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.

Age validation of juvenile Shortfin Mako (Isurus oxyrinchus) tagged and marked with oxytetracycline off southern California

The purpose of this study was to validate aging results of juvenile Shortfin Mako (Isurus oxyrinchus) by vertebral band counts. Vertebrae of 29 juvenile Shortfin Mako marked with oxytetracycline (OTC) were obtained from tag-recapture activities to determine centrum growth-band deposition. Tagging occurred off southern California from 1996 to 2010, and time at liberty of the 29 sharks ranged from 4 months to 4.4 years (mean=1.3 years). Growth information also was obtained from length-frequency modal analyses (MULTIFAN and MIXDIST) by using a 29-year data set of commercial and research catch data, in addition to a tag-recapture growth model (e.g, the GROTAG model). For vertebrae samples used for age validation, shark size at time of release ranged from 79 to 142 cm fork length (FL) and from 98 to 200 cm FL at recapture. Results from band counts of vertebrae distal to OTC marks indicate 2 band pairs (2 translucent and 2 opaque) are formed each year for Shortfin Mako of the size range examined. Length-frequency analyses identified 3 age class modes. Growth rate estimates from 26.5 to 35.5 cm/year were calculated for the first age-class mode (85 cm FL) and from 22.4 to 28.6 cm/year for the second age-class mode (130 cm FL). Results from the tag-recapture growth model revealed fast growth during time at liberty for tagged fish of the 2 youngest age classes. Collectively, these methods suggest rapid growth of juvenile Shortfin Mako in the southern California study area and indicate biannual deposition of growth bands in vertebrae for the first 5 years.

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.