The effects of size-selective fisheries on the stock dynamics of and sperm limitation in sex-changing fish

Fisheries models have traditionally focused on patterns of growth, fecundity, and survival of fish. However, reproductive rates are the outcome of a variety of interconnected factors such as life-history strategies, mating patterns, population sex ratio, social interactions, and individual fecundity and fertility. Behaviorally appropriate models are necessary to understand stock dynamics and predict the success of management strategies. Protogynous sex-changing fish present a challenge for management because size-selective fisheries can drastically reduce reproductive rates. We present a general framework using an individual-based simulation model to determine the effect of life-history pattern, sperm production, mating system, and management strategy on stock dynamics. We apply this general approach to the specific question of how size-selective fisheries that remove mainly males will impact the stock dynamics of a protogynous population with fixed sex change compared to an otherwise identical dioecious population. In this dioecious population, we kept all aspects of the stock constant except for the pattern of sex determination (i.e. whether the species changes sex or is dioecious). Protogynous stocks with fixed sex change are predicted to be very sensitive to the size-selective fishing pattern. If all male size classes are fished, protogynous populations are predicted to crash even at relatively low fishing mortality. When some male size classes escape fishing, we predict that the mean population size of sex-changing stocks will decrease proportionally less than the mean population size of dioecious species experiencing the same fishing mortality. For protogynous species, spawning-per-recruit measures that ignore fertilization rates are not good indicators of the impact of fishing on the population. Decreased mating aggregation size is predicted to lead to an increased effect of sperm limitation at constant fishing mortality and effort. Marine protected areas have the potential to mitigate some effects of fishing on sperm limitation in sex-changing populations.

Latitudinal and seasonal egg-size variation of the anchoveta (Engraulis ringens) off the Chilean coast

The anchoveta Engraulis ringens is widely distributed along the eastern South Pacific (from 4° to 42°S; Serra et al., 1979) and it has also supported one of the largest fisheries of the world over the last four decades. However, there are few interpopulation comparisons for either the adult or the younger stages. Reproductive traits, such as fecundity or spawning season length, are known to vary with latitude for some fish species (Blaxter and Hunter, 1982; Conover, 1990; Fleming and Gross, 1990; Castro and Cowen, 1991), and latitudinal trends for some early life history traits, such as egg size and larval growth rates, have been reported for others clupeiforms and other fishes (Blaxter and Hempel, 1963; Ciechomski, 1973; Imai and Tanaka, 1987, Conover 1990, Houde 1989). However, there is no published information on potential latitudinal trends during the adult or the early life history of the anchoveta, even though this type of information may help in understanding recruitment variability, especially during recurring large scale events (such as El Niño or La Niña) that affect the entire species range.

Diet shifts of juvenile red snapper (Lutjanus campechanus) with changes in habitat and fish size

We examined the diets and habitat shift of juvenile red snapper (Lutjanus campechanus) in the northeast Gulf of Mexico. Fish were collected from open sand-mud habitat (little to no relief), and artificial reef habitat (1-m3 concrete or PVC blocks), from June 1993 through December 1994. In 1994, fish settled over open habitat from June to September, as shown by trawl collections, then began shifting to reef habitat — a shift that was almost completed by December as observed by SCUBA visual surveys. Stomachs were examined from 1639 red snapper that ranged in size from 18.0 to 280.0 mm SL. Of these, 850 fish had empty stomachs, and 346 fish from open habitat and 443 fish from reef habitat contained prey. Prey were identified to the lowest possible taxon and quantified by volumetric measurement. Specific volume of particular prey taxa were calculated by dividing prey volume by individual fish weight. Red snapper shifted diets with increasing size. Small red snapper (<60 mm SL) fed mostly on chaetognaths, copepods, shrimp, and squid. Large red snapper (60–280 mm SL) shifted feeding to fish prey, greater amounts of squid and crabs, and continued feeding on shrimp. We compared red snapper diets for overlapping size classes (70–160 mm SL) of fish that were collected from both habitats (Bray-Curtis dissimilarity index and multidimensional scaling analysis). Red snapper diets separated by habitat type rather than fish size for the size ranges that overlapped habitats. These diet shifts were attributed to feeding more on reef prey than on open-water prey. Thus, the shift in habitat shown by juvenile red snapper was reflected in their diet and suggested differential habitat values based not just on predation refuge but food resources as well.

