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.

New insights into the diets of harbor seals (Phoca vitulina) in the Salish Sea revealed by analysis of fatty acid signatures

Harbor seals (Phoca fvitulina) are an abundant predator along the west coast of North America, and there is considerable interest in their diet composition, especially in regard to predation on valued fish stocks. Available informationon harbor seal diets, primarily derived from scat analysis, suggests that adult salmon (Oncorhynchus spp.), Pacific Herring (Clupea pallasii), and gadids predominate. Because diet assessments based on scat analysis may be biased, we investigated diet composition through quantitative analysis of fatty acid signatures. Blubber samples from 49 harbor seals captured in western North America from haul-outs within the area of the San Juan Islands and southern Strait of Georgia in the Salish Sea were analyzed for fatty acid composition, along with 269 fish and squid specimens representing 27 potential prey classes. Diet estimates varied spatially, demographically, and among individual harbor seals. Findings confirmed the prevalence of previously identified prey species in harbor seal diets, but other species also contributed significantly. In particular, Black (Sebastes melanops) and Yellowtail (S. flavidus) Rockfish were estimated to compose up to 50% of some individual seal diets. Specialization and high predation rates on Black and Yellowtail Rockfish by a subset of harbor seals may play a role in the population dynamics of these regional rockfish stocks that is greater than previously realized.

Fish consumption by harbor seals (Phoca vitulina) in the San Juan Islands, Washington

The harbor seal (Phoca vitulina) is a large-bodied and abundant predator in the Salish Sea ecosystem, and its population has recovered since the 1970s after passage of the Marine Mammal Protection Act and the cessation of bounties. Little is known about how this large predator population may affect the recovery of fish stocks in the Salish Sea, where candidate marine protected areas are being proposed. We used a bioenergetics model to calculate baseline consumption rates in the San Juan Islands, Washington. Salmonids (Oncorhynchus spp.) and herring (Clupeidae) were the 2 most energetically important prey groups for biomass consumed by harbor seals. Estimated consumption of salmonids was 783 (±380 standard deviation [SD]) metric tons (t) in the breeding season and 675 (±388 SD t in the nonbreeding season. Estimated consumption of herring was 646 (±303 SD) t in the breeding season and 2151 (±706 SD) t in the nonbreeding season. Rockfish, a depressed fish stock currently in need of population recovery, composed one of the minor prey groups consumed by harbor seals (84 [±26 SD] t in the nonbreeding season). The variables of seal body mass and proportion of prey in seal diet explained >80% of the total variation in model outputs. Prey groups, such as rockfish, that are targeted for recovery may still be affected by even low levels of predation. This study highlights the importance of salmonids and herring for the seal population and provides a framework for refining consumption estimates and their confidence intervals with future data.

Shifts in the estuarine demersal fish community after a fishery closure in Puget Sound, Washington

Puget Sound is one of the largest and most ecologically significant estuaries in the United States, but the status and trends of many of its biological components are not well known. We analyzed a 21-year time series of data from standardized bottom trawl sampling at a single study area to provide the first assessment of population trends of Puget Sound groundfishes after the closure of bottom trawl fisheries. The expected increase in abundance was observed for only 3 of 14 species after this closure, and catch rates of most (10) of the abundant species declined through time. Many of these changes were stepwise (abrupt) rather than gradual, and many stocks exhibited changes in catch rate during the 3-year period from 1997 through 2000. No detectable change was recorded for either temperature or surface salinity over the entire sampling period. The abrupt density reductions that were observed likely do not reflect changes in demographic rates but may instead represent distributional shifts within Puget Sound.

Three decades of U.S. Gulf of Mexico white shrimp, Litopenaeus setiferus, commercial catch statistics

Gulf of Mexico, white shrimp, Litopenaeus setiferus, catch statistics have been collected by NOAA’s National Marine Fisheries Service for over 50 years. Recent occurrences such as natural and manmade disasters have raised awareness for the need to publish these types of data. Here we report shrimp data collected from 1984 to 2011. These 28 years of catch history are the time series used in the most recent Gulf of Mexico white shrimp stock assessment. Fishing effort for this stock has fluctuated over the period reported, ranging from 54,675 to 162,952 days fished. Catch averaged 55.7 million pounds per year, increasing significantly over the times series. In addition, catch rates have been increasing in recent years, with CPUE levels ranging from 315 lb/day fished in 2002, to 1,175 lb/ day fished in 2008. The high CPUE’s we have measured is one indication that the stock was not in decline during this time period. Consequently, we believe the decline in effort levels is due purely to economic factors. Current stock assessments are now using these baseline data to provide managers with further insights into the Gulf L. setiferus stocks.

