Structural change in an exploited fish community: a consequence of differential fishing effects on species with contrasting life histories

1. An understanding of the links between life histories and responses to exploitation could provide the basis for predicting shifts in community structure by identifying susceptible species and linking life-history tactics with population dynamics.
2. We examined long-term trends in the abundance of species in the North Sea bottom-dwelling (demersal) fish community. Between 1925 & 1996 changes in species composition led to an increase in mean growth rate, while mean maximum size, age at maturity and size at maturity decreased. The demersal fish community was increasingly heavily fished during this period.
3. Trends in mean life-history characteristics of the community were linked to trends in abundance of component species. An approach based on phylogenetic comparisons was used to examine the differential effects of fishing on individual species with contrasting life histories.
4. Those species that decreased in abundance relative to their nearest relative, matured later at a greater size, grew more slowly towards a greater maximum size and had lower rates of potential population increase. The phylogenetically based analyses demonstrated that trends in community structure could be predicted from the differential responses of related species to fishing.
5. This is the first study to link exploitation responses of an entire community to the life histories of individual species. The results demonstrate that fishing has greater effects on slower growing, larger species with later maturity and lower rates of potential population increase. The comparative approach provides a basis for predicting structural change in other exploited communities.     

Demographic diversity and sustainable fisheries

Fish species are diverse. For example, some exhibit early maturation while others delay maturation, some adopt semelparous reproductive strategies while others are iteroparous, and some are long-lived and others short-lived. The diversity is likely to have profound effects on fish population dynamics, which in turn has implications for fisheries management. In this study, a simple density-dependent stage-structured population model was used to investigate the effect of life history traits on sustainable yield, population resilience, and the coefficient of variation (CV) of the adult abundance. The study showed that semelparous fish can produce very high sustainable yields, near or above 50% of the carrying capacity, whereas long-lived iteroparous fish can produce very low sustainable yields, which are often much less than 10% of the carrying capacity. The difference is not because of different levels of sustainable fishing mortality rate, but because of difference in the sensitivity of the equilibrium abundance to fishing mortality. On the other hand, the resilience of fish stocks increases from delayed maturation to early maturation strategies but remains almost unchanged from semelparous to long-lived iteroparous. The CV of the adult abundance increases with increased fishing mortality, not because more individuals are recruited into the adult stage (as previous speculated), but because the mean abundance is more sensitive to fishing mortality than its standard deviation. The magnitudes of these effects vary depending on the life history strategies of the fish species involved. It is evident that any past high yield of long-lived iteroparous fish is a transient yield level, and future commercial fisheries should focus more on fish that are short-lived (including semelparous species) with high compensatory capacity.     

The consequences of balanced harvesting of fish communities

Balanced harvesting, where species or individuals are exploited in accordance with their productivity, has been proposed as a way to minimize the effects of fishing on marine fish communities and ecosystems. This calls for a thorough examination of the consequences balanced harvesting has on fish community structure and yield. We use a size- and trait-based model that resolves individual interactions through competition and predation to compare balanced harvesting with traditional selective harvesting, which protects juvenile fish from fishing. Four different exploitation patterns, generated by combining selective or unselective harvesting with balanced or unbalanced fishing, are compared. We find that unselective balanced fishing, where individuals are exploited in proportion to their productivity, produces a slightly larger total maximum sustainable yield than the other exploitation patterns and, for a given yield, the least change in the relative biomass composition of the fish community. Because fishing reduces competition, predation and cannibalism within the community, the total maximum sustainable yield is achieved at high exploitation rates. The yield from unselective balanced fishing is dominated by small individuals, whereas selective fishing produces a much higher proportion of large individuals in the yield. Although unselective balanced fishing is predicted to produce the highest total maximum sustainable yield and the lowest impact on trophic structure, it is effectively a fishery predominantly targeting small forage fish.     