A method to improve size estimates of walleye pollock (Theragra chalcogramma) Atka mackerel (Pleurogrammus monopterygius) consumed by pinnipeds: digestion correction factors applied to bones and otoliths recovered in scats

The lengths of otoliths and other skeletal structures recovered from the scats of pinnipeds, such as Steller sea lions (Eumetopias jubatus), correlate with body size and can be used to estimate the length of prey consumed. Unfortunately, otoliths are often found in too few scats or are too digested to usefully estimate prey size. Alternative diagnostic bones are frequently recovered, but few bone-size to prey-size correlations exist and bones are also reduced in size by various degrees owing to digestion. To prevent underestimates in prey sizes consumed techniques are required to account for the degree of digestion of alternative bones prior to estimating prey size. We developed a method (using defined criteria and photo-reference material) to assign the degree of digestion for key cranial structures of two prey species: walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). The method grades each structure into one of three condition categories; good, fair or poor. We also conducted feeding trials with captive Steller sea lions, feeding both fish species to determine the extent of erosion of each structure and to derive condition-specific digestion correction factors to reconstruct the original sizes of the structures consumed. In general, larger structures were relatively more digested than smaller ones. Mean size reduction varied between different types of structures (3.3−26.3%), but was not influenced by the size of the prey consumed. Results from the observations and experiments were combined to be able to reconstruct the size of prey consumed by sea lions and other pinnipeds. The proposed method has four steps: 1) measure the recovered structures and grade the extent of digestion by using defined criteria and photo-reference collection; 2) exclude structures graded in poor condition; 3) multiply measurements of structures in good and fair condition by their appropriate digestion correction factors to derive their original size; and 4) calculate the size of prey from allometric regressions relating corrected structure measurements to body lengths. This technique can be readily applied to piscivore dietary studies that use hard remains of fish.

Temporal changes in population density, fecundity, and egg size of the Hawaiian spiny lobster (Panulirus marginatus) at Necker Bank, Northwestern Hawaiian Islands

Fecundity (F, number of brooded eggs) and egg size were estimated for Hawaiian spiny lobster (Panulirus marginatus) at Necker Bank, North-western Hawaiian Islands (NWHI), in June 1999, and compared with previous (1978–81, 1991) estimates. Fecundity in 1999 was best described by the power equations F = 7.995 CL 2.4017, where CL is carapace length in mm (r2=0.900), and F = 5.174 TW 2.758, where TW is tail width in mm (r2=0.889) (both n=40; P< 0.001). Based on a log-linear model ANCOVA, size-specific fecundity in 1999 was 18% greater than in 1991, which in turn was 16% greater than during 1978–81. The additional increase in size-specific fecundity observed in 1999 is interpreted as evidence for further compensatory response to decreased lobster densities and increased per capita food resources that have resulted either from natural cyclic declines in productivity, high levels of harvest by the commercial lobster trap fishery, or both.

Dynamics of bigeye (Thunnus obesus) and yellowfin (T. albacares) tuna in Hawaii’s pelagic fisheries: analysis of tagging data with a bulk transfer model incorporating size-specific attrition

