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.

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.

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.

Assessment of sampling methods to estimate horseshoe crab (Limulus polyphemus L.) egg density in Delaware Bay

Each spring horseshoe crabs (Limulus polyphemus L.) emerge from Delaware Bay to spawn and deposit their eggs on the foreshore of sandy beaches (Shuster and Botton, 1985; Smith et al., 2002a). From mid-May to early June, migratory shorebirds stopover in Delaware Bay and forage heavily on horseshoe crab eggs that have been transported up onto the beach (Botton et al., 1994; Burger et al., 1997; Tsipoura and Burger, 1999). Thus, estimating the quantity of horseshoe crab eggs in Delaware Bay beaches can be useful for monitoring spawning activity and assessing the amount of forage available to migratory shorebirds.

Reproductive biology of the blue swimmer crab (Portunus pelagicus, Decapoda: Portunidae) in five bodies of water on the west coast of Australia

Portunus pelagicus was collected at regular intervals from two marine embayments and two estuaries on the lower west coast of Australia and from a large embayment located approximately 800 km farther north. The samples were used to obtain data on the reproductive biology of this species in three very different environments. Unlike females, the males show a loosening of the attachment of the abdominal flap to the cephalothorax at a prepubertal rather than a pubertal molt. Males become gonadally mature (spermatophores and seminal fluid present in the medial region of the vas deferentia) at a very similar carapace width (CW) to that at which they achieve morphometric maturity, as reflected by a change in the relative size of the largest cheliped. Logistic curves, derived from the prevalence of mature male P. pelagicus, generally had wider confidence limits with morphometric than with gonadal data. This presumably reflects the fact that the morphometric (allometric) method of classifying a male P. pelagicus as mature employs probabilities and is thus indirect, whereas gonadal structure allows a mature male to be readily identified. However, the very close correspondence between the CW50’s derived for P. pelagicus by the two methods implies that either method can be used for management purposes. Portunus pelagicus attained maturity at a significantly greater size in the large embayment than in the four more southern bodies of water, where water temperatures were lower and the densities of crabs and fishing pressure were greater. As a result of the emigration of mature female P. pelagicus from estuaries, the CW50’s derived by using the prevalence of mature females in estuaries represent overestimates for those populations as a whole. Estimates of the number of egg batches produced in a spawning season ranged from one in small crabs to three in large crabs. These data, together with the batch fecundities of different size crabs, indicate that the estimated number of eggs produced by P. pelagicus during the spawning season ranges from about 78,000 in small crabs (CW=80 mm) to about 1,000,000 in large crabs (CW=180 mm).

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.

Marine fish assemblages associated with fish aggregating devices (FADs): effects of fish removal, FAD size, fouling communities, and prior recruits

Fishes are widely known to aggregate around floating objects, including flotsam and fish aggregating devices (FADs).The numbers and diversity of juvenile fishes that associated with floating objects in the nearshore waters of the eastern tropical Pacific were recording by using FADs as an experimental tool. The effects of fish removal, FAD size, and the presence or absence of a fouling community at the FAD over a period of days, and the presence of prior recruits over a period of hours were evaluated by using a series of experiments. The removal of FAD-associated fish assemblages had a significant effect on the number of the dominant species (Abudefduf troschelii) in the following day’s assemblage compared to FADs where the previous day’s assemblage was undisturbed; there was no experimental effect on combined species totals. Fishes do, however, discriminate among floating objects, forming larger, more species-rich assemblages around large FADs compared to small ones. Fishes also formed larger assemblages around FADs possessing a fouling biota versus FADs without a fouling biota, although this effect was also closely tied to temporal factors. FADs enriched with fish accumulated additional recruits more quickly than FADs that were not enriched with fish and therefore the presence of prior recruits had a strong, positive effect on subsequent recruitment. These results suggest that fish recruitment to floating objects is deliberate rather than haphazard or accidental and they sup-port the hypothesis that flotsam plays a role in the interrelationship between environment and some juvenile fishes. These results are relevant to the use of FADs for fisheries, but emphasize that further research is necessary for applied interests.

Geographic and temporal patterns in size and maturity of the longfin inshore squid (Loligo pealeii) off the northeastern United States

Analysis of 32 years of standardized survey catches (1967–98) indicated differential distribution patterns for the longfin inshore squid (Loligo pealeii) over the northwest Atlantic U.S. continental shelf, by geographic region, depth, season, and time of day. Catches were greatest in the Mid-Atlantic Bight, where there were significantly greater catches in deep water during winter and spring, and in shallow water during autumn. Body size generally increased with depth in all seasons. Large catches of juveniles in shallow waters off southern New England during autumn resulted from inshore spawning observed during late spring and summer; large proportions of juveniles in the Mid-Atlantic Bight during spring suggest that substantial winter spawning also occurs. Few mature squid were caught in survey samples in any season; the majority of these mature squid were captured south of Cape Hatteras during spring. Spawning occurs inshore from late spring to summer and the data suggest that winter spawning occurs primarily south of Cape Hatteras.