Comparison of losses of planktivorous fish by predation and seine-fishing in a lake undergoing long-term biomanipulation

SUMMARY 1. Piscivore stocking at artificially high densities and fishing are the two common approaches to reduce the amount of planktivorous and benthivorous fish in lake biomanipulation programmes. Both measures have advantages and disadvantages, but their relative efficacy has not previously been directly compared.
2. We calculated the average annual catch of roach and bream in a lake undergoing long-term biomanipulation (Feldberger Haussee, Germany) by seining each year between 1992 and 1998. We compared this value with a bioenergetics estimate of annual consumption rates of the dominant cohorts of piscivores, pikeperch and pike, in 1997 and 1998. We also determined species composition and length distribution of prey fish in stomachs of the piscivores.
3. Roach was the dominant prey species of both pikeperch and pike, whereas bream was rarely taken by either piscivorous species. Seining removed on average larger specimens of roach than were found in the stomachs of the piscivores.
4. Based on stocking densities of the piscivores, published mortality rates, and individual consumption rates, feeding of pikeperch and pike on roach exceeded the manual removal of roach by seining by a factor of 4–15 (biomass) in 1997 and 1998.
5. Based on these results, a combination of fishing and piscivore enhancement is recommended. Whereas the stocks of adult roach and bream have to be reduced mainly by fishing, the predation of piscivores should be directed predominantly towards the juvenile zooplanktivorous fish. Therefore, small size-classes of piscivorous fish should be promoted by fisheries management, including stocking and harvest regulations.     

Zooplankton responses to predation by larval bluegill: an enclosure experiment

SUMMARY 1. Larval fish are gape-limited predators that forage on prey of specific sizes, and thus may be expected to differentially affect members of a zooplankton community, possibly altering the size-structure or species composition.
2. I used an enclosure experiment to look at the effect of predation by larval bluegill on the dynamics of two zooplankton communities, one dominated by large-bodied individuals and the other by small-bodied individuals. Enclosures containing these zooplankton received a zero, low, medium, or high density of larval bluegill predators.
3. Increasing larval density had a negative effect on zooplankton abundance and abundance declined similarly in the large-bodied and small-bodied communities.
4. Zooplankton size-structure, as estimated by the length of the average zooplankton, increased and then decreased during the experiment, decreasing faster at higher larval fish densities. When zooplankton size-structure was estimated as the length of the average cladoceran, size-structure declined in the large-bodied but not in the small-bodied community and the greatest decline in size-structure was seen in the medium and high larval density treatments.
5. Ordination of each community using multidimensional scaling (MDS) indicated that the trajectory of change in species composition differed between the presence and absence of larval fish. In both communities, the degree of response by individual taxa depended on the density of bluegill larvae. This effect on zooplankton abundance, size-structure and community composition suggests that larval fish may make an important contribution to zooplankton dynamics in many lakes and ponds.     

Predicting the structure and diversity of young-of-the-year fish assemblages in large rivers

1. Interactions between environmental variables and 0+ fish assemblages in the upper River Garonne (France) were quantified during late August 1995.
2. The abundance and diversity of the fish assemblages in floodplain channels were modelled using Artificial Neural Network (ANN) analysis and nine variables: the abundance of the six dominant species, fish specific richness, overall abundance of 0+ fish and the Shannon index of diversity. Multiple regression analysis was also used to assess ANN performance.
3. Using 596 samples, correlation coefficients ( r adjusted) between observed and estimated values of the nine dependent parameters were all highly significant ( P < 0.01). Expected values from the tested data were significantly related to the observed values. The correlation coefficient between observed and estimated values ( r ) varied from 0.70 to 0.85.
4. The ANN provided a high quality prediction, despite the complex nature of the relationship between microhabitat composition and fish abundance.
5. Garson's algorithm was used to provide the explanatory power needed in ecology when using black-box models. Parameters contained in the models (i.e. weighting) were used to determine the relative contributions of explanatory variables and thus to ascertain the structure of fish communities.     

