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.

Distribution of ommastrephid paralarvae in the eastern tropical Pacific

Jumbo squid (Dosidicus gigas) and purpleback squid (Sthenoteuthis oualaniensis) (Teuthida: Ommastrephidae) are thought to spawn in the eastern tropical Pacific. We used 10 years of plankton tow and oceanographic data collected in this region to examine the reproductive habits of these 2 ecologically important squid. Paralarvae of jumbo squid and purpleback squid were found in 781 of 1438 plankton samples from surface and oblique tows conducted by the Southwest Fisheries Science Center (NOAA) in the eastern tropical Pacific over the 8-year period of 1998–2006. Paralarvae were far more abundant in surface tows (maximum: 1588 individuals) than in oblique tows (maximum: 64 individuals). A generalized linear model analysis revealed sea-surface temperature as the strongest environmental predictor of paralarval presence in both surface and oblique tows; the likelihood of paralarval presence increases with increasing temperature. We used molecular techniques to identify paralarvae from 37 oblique tows to species level and found that the purpleback squid was more abundant than the jumbo squid (81 versus 16 individuals).

Staging ovaries of Haddock (Melanogrammus aeglefinus): implications for maturity indices and field sampling practices

We build on recent efforts to standardize maturation staging methods through the development of a field-proof macroscopic ovarian maturity index for Haddock (Melanogrammus aeglefinus) for studies on diel spawning periodicity. A comparison of field and histological observations helped us to improve the field index and methods, and provided useful insight into the reproductive biology of Haddock and other boreal determinate fecundity species. We found reasonable agreement between field and histological methods, except for the regressing and regenerating stages (however, differentiation of these 2 stages is the least important distinction for determination of maturity or reproductive dynamics). The staging of developing ovaries was problematic for both methods partly because of asynchronous oocyte hydration during the early stage of oocyte maturation. Although staging on the basis of histology in a laboratory is generally more accurate than macroscopic staging methods in the field, we found that field observations can uncover errors in laboratory staging that result from bias in sampling unrepresentative portions of ovaries. For 2 specimens, immature ovaries observed during histological examination were incorrectly assigned as regenerating during macroscopic staging. This type of error can lead to miscalculation of length at maturity and of spawning stock biomass, metrics that are used to characterize the state of a fish population. The revised field index includes 3 new macroscopic stages that represent final oocyte maturation in a batch of oocytes and were found to be reliable for staging spawning readiness in the field. The index was found to be suitable for studies of diel spawning periodicity and conforms to recent standardization guidelines.

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.

Establishing species–habitat associations for 4 eteline snappers with the use of a baited stereo-video camera system

With the use of a baited stereo-video camera system, this study semiquantitatively defined the habitat associations of 4 species of Lutjanidae: Opakapaka (Pristipomoides filamentosus), Kalekale (P. sieboldii), Onaga (Etelis coruscans), and Ehu (E. carbunculus). Fish abundance and length data from 6 locations in the main Hawaiian Islands were evaluated for species-specific and size-specific differences between regions and habitat types. Multibeam bathymetry and backscatter were used to classify habitats into 4 types on the basis of substrate (hard or soft) and slope (high or low). Depth was a major influence on bottomfish distributions. Opakapaka occurred at depths shallower than the depths at which other species were observed, and this species showed an ontogenetic shift to deeper water with increasing size. Opakapaka and Ehu had an overall preference for hard substrate with low slope (hard-low), and Onaga was found over both hard-low and hard-high habitats. No significant habitat preferences were recorded for Kalekale. Opakapaka, Kalekale, and Onaga exhibited size-related shifts with habitat type. A move into hard-high environments with increasing size was evident for Opakapaka and Kalekale. Onaga was seen predominantly in hard-low habitats at smaller sizes and in either hard-low or hard-high at larger sizes. These ontogenetic habitat shifts could be driven by reproductive triggers because they roughly coincided with the length at sexual maturity of each species. However, further studies are required to determine causality. No ontogenetic shifts were seen for Ehu, but only a limited number of juveniles were observed. Regional variations in abundance and length were also found and could be related to fishing pressure or large-scale habitat features.