Age determination manual of the Alaska Fisheries Science Center Age and Growth Program

The Age and Growth Program at the Alaska Fisheries Science Center is tasked with providing age data in order to improve the basic understanding of the ecology and fisheries dynamics of Alaskan fish species. The primary focus of the Age and Growth Program is to estimate ages from otoliths and other calcified structures for age-structured modeling of commercially exploited stocks; however, the program has recently expanded its interests to include numerous studies on topics ranging from age estimate validation to the growth and life history of non-target species. Because so many applications rely upon age data and particularly upon assurances as to their accuracy and precision, the Age and Growth Program has developed this practical guide to document the age determination of key groundfish species from Alaskan waters. The main objective of this manual is to describe techniques specific to the age determination of commercially and ecologically important species studied by the Age and Growth Program. The manual also provides general background information on otolith morphology, dissection, and preparation, as well as descriptions of methods used to measure precision and accuracy of age estimates. This manual is intended not only as a reference for age readers at the AFSC and other laboratories, but also to give insight into the quality of age estimates to scientists who routinely use such data.

Our living oceans: report on the status of U.S. living marine resources, 6th edition

The National Marine Fisheries Service (NMFS) is dedicated to the stewardship of living marine resources (LMR’s). This is accomplished through science-based conservation and management, and the promotion of healthy ecosystems. As a steward, NMFS has an obligation to conserve, protect, and manage these resources in a way that ensures their continuation as functioning components of healthy marine ecosystems, affords economic opportunities, and enhances the quality of life for the American public. In addition to its responsibilities within the U.S. Exclusive Economic Zone (EEZ), NMFS plays a supportive and advisory role in the management of LMR’s in the coastal areas under state jurisdiction and provides scientific and policy leadership in the international arena. NMFS also implements international measures for the conservation and management of LMR’s, as appropriate.NMFS receives its stewardship responsibilities under a number of Federal laws. These include the Nation’s primary fisheries law, the Magnuson Fishery Conservation and Management Act. This law was first passed in 1976, later reauthorized as the Magnuson-Stevens Fishery Conservation and Management Act in 1996, and reauthorized again on 12 January 2007 as the Magnuson-Stevens Fishery Conservation and Management Reauthorization Act (MSRA). The MSRA mandates strong action to conserve and manage fishery resources and requires NMFS to end overfishing by 2010 in all U.S. commercial and recreational fisheries, rebuild all overfished stocks, and conserve essential fish habitat.

Bering Sea FOCI Final Report

To develop an understanding of stock structure and recruitment variation in Bering Sea pollock, the Coastal Ocean Program of the National Oceanic and Atmospheric Administration (NOAA) funded an 7-year (1991-1997), interdisciplinary project named Bering Sea Fisheries-Oceanography Coordinated Investigations (BS FOCI; Schumacher and Kendall, 1995) for which NOAA and academic researchers were selected through a competitive process (Macklin, this report). The project goals, based on recommendations from an international symposium on pollock (Aron and Balsiger, 1989) were to (1) determine stock structure in the Bering Sea and its potential relationship to physical oceanography, and (2) examine recruitment processes in the eastern Bering Sea. Both of these have direct implication to management. An integrated set of field, laboratory, and modeling studies were established to accomplish these goals. To address the first goal, project objectives were to establish details of oceanic circulation relevant to larval dispersal and separation of stocks, and determine if unique chemical or genetic indicators existed for different stocks. The recruitment component of BS FOCI, addressing the second goal, focused on understanding causes of variable mortality of pollock larvae in the different habitats of the eastern Bering Sea. The emphasis of recruitment studies was to determine the dominant physical oceanographic features (turbulence, temperature, and transport) that could influence survival of pollock larvae, and investigate factors controlling food production for the larvae. A later component contrasted juvenile habitat in three hydrographic regimes around the Pribilof Islands (Brodeur, this report).

