inferred, prior to population descent at around dawn. These data are in agreement with the hunger-satiation hypothesis and were shared by individuals at the midshelf station (low chlorophyll). Interestingly, the major part of the E. lucens population at the midshelf station was confined to deep water throughout 24 h. While this conflicts with the hunger-satiation hypothesis, such behaviour is required to both ensure population maintenance and avoid surface predation within the Benguela region. Although the examination of some recent literature suggests that asynchronous, hunger-satiation behaviours may be common amongst euphausiids in single-species assemblages, responses to the food environment may be constrained by other factors.     

Behavioural inertia places a top marine predator at risk from environmental change in the Benguela upwelling system

In variable environments, organisms are bound to track environmental changes if they are to survive. Most marine mammals and seabirds are colonial, central-place foragers with long-term breeding-site fidelity. When confronted with environmental change, such species are potentially constrained in their ability to respond to these changes. For example, if environmental conditions deteriorate within their limited foraging range, long-lived species favour adult survival and abandon their current breeding effort, which ultimately influences population dynamics. Should poor conditions persist over several seasons, breeding-site fidelity may force animals to continue breeding in low-quality habitats instead of emigrating towards more profitable grounds. We assessed the behavioural response of a site-faithful central-place forager, the Cape gannet Morus capensis, endemic to the Benguela upwelling system, to a rapid shift in the distribution and abundance of its preferred prey, small pelagic shoaling fish. We studied the distribution and the abundance of prey species, and the diet, foraging distribution, foraging effort, energy requirements, and breeding success of gannets at Malgas Island (South Africa) over four consecutive breeding seasons. Facing a rapid depletion of preferred food within their foraging range, Cape gannets initially increased their foraging effort in search of their natural prey. However, as pelagic fish became progressively scarcer, breeding birds resorted to scavenging readily available discards from a nearby demersal fishery. Their chicks cannot survive on such a diet, and during our 4-year study, numbers of breeding birds at the colony decreased by 40% and breeding success of the remaining birds was very low. Such behavioural inflexibility caused numbers of Cape gannets breeding in Namibia to crash by 95% following over-fishing of pelagic fish in the 1970s. In the context of rapid environmental changes, breeding-site fidelity of long-lived species may increase the risk of local or even global extinction, rendering these species particularly vulnerable to global change.     

Distribution patterns,abundance and population dynamics of the euphausiidsNyctiphanes capensis andEuphausia hanseni in the northern Benguela upwelling system

Distribution patterns, population structure and biomass of the euphausiidsNyctiphanes capensis andEuphausia hanseni were examined off the coast of Namibia, southwest Africa, in relation to temperature, depth and season, from data collected on nine surveys from September 1982 to March 1984. High densities ofN. capensis were found in the shallow coastal waters (<200 m), with the biomass of adults ranging from 675 to 5 706 mg dry wt m^–2. For adultE. hanseni, the biomass was an order of magnitude lower, ranging from 65 to 505 mg dry wt m^–2, with most specimens occurring over the shelf break at depths of 200 to 1000 m. These distribution patterns remained relatively constant throughout the year, despite seasonal differences in upwelling events. Both species displayed continuous breeding, with 43 to 82% of the adult femaleE. hanseni being fertilized, while a much lower proportion ofN. capensis females were reproductively active (0.5 to 26%). Different breeding strategies were adopted by these two euphausiid species, withE. hanseni producing frequent broods (14.8 broods in 6 mo) consisting of relatively large eggs which are released into the sea, andN. capensis exhibiting a lower frequency of spawning, with broods consisting of large numbers of relatively small eggs, protected by a brood pouch. These strategies enable both species to maintain high densities throughout the year in a fluctuating physical environment. Growth rate estimated from size-frequency distributions were 0.003 to 0.063 mm d^–1 forN. capensis and 0.077 to 0.083 mm d^–1 forE. hanseni, suggesting an adult lifespan of approximately 6 mo for both species. Maximum sizes were attained in September, withN. capensis reaching a total length of 21 mm (in contrast to all previous studies onN. capensis, where the maximum size recorded was only 13 mm total length) andE. hanseni a total length of 33 mm.     

Effect of prey densities on cannibalism in Cape hake (Merluccius capensis) off Namibia

The preference of large (>60 cm total length) Cape hake (Merluccius capensis Castelnau, 1861) for small conspecifics under natural conditions was analyzed off southern Namibia (S.E. Atlantic) between 1983 and 1989. The results indicate that cannibalism is not related to the density of the small conspecifics nor to the density of alternative prey, but rather that large hake have a dietary preference for small conspecifics. The main consequence of such cannibalism is a lack of density-dependent regulation of the M. capensis population.     