James's rule and causes and consequences of a latitudinal cline in the demography of John's Snapper (Lutjanus johnii) in coastal waters of Australia

Demographic parameters were derived from sectioned otoliths of John’s Snapper (Lutjanus johnii) from 4 regions across 9° of latitude and 23° of longitude in northern Australia. Latitudinal variation in size and growth rates of this species greatly exceeded longitudinal variation. Populations of John’s Snapper farthest from the equator had the largest body sizes, in line with James’s rule, and the fastest growth rates, contrary to the temperature-size rule for ectotherms. A maximum age of 28.6 years, nearly 3 times previous estimates, was recorded and the largest individual was 990 mm in fork length. Females grew to a larger mean asymptotic fork length (L∞) than did males, a finding consistent with functional gonochorism. Otolith weight at age and gonad weight at length followed the same latitudinal trends seen in length at age. Length at maturity was ~72–87% of L∞ and varied by ~23% across the full latitudinal gradient, but age at first maturity was consistently in the range of 6–10 years, indicating that basic growth trajectories were similar across vastly different environments. We discuss both the need for complementary reproductive data in age-based studies and the insights gained from experiments where the concept of oxygen- and capacity-limited thermal tolerance is applied to explain the mechanistic causes of James’s rule in tropical fish species.

Maternal effects on fecundity and egg quality of the Patagonian stock of Argentine Hake (Merluccius hubbsi)

The influences of age, size, and condition of spawning females on fecundity and oocyte quality were analyzed for the Patagonian stock of Argentine Hake (Merluccius hubbsi). Samples of mature females were collected in the spawning area as part of 2 research surveys conducted in January 2010 and 2011, during the peak of the reproductive season. Batch fecundity (BF) ranged between 40,500 (29 cm total length [TL]) and 2,550,000 (95 cm TL) hydrated oocytes, and was positively correlated with TL, gutted weight, age, hepatosomatic index (HSI), and the relative condition factor (Kn). Relative fecundity ranged between 85 and 1040 hydrated oocytes g–1 and showed significant positive relationships with gutted weight, HSI, and Kn; however, coefficients of determination were low for all regressions. Dry weights of samples of 100 hydrated oocytes ranged between 1.8 and 3.95 mg and were positively correlated with all variables analyzed, including batch and relative fecundity. Multiple regression models created with data of the morphophysiological characteristics of females supported maternal influences on fecundity and egg weights. Within the studied size range (29–95 cm TL), larger individuals had better somatic and egg condition, mainly revealed by higher HSI and hydrated oocytes with larger oil droplets (275.71μm [standard error 1.49]). These results were associated with the higher feeding activity of larger females during the spawning season in comparison with the feeding activity of young individuals (<5 years old); the better nutritional state of larger females, assumed to result from more feeding, was conducive to greater production of high-quality eggs.

Holistic fisheries management: combining macroecology, ecology, and evolutionary biology

Ecosystem-based management is one of many indispensable components of objective, holistic management of human impacts on nonhuman systems. By itself, however, ecosystem-based management carries the same risks we face with other forms of current management; holism requires more. Combining single-species and ecosystem approaches represents progress. However, it is now recognized that management also needs to be evosystem-based. In other words, management needs to account for all coevolutionary and evolutionary interactions among all species; otherwise we fall far short of holism. Fully holistic practices are quite distinct from the approaches to the management of fisheries that are applied today. In this paper, we show how macroecological patterns can guide management consistently, objectively, and holistically. We present one particular macroecological pattern with two applications. The first application is a case study of fisheries from the Baltic Sea involving historical data for two species; the second involves a sample of 44 species of primarily marine fish worldwide. In both cases we evaluate historical fishing rates and determine holistic/systemic sustainable single-species fishing rates to illustrate that conventional fisheries management leads to much more extensive and pervasive overfishing than currently realized; harvests are, on average, over twenty-fold too large to be fully sustainable. In general, our approach involves not only the sustainability of fisheries and related resources but also the sustainability of the ecosystems and evosystems in which they occur. Using macroecological patterns accomplishes four important goals: 1) Macroecology becomes one of the interdisciplinary components of management. 2) Sustainability becomes an option for harvests from populations of individual species, species groups, ecosystems, and the entire marine environment. 3) Policies and goals are reality-based, holistic, or fully systemic; they account for ecological as well as evolutionary factors and dynamics (including management itself). 4) Numerous management questions can be addressed.