Synthesis of summer flounder habitat parameters

The summer flounder, Paralichthys dentatus, is overexploited and is currently at very low levels of abundance. This is reflected in the compressed age structure of the population and the low catches in both commercial and recreational fisheries. Declining habitat quantity and quality may be contributing to these declines, however we lack a thorough understanding of the role of habitats in the population dynamics of this species. Stock structure is unresolved and current interpretations, depending on the technique and study area, suggest that there may be two or three spawning populations. If so, these stocks may have differing habitat requirements. In response to this lack of knowledge, this document summarizes and synthesizes the available information on summer flounder habitat in all life history stages (eggs, larvae, juveniles and adults) and identifies areas where further research is needed. Several levels of investigation were conducted in order to produce this document. First, an extensive search for summer flounder habitat information was made, which included both the primary and gray literature as well as unanalyzed data. Second, state and federal fisheries biologists and resource managers in all states within the primary range of summer flounder (Massachusetts to Florida) were interviewed along with a number of fish ecologists and summer flounder experts from the academic and private sectors. Finally, information from all sources was analyzed and synthesized to form a coherent overview. This document first presents an overview of the economic importance and current status of summer flounder (Chapter 1). It then summarizes our present state of knowledge of summer flounder distribution, life history patterns and stock identification (Chapter 2). This is followed by a synopsis of habitat requirements during each life history stage. For convenience, this is presented by general habitat as offshore eggs (Chapter 3), offshore larvae (Chapter 4), estuarine larvae (Chapter 5), estuarine juveniles (Chapter 6), offshore juveniles (Chapter 7) and estuarine and offshore adults (Chapter 8). In several instances, previously undigested data sets are analyzed to provide more detailed information, especially for estuarine juveniles. The information is then discussed in terms of its relevance to resource managers (Chapter 9).

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.

Sex-specific growth and mortality, spawning season, and female maturation of the stripey bass (Lutjanus carponotatus) on the Great Barrrier Reef

Sex-specific demography and reproductive biology of stripey bass (Lutjanus carponotatus) (also known as Spanish flag snapper, FAO) were examined at the Palm and Lizard island groups, Great Barrier Reef (GBR).Total mortality rates were similar between the sexes. Males had larger L∞ at both island groups and Lizard Island group fish had larger overall L∞. Female:male sex ratios were 1.3 and 1.1 at the Palm and Lizard island groups, respectively. The former is statistically different from 1, but is unlikely significantly different in a biological sense. Females matured on average at 2 years of age and 190 mm fork length at both locations. Female gonadal lipid body indices peaked from August through October, preceding peak gonadosomatic indices in October, November, and December that were twice as great as in any other month. However, ovarian staging revealed 50% or more ovaries were ripe from September through February, suggesting a more protracted spawning season and highlighting the different interpretations that can arise between gonad weight and gonad staging methods. Gonadosomatic index increases slightly with body size and larger fish have a longer average spawning season, which suggests that larger fish produce greater relative reproductive output. Lizard Island group females had ovaries nearly twice as large as Palm Island group females at a given body size. However, it is unclear whether this reflects spatial differences akin to those observed in growth or effects of sampling Lizard Island group fish closer to their date of spawning. These results support an existing 250 mm minimum size limit for L. carponotatus on the GBR, as well as the timing of a proposed October through December spawning closure for the fishery. The results also caution against assessing reef-fish stocks without reference to sex-, size-, and location-specific biological traits.

Examination of the foraging habits of Pacific harbor seal (Phoca vitulina richardsi) to describe their use of the Umpqua River, Oregon, and their predation on salmonids

The increase in harbor seal (Phoca vitulina richardsi) abundance, concurrent with the decrease in salmonid (Oncorhynchus spp.) and other fish stocks, raises concerns about the potential negative impact of seals on fish populations. Although harbor seals are found in rivers and estuaries, their presence is not necessarily indicative of exclusive or predominant feeding in these systems. We examined the diet of harbor seals in the Umpqua River, Oregon, during 1997 and 1998 to indirectly assess whether or not they were feeding in the river. Fish otoliths and other skeletal structures were recovered from 651 scats and used to identify seal prey. The use of all diagnostic prey structures, rather than just otoliths, increased our estimates of the number of taxa, the minimum number of individuals and percent frequency of occurrence (%FO) of prey consumed. The %FO indicated that the most common prey were pleuronectids, Pacific hake (Merluccius productus), Pacific stag-horn sculpin (Leptocottus armatus), osmerids, and shiner surfperch (Cymatogaster aggregata). The majority (76%) of prey were fish that inhabit marine waters exclusively and fish found in marine and estuarine areas (e.g. anadromous spp.) which would indicate that seals forage predominantly at sea and use the estuary for resting and opportunistic feeding. Salmonid remains were encountered in 39 samples (6%); two samples contained identifiable otoliths, which were determined to be from chi-nook salmon (O. tshawytscha). Because of the complex salmonid composition in the Umpqua River, we used molecular genetic techniques on salmonid bones retrieved from scat to discern species that were rare from those that were abundant. Of the 37 scats with salmonid bones but no otoliths, bones were identified genetically as chinook or coho (O. kisutch) salmon, or steelhead trout (O. mykiss) in 90% of the samples.