色林错渔业生产的现状与可持续利用的对策

色林错裸鲤（Gymnocypris selincuoensis)是藏北色林错湖泊中惟一的一种鱼类。本文介绍了色林错渔业的开发利用情况,并对色林错裸鲤最小捕捞年龄、捕捞强度、最小网目以及最佳年捕捞产量进行了探讨。从可持续发展的角度,我们认为对色林错鱼类资源的利用必须以保证现有湖泊生态系统的稳定为核心,在满足维持一个足够数量及年龄结构的繁殖群体的基础上,以获得最佳持续经济利益为目标。以往强调甚至采用的最大持续产量理论在高原极端环境条件下并不能作为鱼类资源利用的追求目标或确定渔产量的标准,而只能作为对其捕捞强度的参考。总的允许渔获量应当根据最适捕捞死亡率F0．2来确定。本文最后提出了色林错湖泊鱼类资源可持续利用的8条具体措施。     

Patterns and prediction of population recovery in marine reserves

Marine reserves (no-take zones) are widely recommended asconservation and fishery management tools. One potential benefitof marine reserves is that they can reduce fishing mortality.This can lead to increases in the abundance of spawners,providing insurance against recruitment failure and maintainingor enhancing yields in fished areas. This paper considers thefactors that influence recovery following marine reserveprotection, describes patterns of recovery in numbers andbiomass, and suggests how recovery rates can be predicted.Population recovery is determined by initial population size, theintrinsic rate of population increase r, and the degree ofcompensation (increases in recruits per spawner as spawnerabundance falls) or depensation (lower than expected recruitmentat low abundance, Allee effect) in the spawner-recruitrelationship. Within a reserve, theoretical recovery rates arefurther modified by metapopulation structure and the success ofindividual recruitment events. Recovery also depends on theextent of reductions in fishing mortality (F) as determined bythe relationship between patterns of movement, migration, anddensity-dependent habitat use (buffer effect) in relation to thesize, shape and location of the reserve. The effects ofreductions in F on population abundance have been calculatedusing a variety of models that incorporate transfer rates betweenthe reserve and fished areas, fishing mortality outside thereserve and life history parameters of the population. Thesemodels give useful indications of increases in production andbiomass (as yield per recruit and spawners per recruitrespectively) due to protection, but do not address recruitment.Many reserves are very small in relation to the geographicalrange of fish or invertebrate populations. In these reserves itmay be impossible to distinguish recovery due to populationgrowth from that due to redistribution. Mean rates of recoverycan be predicted from r, but the methods are data intensive. Thisis ironic when marine reserves are often favoured for managementor conservation in data-poor situations where conventional stockassessment is impossible. In these data-poor situations, it maybe possible to predict recovery rates from very low populationsizes by using maximum body size or age at maturity as simplecorrelates of the intrinsic rate of natural increase.     

Natural fishing experiments in marine reserves 1983 – 1993: roles of life history and fishing intensity in family responses

 This study examined the effect of fishing on the abundance and species richness of families of coral reef fish at two islands (Sumilon and Apo) in the Philippines from 1983 to 1993. Natural fishing experiments occurred in marine reserves at each island, where long term estimates of fishing intensity were available. Responses to fishing were interpreted in terms of life histories of fish. The intensity of fishing and fish life histories were generally good predictors of the differential rates of decline and recovery of abundance in response to fishing. Large predators had vulnerable life histories (low rates of natural mortality, growth and recruitment) and were subjected to high intensity fishing. They declined significantly in density when fished and increased significantly but slowly when protected from fishing. Caesionidae, a family with a life history resilient to fishing (high rates of natural mortality, growth and recruitment) but fished intensively also declined rapidly in abundance when fished. Thus, knowledge of life history alone was insufficient to predict response to fishing. Acanthuridae were fished relatively hard and had a life history of intermediate vulnerability but displayed weak responses to fishing. Thus level of fishing intensity alone was also not sufficient to predict response to fishing. For Chaetodontidae, effects of fishing conformed to expectations based on life history and fishing intensity at one island but not the other. Three families with intermediate vulnerability and subjected to intermediate to light fishing (F. Scaridae, Labridae and Mullidae) displayed predictably weak responses to fishing, or counter-intuitive responses (e.g., increasing in abundance following fishing). These counter-intuitive responses were unlikely to be secondary effects of increase in prey in response to declines of predators. Two lightly-fished families with resilient life histories (F. Pomacentridae, Sub F. Anthiinae) predictably displayed weak numerical responses to fishing except during a period of use of explosives and drive nets. Accepted: 30 June 1998     