The role of fungi in processing marine organic matter in the upwelling ecosystem off Chile

In a study that spanned from March 2007 through November 2009, we report high fungal biomass and over 90% of extracellular enzymatic activity occurring in the size classes dominated by fungi during periods of high autotrophic biomass in surface waters of the upwelling ecosystem off central-southern Chile (36°30.80′S–73°07.70′W). Fungal biomass in the water column was determined by the abundance of hyphae and was positively correlated with the concentration of the fungal biomarker 18:2ω6. High fungal biomass during active upwelling periods was comparable to that of prokaryotes (bacteria plus archaea) and was associated with an increase in phytoplankton biomass and in extracellular enzymatic hydrolysis in waters from the depth of maximum fluorescence. We show fungi as a new microbial component in the coastal upwelling ecosystem of the Humboldt Current System off central Chile. Our results suggest that the temporal pattern in fungal biomass in the water column during a year cycle is a reflection of their capacity to hydrolyze organic polymers and, in consequence, fungal biomass and activity respond to a seasonal cycle of upwelling in this ecosystem.     

The application of predicted habitat models to investigate the spatial ecology of demersal fish assemblages

Benthic habitats are known to influence the abundance and richness of demersal fish assemblages; however, little is known about how habitat structure and composition influences these distributions at very fine scales. We examined how the benthic environment structures marine fish assemblages using high-resolution bathymetry and accurate predicted benthic habitat maps. Areas characterised by a mosaic of habitat patches supported the highest richness of demersal fishes. A total of 37.4% of the variation in the distribution of the fish assemblage was attributed to 6 significant variables. Depth explained 23.0% of the variation, with the boulders explaining 12.6% and relief 1.4%. The remaining measures (seawhips, light/exposure and solid reef) provided a small (<1.0%) but significant contribution. Identifying components of the benthic environment important in structuring fish assemblages and understanding how they influence the spatial distribution of marine fishes is imperative for better management of demersal fish populations.     

Feeding strategies of some demersal fishes of the continental slope and rise off the Mid-Atlantic Coast of the USA

Stomach contents of 729 fishes comprising 16 species were examined from the continental slope and rise off the Middle Atlantic States of the USA. Two main feeding modes among demersal deep-sea fishes were evident: those feeding primarily on pelagic food items, and those feeding on benthic invertebrates. Both pelagic and benthic predators were euryphagous. Most pelagic predators also fed on the epibenthos. These findings support Dayton and Hessler's (1972) suggestion that benthic predators should have a generalized diet which may be responsible for the high diversity found in the deep-sea infauna. The mesopelagic fauna is an important food source for some demersal fishes on the continental slope. Pelagic prey, which are also important to ecologically dominant demersal fishes on the lower slope and continental rise, may be nutritionally supported by suspended particulate organic matter in a nepheloid layer close to the bottom, and they may exist in much higher concentrations than in the bathypelagic zone above.Virginia Institute of Marine Science Contribution No. 835.     

Carbon and nitrogen exchange between sandy beach clams (Donax serra) and kelp beds in the Benguela coastal upwelling region

Carbon consumption and nitrogen requirements were estimated for populations of the sandy beach bivalve Donax serra on nine beaches of the west coast of South Africa. Subtidal populations composed mainly of adult clams were responsible for the bulk of standing stock (3538 g C m⁻¹), annual carbon consumption (13 444 g C m⁻¹ yr⁻¹), faeces production (6478 g C m⁻¹ yr⁻¹ ) and nitrogen regeneration (2525 g N m⁻¹ yr⁻¹). Kelp detritus, bacteria and kelp consumers' faeces available in the water column surpass several times the carbon and nitrogen requirements of intertidal and subtidal clam populations. Individual Donax serra pop ulations, in turn, may regenerate up to 3.2% of the total nitrogen requirements of all primary producers from kelp beds and 14% of the requirements of phytoplankton. These high standing stocks of clams are presumably supported mainly by organic matter originating from kelp which, in contrast to phytoplankton, is in constant supply and comprises the largest proportion of the annual production of particulate organic matter on this coast. Wide and shallow continental shelves with gentle slopes probably limit the penetration of upwelled waters to the nearshore waters, decreasing the influence of external inputs and increasing the importance of internal flows of nutrients and carbon within the nearshore zone. In this context, sandy beaches, rocky shores and kelp beds may be more closely interlinked compartments of a larger ecosystem encompassing the whole nearshore than traditionally thought. Received: 28 August 1996 / Accepted: 7 October 1996     

Reconstructing the movements of free-ranging demersal fish in the North Sea: a data-matching and simulation method