Tag release and recapture data of bigeye (Thunnus obesus) and yellowfin tuna (T. albacares) from the Hawaii Tuna Tagging Project (HTTP) were analyzed with a bulk transfer model incorporating size-specific attrition to infer population dynamics and transfer rates between various fishery components. For both species, the transfer rate estimates from the offshore handline fishery areas to the longline fishery area were higher than the estimates of transfer from those same areas into the inshore fishery areas. Natural and fishing mortality rates were estimated over three size classes: yellowfin 20–45, 46–55, and ≥56 cm and bigeye 29–55, 56–70, and ≥71 cm. For both species, the estimates of natural mortality were highest in the smallest size class. For bigeye tuna, the estimates decreased with increasing size and for yellowfin tuna there was a slight increase in the largest size class. In the Cross Seamount fishery, the fishing mortality rate of bigeye tuna was similar for all three size classes and represented roughly 12% of the gross attrition rate (includes fishing and natural mortality and emigration rates). For yellowfin tuna, fishing mortality ranged between 7% and 30%, the highest being in the medium size class. For both species, the overall attrition rate from the entire fishery area was nearly the same. However, in the specific case of the Cross Seamount fishery, the attrition rate for yellowfin tuna was roughly twice that for bigeye. This result indicates that bigeye tuna are more resident at the Seamount than yellowfin tuna, and larger bigeye tunas tend to reside longer than smaller individuals. This may result in larger fish being more vulnerable to capture in the Seamount fishery. The relatively low level of exchange between the Sea-mount and the inshore and longline fisheries suggests that the fishing activity at the Seamount need not be of great management concern for either species. However, given that the current exploitation rates are considered moderate (10–30%), and that Seamount aggregations of yellowfin and bigeye tuna are highly vulnerable to low-cost gear types, it is recommended that further increases in fishing effort for these species be monitored at Cross Seamount.

The relative value of different estuarine nursery areas in North Carolina for transient juvenile marine fishes

Offshore winter-spawned fishes dominate the nekton of south-eastern United States estuaries. Their juveniles reside for several months in shallow, soft bottom estuarine creeks and bays called primary nursery areas. Despite similarity in many nursery characteristics, there is, between and within species, variability in the occupation of these habitats. Whether all occupied habitats are equally valuable to individuals of the same species or whether most recruiting juveniles end up in the best habitats is not known. If nursery quality varies, then factors controlling variation in pre-settlement fish distribution are important to year-class success. If nursery areas have similar values, interannual variation in distribution across nursery creeks should have less effect on population sizes or production. I used early nursery period age-specific growth and mortality rates of spot (Leiostomus xanthurus) and Atlantic croaker (Micropogonias undulatus)—two dominant estuarine fishes—to assess relative habitat quality across a wide variety of nursery conditions, assuming that fish growth and mortality rates were direct reflections of overall physical and biological conditions in the nurseries. I tested the hypothesis that habitat quality varies for these fishes by comparing growth and mortality rates and distribution patterns across a wide range of typical nursery habitats at extreme ends of two systems. Juvenile spot and Atlantic croaker were collected from 10 creeks in the Cape Fear River estuary and from 18 creeks in the Pamlico Sound system, North Carolina, during the 1987 recruitment season (mid-March–mid-June). Sampled creeks were similar in size, depth, and substrates but varied in salinities, tidal regimes, and distances from inlets. Spot was widely distributed among all the estuarine creeks, but was least abundant in the creeks in middle reaches of both systems. Atlantic croaker occurred in the greatest abundance in oligohaline creeks of both systems. Instantaneous growth rates derived from daily otolith ages were generally similar for all creeks and for both species, except that spot exhibited a short-term growth depression in the upriver Pamlico system creeks—perhaps the result of the long migration distance of this species to this area. Spot and Atlantic croaker from upriver oligohaline creeks exhibited lower mortality rates than fish from downstream polyhaline creeks. These results indicated that even though growth was similar at the ends of the estuaries, the upstream habitats provided conditions that may optimize fitness through improved survival.