Thermal affinity as the dominant factor changing Mediterranean fish abundances

Recent decades have seen profound changes in species abundance and community composition. In the marine environment, the major anthropogenic drivers of change comprise exploitation, invasion by nonindigenous species, and climate change. However, the magnitude of these stressors has been widely debated and we lack empirical estimates of their relative importance. In this study, we focused on Eastern Mediterranean, a region exposed to an invasion of species of Red Sea origin, extreme climate change, and high fishing pressure. We estimated changes in fish abundance using two fish trawl surveys spanning a 20‐year period, and correlated these changes with estimated sensitivity of species to the different stressors. We estimated sensitivity to invasion using the trait similarity between indigenous and nonindigenous species; sensitivity to fishing using a published composite index based on the species’ life‐history; and sensitivity to climate change using species climatic affinity based on occurrence data. Using both a meta‐analytical method and random forest analysis, we found that for shallow‐water species the most important driver of population size changes is sensitivity to climate change. Species with an affinity to warm climates increased in relative abundance and species with an affinity to cold climates decreased suggesting a strong response to warming local sea temperatures over recent decades. This decrease in the abundance of cold‐water‐associated species at the trailing “warm” end of their distribution has been rarely documented. Despite the immense biomass of nonindigenous species and the presumed high fishing pressure, these two latter factors seem to have only a minor role in explaining abundance changes. The decline in abundance of indigenous species of cold‐water origin indicates a future major restructuring of fish communities in the Mediterranean in response to the ongoing warming, with unknown impacts on ecosystem function.     

Fish-Community Response To Protective Liming in Thrush Lake, Minnesota

A protective limestone treatment was applied to an acid-sensitive lake in northeastern Minnesota as part of the Acid Precipitation Mitigation Program. This 6–year study evaluated the impact of that treatment on lakes in the upper Midwest that experience episodes of acid stress but have not lost basic species integrity and community structure. Several changes in the fish community can be directly or indirectly attributed to the addition of 4.6 tonnes of calcium carbonate early in the third year of the study. An almost 30–fold increase in the population of Pimephales promelas(fathead minnow) a year after liming, based on mark-recapture estimates from trap netting and snorkeling, was attributed to a pH increase and a three-fold increase in the calcium concentration of the epilimnion. After the initial increase, the abundance of fathead minnows declined in subsequent years, as did the elevated pH and calcium concentrations. The Salvelimis fontinalis(brook trout) population also increased in the lake following application of limestone, but this was due in part to closing the lake to fishing. An increase in survival of stocked brook trout to age 1+ and an increase in growth of older brook trout after liming were attributed to the increased forage that the fathead minnows provided. Fathead minnows may have also reduced predation pressure on young brook trout by older brook trout. This study demonstrated that liming of a slightly acidic lake did not adversely affect the integrity of the fish community, and in fact may have increased the abundance and biomass of the forage fish community and indirectly increased the survival, abundance, and growth of brook trout.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.     