Report of 2009 Bottlenose Dolphin (Tursiops truncatus) Life History Workshop: expanding network capabilities

The Virginia Aquarium & Marine Science Center Foundation’s Stranding Response Program (VAQS) was awarded a grant in 2008 to conduct life history analysis on over 10 years of Tursiops truncatus teeth and gonad samples from stranded animals in Virginia. A major part of this collaborative grant included a workshop involving life historians from Hubbs-Sea World Research Institute (HSWRI), NOS, Texas A & M University (TAMU), and University of North Carolina Wilmington (UNCW). The workshop was held at the NOAA Center for Coastal Environmental Health & Biomolecular Research in Charleston, SC on 7-9 July 2009. The workshop convened to 1) address current practices among the groups conducting life history analysis, 2) decide on protocols to follow for the collaborative Prescott grant between VAQS and HSWRI, 3) demonstrate tissue preparation techniques and discuss shortcuts and pitfalls, 4) demonstrate data collection from prepared testes, ovaries, and teeth, and 5) discuss data analysis and prepare an outline and timeline for a future manuscript. The workshop concluded with discussions concerning the current collaborative Tursiops Life History Prescott grant award and the beginnings of a collaborative Prescott proposal with members of the Alliance of Marine Mammal Parks and Aquariums to further clarify reproductive analyses. This technical memorandum serves as a record of this workshop.

Oogenesis and spawn formation in the invasive lionfish, Pterois miles and Pterois volitans

The Indo-Pacific lionfish, Pterois miles and P. volitans, have recently invaded the U.S. east coast and the Caribbean and pose a significant threat to native reef fish communities. Few studies have documented reproduction in pteroines from the Indo-Pacific. This study provides a description of oogenesis and spawn formation in P. miles and P. volitans collected from offshore waters of North Carolina, U.S.A and the Bahamas. Using histological and laboratory observations, we found no differences in reproductive biology between P. miles and P. volitans. These lionfish spawn buoyant eggs that are encased in a hollow mass of mucus produced by specialized secretory cells of the ovarian wall complex. Oocytes develop on highly vascularized peduncles with all oocyte stages present in the ovary of spawning females and the most mature oocytes placed terminally, near the ovarian lumen. Given these ovarian characteristics, these lionfish are asynchronous, indeterminate batch spawners and are thus capable of sustained reproduction throughout the year when conditions are suitable. This mode of reproduction could have contributed to the recent and rapid establishment of these lionfish in the northwestern Atlantic and Caribbean.

Deep coral and associated species taxonomy and ecology: (DeepCAST) II Expedition Report, Roatan, Honduras, May 21-28, 2011

NOAA has a mandate to explore and understand deep-sea coral ecology under Magnuson-Stevens Sustainable Fisheries Conservation Act Reauthorization of 2009. Deep-sea corals are increasingly considered a proxy for marine biodiversity in the deep-sea because corals create complex structure, and this structure forms important habitat for associated species of shrimp, crabs, sea stars, brittle stars, and fishes. Yet, our understanding of the nature of the relationships between deep-corals and their associated species is incomplete. One of the primary challenges of conducting any type of deep-sea coral (DSC) research is access to the deep-sea. The deep-sea is a remote environment that often requires long surface transits and sophisticated research vehicles like submersibles and remotely operated vehicles (ROVs). The research vehicles often require substantial crew, and the vehicles are typically launched from large research vessels costing many thousands of dollars a day. To overcome the problem of access to the deep-sea, the Deep Coral and Associated Species Taxonomy and Ecology (DeepCAST) Expeditions are pioneering the use of shore-based submersibles equipped to do scientific research. Shore-based subs alleviate the need for expensive ships because they launch and return under their own power. One disadvantage to the approach is that shore-based subs are restricted to nearby sites. The disadvantage is outweighed, however, by the benefit of repeated observations, and the opportunity to reduce the costs of exploration while expanding knowledge of deep-sea coral ecology.