Diet composition of large striped bass (Morone saxatilis) in Chesapeake Bay

Large (>458 mm) striped bass (Morone saxatilis) are dominant predators in Chesapeake Bay. In recent years, the Chesapeake Bay stock of striped bass has increased dramatically, raising concerns about their predatory impact and their forage requirements. In response to these concerns and the need for more recent ecological studies, this investigation was conducted to characterize feeding habits of large striped bass in Chesapeake Bay. Stomach contents from 1225 striped bass from 458 to 1151 mm TL were examined in the spring and fall of 1997 and 1998. Striped bass consumed 52 different species of vertebrates and invertebrates; however, only a few species of clupeoid and sciaenid fishes dominated diets across both the seasons and size ranges of striped bass examined. Of finfish species, menhaden (Brevoortia tyrannus) was the dominant prey in most areas and gizzard shad (Dorosoma cepedianum) replaced menhaden in importance in lower salinity waters. Spot (Leiostomus xanthurus) and other sciaenid fishes and anadromous herrings (Alosa spp.) also contibuted large percentages of striped bass diet. Although pelagic schooling fishes formed the majority of the diet, benthic fishes contributed a higher percentage to the diet than in previous studies of striped bass diet composition.

Estimation of bycatch in shrimp trawl fisheries: a comparison of estimation methods using field data and simulated data

Bycatch, or the incidental catch of nontarget organisms during fi shing operations, is a major issue in U.S. shrimp trawl fisheries. Because bycatch is typically discarded at sea, total bycatch is usually estimated by extrapolating from an observed bycatch sample to the entire fleet with either mean-per-unit or ratio estimators. Using both field observations of commercial shrimp trawlers and computer simulations, I compared five methods for generating bycatch estimates that were used in past studies, a mean-per-unit estimator and four forms of the ratio estimator, respectively: 1) the mean fish catch per unit of effort, where unit effort was a proxy for sample size, 2) the mean of the individual fish to shrimp ratios, 3) the ratio of mean fish catch to mean shrimp catch, 4) the mean of the ratios of fish catch per time fished (a variable measure of effort), and 5) the ratio of mean fish catch per mean time fished. For field data, different methods used to estimate bycatch of Atlantic croaker, spot, and weakfish yielded extremely different results, with no discernible pattern in the estimates by method, geographic region, or species. Simulated fishing fleets were used to compare bycatch estimated by the fi ve methods with “actual” (simulated) bycatch. Simulations were conducted by using both normal and delta lognormal distributions of fish and shrimp and employed a range of values for several parameters, including mean catches of fish and shrimp, variability in the catches of fish and shrimp, variability in fishing effort, number of observations, and correlations between fish and shrimp catches. Results indicated that only the mean per unit estimators provided statistically unbiased estimates, while all other methods overestimated bycatch. The mean of the individual fish to shrimp ratios, the method used in the South Atlantic Bight before the 1990s, gave the most biased estimates. Because of the statistically significant two- and 3-way interactions among parameters, it is unlikely that estimates generated by one method can be converted or corrected to estimates made by another method: therefore bycatch estimates obtained with different methods should not be compared directly.

Does the size of subsamples taken from multispecies trawl catches affect estimates of catch composition and abundance?

Although subsampling is a common method for describing the composition of large and diverse trawl catches, the accuracy of these techniques is often unknown. We determined the sampling errors generated from estimating the percentage of the total number of species recorded in catches, as well as the abundance of each species, at each increase in the proportion of the sorted catch. We completely partitioned twenty prawn trawl catches from tropical northern Australia into subsamples of about 10 kg each. All subsamples were then sorted, and species numbers recorded. Catch weights ranged from 71 to 445 kg, and the number of fish species in trawls ranged from 60 to 138, and invertebrate species from 18 to 63. Almost 70% of the species recorded in catches were “rare” in subsamples (less than one individual per 10 kg subsample or less than one in every 389 individuals). A matrix was used to show the increase in the total number of species that were recorded in each catch as the percentage of the sorted catch increased. Simulation modelling showed that sorting small subsamples (about 10% of catch weights) identified about 50% of the total number of species caught in a trawl. Larger subsamples (50% of catch weight on average) identified about 80% of the total species caught in a trawl. The accuracy of estimating the abundance of each species also increased with increasing subsample size. For the “rare” species, sampling error was around 80% after sorting 10% of catch weight and was just less than 50% after 40% of catch weight had been sorted. For the “abundant” species (five or more individuals per 10 kg subsample or five or more in every 389 individuals), sampling error was around 25% after sorting 10% of catch weight, but was reduced to around 10% after 40% of catch weight had been sorted.