Arthropod community diversity and trophic structure: a comparison between extremes of plant stress

Abstract.  1. Previous studies have shown that plant stress and plant vigour impact the preference and performance of many insect species. Global climate-change scenarios suggest that some regions such as continental interiors may become increasingly subject to severe drought. In combination, these two issues suggest that drought-driven plant stress may impact insect communities on a landscape scale. While there have been many population studies relating plant stress to the life history of individual herbivore species, far less is known about how plant stress affects entire communities.
2. To study the effect of plant stress on arthropod communities, arthropods were sampled from the canopies of pinyon pines ( Pinus edulis ) growing at sites with a history of chronically high environmental stress (e.g. lower water potentials, soil moisture, and reduced growth rates), and those growing under more favourable conditions. Sampling in these environments yielded >59 000 arthropods, representing 287 species from 14 orders and 80 families, and revealed three major community patterns.
3. First, chronic stress significantly altered community composition. Second, trees growing under high stress supported about 1/10th the number of arthropods, and roughly half the species as trees growing under more favourable conditions. Third, of the 33 abundant herbivore species that exhibited a significantly skewed distribution towards either high- or low-stress trees, 73% were skewed with higher numbers on low-stress trees.
4. The pattern of potentially reduced arthropod diversity and abundance on stressed pines observed in this study may further compound the loss of species resulting from the recent, landscape-scale drought-induced mortality of pines in the southwestern USA.     

Temperature effects on chemical signalling in a predator–prey system

SUMMARY 1. We investigated the effect of temperature on chemical signalling in a predator–prey model system (planktivorous fish and Daphnia galeata ). Life-history changes in Daphnia in response to chemical cues (kairomones) derived from fish have become a paradigm for chemically induced anti-predator defences.
2. As temperature can affect both predator and prey, we carried out two experiments to disentangle these effects. In order to test for temperature effects on the predator, we kept prey at a single temperature and exposed them to kairomones from fish exposed to two different temperatures. Daphnia exhibited a higher intrinsic rate of population increase ( r ) when exposed to fish kairomones produced at high rather than low temperature. Assuming a positive correlation between r (because of an earlier maturation and/or increased clutch sizes) and kairomone concentration, our results suggest that kairomone production increases with rising temperature.
3. In the second experiment, to study the influence of temperature on the prey, Daphnia were kept at two different temperatures and exposed to fish kairomones produced at one constant temperature. We found no interaction between the effects of fish kairomone and temperature on Daphnia life history, suggesting that temperature does not directly alter life-history responses to fish kairomones.
4. Our results suggest that temperature influences Daphnia life history through its effects on fish kairomone concentration, but that temperature does not affect the strength of the response of Daphnia to the presence of fish.     

Interaction effects of fish, nutrients, mixing and sediments on autotrophic picoplankton and algal composition

1. We examined the effects of nutrients, turbulent mixing, mosquitofish, Gambusia affinis Baird and Girard and sediments on algal composition, algal biomass and autotrophic picoplankton (APP) abundance in a 6-week experiment of factorial design in twenty-four 5-m³ outdoor mesocosms during late autumn 1995.
2. Turbulent mixing decreased surface temperature and increased turbidity, which also was increased by the addition of sediments. Total algal biomass was significantly enhanced by nutrients and mixing, and decreased by the sediment treatment. In the mixing × nutrient treatment, algal biomass increased more than expected from the individual effects, while the fish × mixing and mixing × sediment treatments increased algal biomass less than expected.
3.  Cryptomonas (cryptomonad) blooms were observed in the unmixed, high nutrient treatment; Synedra (diatom) blooms were observed in the high nutrient, high sediment treatment; Ulothrix (green algae) blooms were observed in the mixed, high nutrient, low sediment treatment.
4. Eukaryotic APP abundances were increased by sediment addition and by turbulent mixing, and increased synergistically by mixing × sediment and mixing × nutrient interactions. Prokaryotic APP abundances were decreased by nutrient enhancement and by a mixing × nutrient interaction. There were no main effects of fish on APP abundance, but fish were involved in some of the two–way interactions.
5. The large number of significant interaction effects indicates that APP and other phytoplankton are regulated by a complex set of interdependent factors which should be considered simultaneously in studies of phytoplankton population dynamics and community composition.     