Demersal fish cannot be readily tracked using data loggers that provide satellite-based or light-based geolocation. Moreover, fish that are highly mobile within the water column cannot readily be located with other methods, such as the tidal location method (TLM). As an alternative, we describe a process that provides estimates of geographic location by simulating movement paths through geographic locations that match temperature and depth data recorded by data loggers. Depths and temperatures recorded by data loggers were compared with a North Sea temperature and depth database to identify all locations with matching data. A movement rate filter was then applied to eliminate spurious locations and simulations of possible movement paths through the remaining positions were used to generate estimates of the likelihood of a particular location having been occupied. The performance of the technique was assessed by reconstructing movement paths of artificial migrations and by using depth and temperature data collected at known locations in the North Sea. Estimates of the positional accuracy and error were comparable to the North Sea TLM. Reconstructions of the migrations of cod tagged and released in the North Sea were successfully achieved with the method. This method has application in defining the movements and migrations of commercial species in any sea area where databases of commonly measured environmental variables are available.     

Comparison of fine-scale acoustic monitoring systems using home range size of a demersal fish

We compared the results from fixed acoustic transmitters and transmitters implanted in lingcod Ophiodon elongatus provided by two fine-scale passive acoustic monitoring systems: the older Vemco^? Radio Acoustic Positioning (VRAP) system and the newer VR2W Positioning System (VPS) with either three or four receivers. The four-receiver VPS method calculated five times more positions of lingcod than VRAP and more than twice as many as the three-receiver VPS. Calculated positions of fixed transmitters were less precise with VRAP than either VPS approach. Measurements of home range for lingcod were similar between the four-receiver VPS and VRAP, which were both greater than the three-receiver VPS. Comparisons varied when lingcod were in/near complex habitats. As new technology develops, it is important to understand how new methods compare to previous methods. This may be important when describing patterns of movement or habitat use in the context of changes in habitat or management efforts.     

Two-way coupling versus one-way forcing of plankton and fish models to predict ecosystem changes in the Benguela

‘End-to-end’ models have been adopted in an attempt to capture more of the processes that influence the ecology of marine ecosystems and to make system wide predictions of the effects of fishing and climate change. Here, we develop an end-to-end model by coupling existing models that describe the dynamics of low (ROMS–N₂P₂Z₂D₂) and high trophic levels (OSMOSE). ROMS–N₂P₂Z₂D₂ is a biogeochemical model representing phytoplankton and zooplankton seasonal dynamics forced by hydrodynamics in the Benguela upwelling ecosystem. OSMOSE is an individual-based model representing the dynamics of several species of fish, linked through opportunistic and size-based trophic interactions. The models are coupled through a two-way size-based predation process. Plankton provides prey for fish, and the effects of predation by fish on the plankton are described by a plankton mortality term that is variable in space and time. Using the end-to-end model, we compare the effects of two-way coupling versus one-way forcing of the fish model with the plankton biomass field. The fish-induced mortality on plankton is temporally variable, in part explained by seasonal changes in fish biomass. Inclusion of two-way feedback affects the seasonal dynamics of plankton groups and usually reduces the amplitude of variation in abundance (top-down effect). Forcing and coupling lead to different predicted food web structures owing to changes in the dominant food chain which is supported by plankton (bottom-up effect). Our comparisons of one-way forcing and two-way coupling show how feedbacks may affect abundance, food web structure and food web function and emphasise the need to critically examine the consequences of different model architectures when seeking to predict the effects of fishing and climate change.     

Economics of harvesting age-structured fish populations

A generic age-structured model is developed to derive analytical results on optimal harvesting. Given two age classes, knife-edge selectivity, and no stock-dependent harvesting cost, the steady state is a unique saddle point. Adding harvesting cost does not alter the uniqueness, given that the utility is linear. Under specific conditions such as nonselective gear, optimal harvesting is proved to be a stationary cycle that represents pulse fishing. Optimal steady states are different if age-structured information is ignored and optimization is based on traditional biomass variables. This implies that the existence of optimal sustainable harvesting depends on age-structured information. Given a specific set of conditions such as low interest rate and knife-edge selectivity, optimal harvesting converges toward a unique saddle point independently of the number of age classes.     

Economics of harvesting age-structured fish populations

A generic age-structured model is developed to derive analytical results on optimal harvesting. Given two age classes, knife-edge selectivity, and no stock-dependent harvesting cost, the steady state is a unique saddle point. Adding harvesting cost does not alter the uniqueness, given that the utility is linear. Under specific conditions such as nonselective gear, optimal harvesting is proved to be a stationary cycle that represents pulse fishing. Optimal steady states are different if age-structured information is ignored and optimization is based on traditional biomass variables. This implies that the existence of optimal sustainable harvesting depends on age-structured information. Given a specific set of conditions such as low interest rate and knife-edge selectivity, optimal harvesting converges toward a unique saddle point independently of the number of age classes.