Seascape ecology of coastal biogenic habitats: advances, gaps, and challenges

We review the progress made in the emerging field of coastal seascape ecology, i.e. the application of landscape ecology concepts and techniques to the coastal marine environment. Since the early 1990s, the landscape ecology approach has been applied in several coastal subtidal and intertidal biogenic habitats across a range of spatial scales. Emerging evidence indicates that animals in these seascapes respond to the structure of patches and patch mosaics in different ways and at different spatial scales, yet we still know very little about the ecological significance of these relationships and the consequences of change in seascape patterning for ecosystem functioning and overall biodiversity. Ecological interactions that occur within patches and among different types of patches (or seascapes) are likely to be critically important in maintaining primary and secondary production, trophic transfer, biodiversity, coastal protection, and supporting a wealth of ecosystem goods and services. We review faunal responses to patch and seascape structure, including effects of fragmentation on 5 focal habitats: seagrass meadows, salt marshes, coral reefs, mangrove forests, and oyster reefs. Extrapolating and generalizing spatial relationships between ecological patterns and processes across scales remains a significant challenge, and we show that there are major gaps in our understanding of these relationships. Filling these gaps will be crucial for managing and responding to an inevitably changing coastal environment. We show that critical ecological thresholds exist in the structural patterning of biogenic ecosystems that, when exceeded, cause abrupt shifts in the distribution and abundance of organisms. A better understanding of faunal–seascape relationships, including the identifications of threshold effects, is urgently needed to support the development of more effective and holistic management actions in restoration, site prioritization, and forecasting the impacts of environmental change.

Patterns of scale-dependency and the influence of map resolution on the seascape ecology of reef fish

Detection and perception of ecological relationships between biota and their surrounding habitats is sensitive to analysis scale and resolution of habitat data. We measured strength of univariate linear correlations between reef fish and seascape variables at multiple spatial scales (25 to 800 m). Correlation strength was used to identify the scale that best associates fish to their surrounding habitat. To evaluate the influence of map resolution, seascape variables were calculated based on 4 separate benthic maps produced using 2 levels of spatial and thematic resolution, respectively. Individual seascape variables explained only 25% of the variability in fish distributions. Length of reef edge was correlated with more aspects of the fish assemblage than other features. Area of seagrass and bare sand correlated with distribution of many fish, not just obligate users. No fish variables correlated with habitat diversity. Individual fish species achieved a wider range of correlations than mobility guilds or the entire fish assemblage. Scales of peak correlation were the same for juveniles and adults in a majority of comparisons. Highly mobile species exhibited broader scales of peak correlation than either resident or moderately mobile fish. Use of different input maps changed perception of the strength and even the scale of peak correlations for many comparisons involving hard bottom edge length and area of sand, whereas results were consistent regardless of map type for comparisons involving area of seagrass and habitat diversity.

Practicing coastal seascape ecology

Landscape ecology concepts developed from terrestrial systems have recently emerged as theoretical and analytical frameworks that are equally useful for evaluating the ecological consequences of spatial patterns and structural changes in the submerged landscapes of coastal ecosystems. The benefits of applying a spatially-explicit perspective to resource management and restoration planning in the coastal zone are rapidly becoming apparent. This Theme Section on the application of landscape ecology to the estuarine and coastal environment emerged from a special symposium at the Coastal and Estuarine Research Federation (CERF) 20th Biennial Conference (Estuaries and Coasts in a Changing World) held in Portland, Oregon, USA, in November 2009. The 7 contributions in this Theme Section collectively provide substantial insights into the current status and application of the landscape approach in shallow marine environments, and identify significant knowledge gaps, as well as potential directions for the future advancement of ‘seascape ecology’.

Age, growth and mortality of the toadfish, Halobatrachus didactylus (Schneider, 1801) (Pisces: Batrachoididae), in the Bay of Cádiz (southwestern Spain)

Age, growth and mortality of the toadfish, Halobatrachus didactylus, were determined by examination of the whole sagittal otoliths of fish sampled in the Bay of Cádiz (southwestern Spain) from March 1999 to March 2000. A total of 844 specimens (425 males, 416 females, and 3 of indeterminate sex), ranging from 95 to 470 mm in total length were examined. Eighty-nine percent of the otoliths could be read allowing an age estimation. The opaque zone was formed between April and May coincident with the maximum reproductive peak, while the translucent zone formed mainly in summer-fall (June to December). Maximum ages for males and females were 12 and 10 years, respectively. The samples were dominated by 2- to 6-year-old specimens. Males matured at an age of approximately 2 years and females at 3 years. Fish total length and otolith radius were closely related. The von Bertalanffy growth curve was used to describe growth. The parameters were derived from back-calculated length-at-age. Significant differences in the growth parameters were found between sexes. Although the growth analysis revealed that this species is slow-growing, males reached larger sizes than females. Females appeared to experience higher natural mortality rates than males.