Age and size composition, growth rate, reproductive biology, and habitats of the West Australian dhufish (Glaucosoma hebraicum) and their relevance to the management of this species

Samples of the commercially and recreationally important West Australian dhufish (Glaucosoma hebraicum) were obtained from the lower west coast of Australia by a variety of methods. Fish <300 mm TL were caught over flat, hard substrata and low-lying limestone reefs, whereas larger fish were caught over larger limestone and coral reef formations. Maximum total lengths, weights, and ages were 981 mm, 15.3 kg, and 39 years, respectively, for females and 1120 mm, 23.2 kg, and 41 years, respectively, for males. The von Bertalanffy growth curves for females and males were significantly different. The values for L∞, k, and t0 in the von Bertalanffy growth equations were 929 mm, 0.111/year, and –0.141 years, respectively, for females, and 1025 mm, 0.111/year, and –0.052 years, respectively, for males. Preliminary estimates of total mortality indicated that G. hebraicum is now subjected to a level of fishing pressure that must be of concern to fishery managers. Glaucosoma hebraicum, which spawns between November and April and predominantly between December and March, breeds at a wide range of depths and is a multiple spawner. The L50’s for females and males at first maturity, i.e. 301 and 320 mm, respectively, were attained by about the end of the third year of life and are well below the minimum legal length (MLL) of 500 mm. Because females and males did not reach the MLL until the end of their seventh and sixth years of life, respectively, they would have had, on average, the opportunity of spawning during four and three spawning seasons, respectively, before they reached the MLL. However, because G. hebraicum caught in water depths >40 m typically die upon release, a MLL is of limited use for conserving this species. Alternative approaches, such as restricting fishing activity in highly fished areas, reducing daily bag limits for recreational fishermen, introducing quotas or revising specific details of certain commercial hand-line licences (or doing both) are more likely to provide effective conservation measures.

Recruitment season, size, and age of young American eels (Anguilla rostrata) entering an estuary near Beaufort, North Carolina

Net catches from 1985–86 to 1994–95 at Pivers Island, North Carolina, indicated that glass-eel stage American eels (Anguilla rostrata) were recruited to the estuary from November to early May, with peak numbers in January, February, and March. There was no declining trend in recruitment over the years of sampling. Except for one year, there was no clear seasonal decrease in mean length. But shorter glass eels were older than longer glass eels, as judged by age within the glass eel growth zone of the otolith, suggesting that smaller fish took longer to arrive. The mean age of glass eels collected from the lower estuary and a freshwater site 9.5 km upriver differed by 8.4 d (36.2 vs. 44.6, respectively). Outer increments (30–35) of the otolith growth zone of glass eels from North Carolina were significantly wider than corresponding increments of otoliths from New Brunswick. Mean total ages of North Carolina, New Jersey, and New Brunswick elvers were 175.4, 201.2, and 209.3 d, corresponding to mean lengths of 55.9, 60.9, and 58.1 mm TL, respectively. The mean durations of glass-eel growth zones (44.6, 62.3, and 69.8) were in close agreement with those from previous studies, but total ages were not. This suggested that perhaps some finer (leptocephalus stage) increments were not detected by light microscopy, differences occurred in seasonal increment deposition, or absorption of the otolith material may have taken place during metamorphosis, rendering the aging of larvae inaccurate. Judging from the long recruitment period and seasonal uniformity in both mean age and length found in our study, the spawning period of American eels may be somewhat more protracted than previously considered.

Do fluctuations in the somatic growth rate of rock lobster (Jasus lalandii) encompass all size classes? A re-assessment of juvenile growth

Catch rates in the South African rock lobster (Jasus lalandii) fishery declined after 1989 in response to reduced adult somatic growth rates and a consequent reduction in recruitment to the fishable population. Although spatial and temporal trends in adult growth are well described, little is known about how juvenile growth rates have been affected. In our study, growth rates of juvenile rock lobster on Cape Town harbor wall were compared with those recorded at the same site more than 25 years prior to our study, and with those on a nearby natural nursery reef. We found that indices of somatic growth measured during 1996–97 at the harbor wall had declined significantly since 1971–72. Furthermore, growth was slower among juvenile J. lalandii at the harbor wall than those at the natural nursery reef. These results suggest that growth rates of juvenile and adult J. lalandii exhibit similar types of spatiotemporal patterns. Thus, the recent coastwide decline in adult somatic growth rates might also encompass smaller size classes.

Seasonal growth of King George whiting (Sillaginodes punctata) estimated from length-at-age samples of the legal-size harvest

Samples of 11,000 King George whiting (Sillaginodes punctata) from the South Australian commercial and recreational catch, supplemented by research samples, were aged from otoliths. Samples were analyzed from three coastal regions and by sex. Most sampling was undertaken at fish processing plants, from which only fish longer than the legal minimum length were obtained. A left-truncated normal distribution of lengths at monthly age was therefore employed as model likelihood. Mean length-at-monthly-age was described by a generalized von Bertalanffy formula with sinusoidal seasonality. Likelihood standard deviation was modeled to vary allometrically with mean length. A range of related formulas (with 6 to 8 parameters) for seasonal mean length at age were compared. In addition to likelihood ratio tests of relative fit, model selection criteria were a minimum occurrence of high uncertainties (>20% SE), of high correlations (>0.9, >0.95, and >0.99) and of parameter estimates at their biological limits, and we sought a model with a minimum number of parameters. A generalized von Bertalanffy formula with t0 fixed at 0 was chosen. The truncated likelihood alleviated the overestimation bias of mean length at age that would otherwise accrue from catch samples being restricted to legal sizes.