Molecular methods for the genetic identification of salmonid prey from Pacific harbor seal (Phoca vitulina richardsi) scat

Twenty-six stocks of Pacific salmon and trout (Oncorhynchus spp.), representing evolutionary significant units (ESU), are listed as threatened or endangered under the Endangered Species Act (ESA) and six more stocks are currently being evaluated for listing. The ecological and economic consequences of these listings are large; therefore considerable effort has been made to understand and respond to these declining populations. Until recently, Pacific harbor seals (Phoca vitulina richardsi) on the west coast increased an average of 5% to 7% per year as a result of the Marine Mammal Protection Act of 1972 (Brown and Kohlman2). Pacific salmon are seasonally important prey for harbor seals (Roffe and Mate, 1984; Olesiuk, 1993); therefore quantifying and understanding the interaction between these two protected species is important for Morphobiologically sound management strategies. Because some Pacific salmonid species in a given area may be threatened or endangered, while others are relatively abundant, it is important to distinguish the species of salmonid upon which the harbor seals are preying. This study takes the first step in understanding these interactions by using molecular genetic tools for species-level identification of salmonid skeletal remains recovered from Pacific harbor seal scats.

Translocation as a strategy to rehabilitate the queen conch (Strombus gigas) population in the Florida Keys

Queen conch (Strombus gigas) stocks in the Florida Keys once supported commercial and recreational fisheries, but overharvesting has decimated this once abundant snail. Despite a ban on harvesting this species since 1985, the local conch population has not recovered. In addition, previous work has reported that conch located in nearshore Keys waters are incapable of spawning because of poor gonadal condition, although reproduction does occur offshore. Queen conch in other areas undergo ontogenetic migrations from shallow, nearshore sites to offshore habitats, but conch in the Florida Keys are prevented from doing so by Hawk Channel. The present study was initiated to determine the potential of translocating nonspawning nearshore conch to offshore sites in order to augment the spawning stock. We translocated adult conch from two nearshore sites to two offshore sites. Histological examinations at the initiation of this study confirmed that nearshore conch were incapable of reproduction, whereas offshore conch had normal gonads and thus were able to reproduce. The gonads of nearshore females were in worse condition than those of nearshore males. However, the gonadal condition of the translocated nearshore conch improved, and these animals began spawning after three months offshore. This finding suggests that some component of the nearshore environment (e.g., pollutants, temperature extremes, poor food or habitat quality) disrupts reproduction in conch, but that removal of nearshore animals to suitable offshore habitat can restore reproductive viability. These results indicate that translocations are preferable to releasing hatchery-reared juveniles because they are more cost-effective, result in a more rapid increase in reproductive output, and maintain the genetic integrity of the wild stock. Therefore, translocating nearshore conch to offshore spawning aggregations may be the key to expediting the recovery of queen conch stocks in the Florida Keys.

Impact of hatchery releases on the recreational fishery for Pacific threadfin (Polydactylus sexfilis) in Hawaii

The Pacific threadfin (Polydactylus sexfilis) is considered one of the premier Hawaiian food fishes but even with catch limits, seasonal closures, and size limits, catches have declined dramatically since the 1960s. It was identified as the top candidate species for stock enhancement in Hawaii, based on the decline in stocks, high market value, and importance of the fishery. In the stock enhancement program for Pacific threadfin, over 430,000 fingerlings of various sizes were implanted with coded wire tags and released in nursery habitats along the windward coast of Oahu between 1993 and 1998. Because few Pacific threadfin were present in creel surveys conducted between 1994 and 1998, Oahu fishermen were offered a $10 reward for each threadfin that was caught (for both hatchery-reared and wild fish). A total of 1882 Pacific threadfin were recovered from the reward program between March 1998 and May 1999, including 163 hatchery-reared fish, an overall contribution of 8.7% to the fishery. Hatchery-reared fish accounted for as high as 71% of returns in the release areas. Hatchery-reared fish were recovered on average 11.5 km (SD=9.8 km) from the release site, although some had moved as far away as 42 km. Average age for recovered hatchery-reared fish was 495 days; the oldest was 1021 days. Cultured Pacific threadfin juveniles survived and recruited successfully to the recreational fishery, accounting for 10% of fishermen’s catches on the windward side of Oahu. Recruitment to the fishery was highest for the 1997 release year; few juveniles from earlier releases were observed. Presence of a few large, fully developed females in the recreational fishery suggested that hatchery-reared fish can survive, grow, and reproductively contribute to the population. Implementation of an enhancement program that is focused on juveniles and perhaps large females, as part of an integrated fishery management strategy, could speed the recovery of this fish population.