Changes in the fish community of the Kpong Headpond, lower Volta River, Ghana after 25 years of impoundment

The Kpong Headpond was the second created on the Volta River after Akosombo Dam, primarily as a source of hydroelectric power generation and potable water supply, and additionally, it has supported some fish production in Ghana since impoundment. The changes in fish community of the Kpong Headpond were studied to provide baseline information for strategies formulation to support the socio-economic development of the reservoir. The study identified changes in the fish community of the reservoir by comparing occurrence, composition, relative abundance and relative importance estimates of fish species, families and trophic groups, from available previous studies in the reservoir. From the collated information all fishes identified in the reservoir were categorised based on occurrence and importance as disappeared, appeared, permanent, declined or important, to show current status. The results indicated that the fish community has experienced a shift in the composition and relative abundance of important species, families and trophic groups in terms of number and weight, while remaining ecologically balanced. Representatives of the families Osteoglossidae, Centropomidae and Characidae have declined while representatives of the families Claroteidae, Cyprinidae and Cichlidae have increased. The aufwuch-detritus and herbivores declined while semi-pelagic omnivores increased resulting in a shift in dominance to benthic and semi pelagic omnivores. The appearance of five species and the disappearance of 25 others indicated a dynamic restructuring of the fish community in the reservoir, as expected. Enforcement of fishing regulations including the use of appropriate gear and fishing methods, fishery access control, promotion of culture-based fisheries and improvement in fisher education are recommended topics for sustainable fisheries in the reservoir.     

Consequences of large-scale processes for the conservation of bird populations

1.  Detailed studies of population ecology are usually carried out in relatively restricted areas in which emigration and immigration play a role. We used a modelling approach to explore the population consequences of such dispersal and applied ideas from our simulations to the conservation of wild birds.
2.  Our spatial model incorporates empirically derived variation in breeding output between habitats, density dependence and dispersal. The outputs indicate that dispersal can have considerable consequences for population abundance and distribution. The abundance of a species within a patch can be markedly affected by the surrounding habitat matrix.
3.  Dispersal between habitats may result in lower population densities at the edge of good quality habitat blocks and could partially explain why some species are restricted to large habitat fragments.
4.  Habitat deterioration may not only lead to population declines within that habitat but also in adjacent habitats of good quality. This may confound studies attempting to diagnose population declines.
5.  Although mobile species have the advantages of colonizing sites within metapopulations, dispersal into poorer quality territories may markedly reduce total populations.
6.  There are two main approaches to conservation: one is to concentrate on establishing and maintaining protected areas, while the other involves conservation of the wider countryside. If dispersal is an important process then protecting only isolated areas may be insufficient to maintain the populations within them.     

When does fishing lead to more fish? Community consequences of bottom trawl fisheries in demersal food webs

Bottom trawls are a globally used fishing gear that physically disturb the seabed and kill non-target organisms, including those that are food for the targeted fish species. There are indications that ensuing changes to the benthic invertebrate community may increase the availability of food and promote growth and even fisheries yield of target fish species. If and how this occurs is the subject of ongoing debate, with evidence both in favour and against. We model the effects of trawling on a simple ecosystem of benthivorous fish and two food populations (benthos), susceptible and resistant to trawling. We show that the ecosystem response to trawling depends on whether the abundance of benthos is top-down or bottom-up controlled. Fishing may result in higher fish abundance, higher (maximum sustainable) yield and increased persistence of fish when the benthos which is the best-quality fish food is also more resistant to trawling. These positive effects occur in bottom-up controlled systems and systems with limited impact of fish feeding on benthos, resembling bottom-up control. Fishing leads to lower yields and fish persistence in all configurations where susceptible benthos are more profitable prey. Our results highlight the importance of mechanistic ecosystem knowledge as a requirement for successful management.