Review of size- and age-dependence in batch spawning: implications for stock assessment of fish species exhibiting indeterminate fecundity

Most assessments of fish stocks use some measure of the reproductive potential of a population, such as spawning biomass. However, the correlation between spawning biomass and reproductive potential is not always strong, and it likely is weakest in the tropics and subtropics, where species tend to exhibit indeterminate fecundity and release eggs in batches over a protracted spawning season. In such cases, computing annual reproductive output requires estimates of batch fecundity and the annual number of batches—the latter subject to spawning frequency and duration of spawning season. Batch fecundity is commonly measured by age (or size), but these other variables are not. Without the relevant data, the annual number of batches is assumed to be invariant across age. We reviewed the literature and found that this default assumption lacks empirical support because both spawning duration and spawning frequency generally increase with age or size. We demonstrate effects of this assumption on measures of reproductive value and spawning potential ratio, a metric commonly used to gauge stock status. Model applications showed substantial sensitivity to age dependence in the annual number of batches. If the annual number of batches increases with age but is incorrectly assumed to be constant, stock assessment models would tend to overestimate the biological reference points used for setting harvest rates. This study underscores the need to better understand the age- or size-dependent contrast in the annual number of batches, and we conclude that, for species without evidence to support invariance, the default assumption should be replaced with one that accounts for age- or size-dependence.

A comparison of size and age of red snapper (Lutjanus campechanus) with the age of artificial reefs in the northern Gulf of Mexico

Despite extensive study, it still is not clear whether artificial reefs produce new fish biomass or whether they only attract various species and make them more vulnerable to fishing mortality. To further evaluate this question, the size and age of red snapper (Lutjanus campechanus) were sampled from April to November 2010 at artificial reefs south of Mobile Bay off the coast of Alabama and compared with the age of the artificial reef at the site of capture. Red snapper were collected with hook and line and a fish trap and visually counted during scuba-diver surveys. In the laboratory, all captured red snapper were weighed and measured, and the otoliths were removed for aging. The mean age of red snapper differed significantly across reefs of different ages, with older reefs having older fish. The mean age of red snapper at a particular reef was not related to reef depth or distance to other reefs. The positive correlation between the mean age of red snapper and the age of the reef where they were found supports the contention that artificial reefs in the northern Gulf of Mexico enhance production of red snapper. The presence of fish older than the reef indicates that red snapper are also attracted to artificial reefs.

Changes in size and age at maturity of the northern stock of Tilefish (Lopholatilus chamaeleonticeps) after a period of overfishing

The modern fishery for Tilefish (Lopholatilus chamaeleonticeps) developed during the 1970s, offshore of southern New England, in the western North Atlantic Ocean. The population quickly became over exploited, with documented declines in catch rates and changes in demographic traits. In an earlier study, median size at maturity (L50) of males declined from 62.6 to 38.6 cm fork length (FL) and median age at maturity (A50) of males declined from 7.1 to 4.6 years between 1978 and 1982. As part of a cooperative research effort to improve the data-limited Tilefish assessment, we updated maturity parameter estimates through the use of an otolith aging method and macroscopic and microscopic evaluations of gonads. The vital rates for this species have continued to change, particularly for males. By 2008, male L50 and A50 had largely rebounded, to 54.1 cm FL and 5.9 years. Changes in female reproductive schedules were less variable among years, but the smallest L50 and youngest A50 were recorded in 2008. Tilefish are dimorphic, where the largest fish are male, and male spawning success is postulated to be socially mediated. These traits may explain the initial rapid decline and the subsequent rebound in male L50 and A50 and less dramatic effects on females. Other factors that likely contribute to the dynamics of maturity parameter estimates are the relatively short period of overfishing and the amount of time since efforts to rebuild this fishery began, as measured in numbers of generations. This study also confirms the gonochoristic sexual pattern of the northern stock, and it reveals evidence of age truncation and relatively high proportions of immature Tilefish in the recent catch.

Age, size, and sexual maturity of channeled whelk (Busycotypus canaliculatus) in Buzzards Bay, Massachusetts

With the southern New England lobster fishery in distress, lobster fishermen have focused more effort toward harvesting channeled whelk (Busycotypus canaliculatus). However, minimal research has been conducted on the life history and growth rates of channeled whelk. Melongenid whelks generally grow slowly and mature late in life, a characteristic that can make them vulnerable to overfishing as fishing pressure increases. We sampled channeled whelk from Buzzards Bay, Massachusetts, in August 2010 and in July 2011, studied their gonad development by histology, and aged them by examining opercula. Males had a slower growth rate and a lower maximum size than females. Male whelk reached 50% maturity (SM50) at 115.5 mm shell length (SL) and at the age of 6.9 years. Female whelk reached SM50 at 155.3 mm SL and at the age of 8.6 years. With a minimum size limit of 69.9 mm (2.75 in) in shell width, males entered the fishery at 7.5 years, a few months after SM50, but females entered the fishery at 6.3 years, approximately 2 years before SM50. Increased fishing pressure combined with slow growth rates and the inability to reproduce before being harvested can easily constrain the long-term viability of the channeled whelk fishery in Massachusetts.

Modeling seabird group size: implications for ecological impact assessments

The purpose of this project is to model seabird flock size data to provide recommendations to the Bureau of Ocean and Energy Management for offshore wind turbine placement. Our hypothesis is that ecological characteristics influence which statistical distribution will provide the best fit to seabird flock size data. To test this, seabird species can be grouped based on shared ecological traits, such as foraging mechanism or diet.

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.