The impact of intrapopulation variability in reproductive traits on population reproductive potential of Grand Bank American plaice (Hippoglossoides platessoides) and yellowtail flounder (Limanda ferruginea)

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Abstract

In geographically extensive fish populations the potential exists for reproductive traits to vary over the population’s range but, the impact that such intrapopulation variability has on overall population reproductive potential has not been formally assessed. Here intrapopulation spatial variability in size at maturity and fecundity are demonstrated for Grand Bank American plaice (Hippoglossoides platessoides) and yellowtail flounder (Limanda ferruginea). Recognition of intrapopulation variability in these reproductive traits coupled with spatial variability in abundance resulted in an increase in estimated population total egg production (TEP) by as much as 1014 eggs for American plaice and 1015 eggs for yellowtail flounder as compared to assessment of TEP for the population as a whole. Results highlight the need to explore variability in life history traits not only between but, also within populations and emphasizes the need for sufficient spatial coverage during sampling in order to assess the reproductive potential of fish populations.

Introduction

The nature of perceived relationships between animals and their environment can depend on the scale at which the system is viewed (Levin, 1992). The scale of observation, however, is usually set a priori and can lead to important aspects of the system being overlooked (Peterson and Parker, 1998). In fishes, for example, spatial variability in life history traits is often assessed at the population level, with less attention directed at potential spatial variability within populations. Even less emphasis has been placed on understanding the impact that potential intrapopulation variability in life history traits might have on population productivity.

Reproduction in fishes is heavily dependent on environmental conditions. For example, conditions that support increased growth rate and higher fish condition, promote maturation at an earlier age/size (Gadgil and Bossert, 1970, Rijnsdorp, 1993, Morgan and Colbourne, 1999, Olsen et al., 2004). In addition, fish of larger size and better condition produce more eggs than smaller and poorer conditioned individuals (Gundersen et al., 1999, Kraus et al., 2002, Blanchard et al., 2003, Lambert et al., 2006) and may have a higher probability of successfully spawning (Rideout et al., 2005). Temperature can also influence fish reproduction (Hodder, 1965, Kjesbu et al., 1998, Yoneda and Wright, 2005a, Yoneda and Wright, 2005b), not only through its impact on growth rate, but also by altering the animal's energy budget or by directly affecting the biochemical processes within developing gametes.

Because environmental conditions can be highly variable over space and time, scale of observation is an important consideration in studying the reproductive and recruitment processes in fishes (Schneider, 2001). Size at maturity and fecundity at length data are typically presented as population values with little consideration for intrapopulation variability. Only a limited number of studies on fishes have reported maturation to be heterogeneous across the range of a population (Korsbrekke, 1999, Bromley, 2000, Gerritsen et al., 2003, Armstrong et al., 2004, Boulcott et al., 2007). Even fewer have demonstrated differences in fecundity between areas within a population (Pinhorn, 1984, Rijnsdorp, 1991, Kjesbu et al., 1998, Nash et al., 2000). It has been suggested that such regional differences might have significant consequences for resource management (Boulcott et al., 2007) but the impact that failure to recognize this variability has on perception of stock reproductive potential has rarely been examined.

American plaice (Hippoglossoides platessoides) and yellowtail flounder (Limanda ferruginea) are two commercially important flatfish species with overlapping distributions off the east coast of Canada. For each species, individuals over the entire Grand Bank, a large geographical area encompassing three Northwest Atlantic Fisheries Organization (NAFO) Divisions (3LNO), are treated as a single population (Fig. 1). Temporal variability in both maturity (Morgan and Colbourne, 1999, Walsh and Morgan, 1999) and fecundity (Rideout and Morgan, 2007) have been reported for the Grand Bank populations of both species. Large scale spatial variability in these reproductive traits has also been demonstrated, with differences between these and other nearby populations (Rideout and Morgan, 2007). The large geographic area covered by the Grand Bank populations of flatfish suggests the potential for environmental conditions to vary over the range of the populations, and hence create the potential for reproductive traits to vary over a smaller spatial scale within the populations. Intrapopulation spatial variability in reproductive traits has been reported for Grand Bank American plaice based on limited sampling (Pitt, 1964, Pitt, 1966) but has not been examined for yellowtail flounder. Here we examine not only intrapopulation variability in maturation and fecundity for both American plaice and yellowtail flounder, but also investigate the potential impacts that failure to recognize intrapopulation variability in life history traits has on the perceived reproductive potential of these fish populations.

Section snippets

Collection and analysis of maturity data

Ovaries collected between 1993 and 1998 during spring stratified random bottom trawl surveys of NAFO Divisions 3LNO (Fig. 1) were classed as either juvenile or adult based on the macroscopic criteria of Templeman et al. (1978). In this classification scheme the immature or juvenile stage is reserved for individuals that show no signs of upcoming or previous spawning. All other stages show some evidence of maturing to spawn or of having spawned in the past and are classed as mature (adult). A

Maturation

Analysis of American plaice length at maturity data, combined for all years, revealed a significant difference (χ2 = 503, df = 2, p < 0.0001) between Divisions within the Grand Bank population (Fig. 2). Fish in 3L matured smaller (L50 = 31 cm) than those in 3N and 3O (L50 = 37 cm). Yearly comparisons showed that 3L was different from 3N and 3O in every year, while fish in 3O matured at a smaller size than those in 3N in 1996 and 1998.

Significant spatial differences in size at maturity (χ2 = 21, df = 1, p < 

Discussion

Recognition of natural variability in reproductive parameters for fish populations can greatly affect perceptions about stock reproductive potential (Morgan and Brattey, 2005). Here it was demonstrated that failure to account for intrapopulation differences can also have large consequences for perception of reproductive potential of the whole population. Accounting for this intrapopulation variability suggested a difference in TEP of up to 1014 and 1015 eggs for American plaice and yellowtail

Acknowledgements

We thank the many ships crew and technical staff involved in the collection and processing of these data. Funding for this project was provided by Fisheries and Oceans Canada, Fisheries Products International Ltd., the Canadian Centre for Fisheries Innovation, Memorial University of Newfoundland, and the Fisheries Science Collaborative Program. B. Nakashima and S. Walsh and three anonymous reviewers gave helpful comments on an earlier version of the paper.

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