Size-selective harvesting alters life histories of a temperate sex-changing fish.

Selective mortality, whether caused naturally by predation or through the influence of harvest practices, initiates changes within populations when individuals possessing certain heritable traits have increased fitness. Theory predicts that increased mortality rates will select for changes in a number of different life history characteristics. For example, fishing often targets larger individuals and has been shown repeatedly to alter population size structure and growth rates, and the timing of maturation. For sex-changing species, selective fishing practices can affect additional traits such as the mature population sex ratio and the timing of sexual transformation. Using historical comparisons, we examined the effects of exploitation on life history characteristics of California sheephead, Semicossyphus pulcher, a temperate protogynous (female-male sex changer) labrid that inhabits nearshore rocky environments from central California, USA, to southern Baja California, Mexico. Recreational fishing intensified and an unregulated commercial live-fish fishery developed rapidly in southern California between the historical and current studies. Collections of S. pulcher from three locations (Bahia Tortugas, Catalina Island, and San Nicolas Island) in 1998 were compared with data collected 20-30 years previously to ascertain fishery-induced changes in life history traits. At Bahia Tortugas, where fishing by the artisanal community remained light and annual survivorship stayed high, we observed no changes in size structure or shifts in the timing of maturation or the timing of sex change. In contrast, where recreational (Catalina) and commercial (San Nicolas) fishing intensified and annual survivorship correspondingly declined, males and females shifted significantly to smaller body sizes, females matured earlier and changed sex into males at both smaller sizes and younger ages and appeared to have a reduced maximum lifespan. Mature sex ratios (female:male) increased at San Nicolas, despite a twofold reduction in the mean time spent as a mature female. Proper fisheries management requires measures to prevent sex ratio skew, sperm limitation, and reproductive failure because populations of sequential hermaphrodites are more sensitive to size-selective harvest than separate-sex species. This is especially true for S. pulcher, where different segments of the fishery (commercial vs. recreational) selectively target distinct sizes and therefore sexes in different locations.     

Sex ratio of juvenile loggerhead turtles in the Mediterranean Sea: is it really 1:1?

Sex ratios are a crucial parameter for evaluating population viability. In species with complex life history patterns and temperature sex determination mechanisms, such as the loggerhead turtle (Caretta caretta), sex ratios may vary within a population and among populations. In the Mediterranean, juvenile sex ratios appear to not differ significantly from 1:1, although estimates for hatchling sex ratios are highly female biased. The immigration of males from the Atlantic has been suggested as a possible cause of such variation. Here, we present results of a multi-year investigation (2000–2011) on the sex ratios of loggerhead turtles foraging along the south Tyrrhenian coast, Western Mediterranean, with the aim of providing a better understanding of the potentially underlying forces that drive regional and age-dependent differences in sex ratios. Sex was determined through visual examination of the gonads in 271 dead turtles (curved carapace length range 29.5–89 cm). A fragment of the mitochondrial DNA control region was sequenced from 61 specimens to characterise the demographic composition of this foraging assemblage by applying a many-to-many mixed stock analysis approach. No significant association was found between sex ratios and years or size classes, although the largest size was male biased. Juvenile sex ratio was 1.56:1, which was different from an even sex ratio but still less female biased than hatchling sex ratios from Mediterranean beaches. Results of the mixed stock analysis indicate that juvenile sex ratios in the Mediterranean are largely unaffected by immigration of Atlantic individuals into the basin, as previously suggested. Continued long-term monitoring of juvenile sex ratios is necessary to detect biologically significant sex ratio shifts in the Mediterranean loggerhead turtle population.     

Econometric modeling of fisheries with complex life histories: Avoiding biological management failures

Economics of the fishery has focused on the wastefulness of common pool resource exploitation. Pure open access fisheries dissipate economic rents and degrade biological stocks. Biologically managed fisheries also dissipate rents but are thought to hold biological stocks at desired levels. We develop and estimate an empirical bioeconomic model of the Gulf of Mexico gag fishery that questions the presumptive success of biological management. Unlike previous bioeconomic life history studies, we provide a way to circumvent calibration problems by embedding our estimation routine directly in the dynamic bioeconomic model. We nest a standard biological management model that accounts for complex life history characteristics of the gag. Biological intuition suggests that a spawning season closure will reduce fishing pressure and increase stocks, and simulations of the biological management model confirm this finding. However, simulations of the empirical bioeconomic model suggest that these intended outcomes of the spawning closure do not materialize. The behavioral response to the closure appears to be so pronounced that it offsets the restriction in allowable fishing days. Our results indicate that failure to account for fishing behavior may play an important role in fishery management failures.     

A simulation model to explore the relative value of stock enhancement versus harvest regulations for fishery sustainability

Harvest restrictions and stock enhancement are commonly proposed management responses for sustaining degraded fisheries, but comparisons of their relative effectiveness have seldom been considered prior to making policy choices. We built a population model that incorporated both size-dependent harvest restrictions and stock enhancement contributions to explore trade-offs between minimum length limits and stock enhancement for improving population sustainability and fishery metrics (e.g., catch). We used a Murray cod Maccullochella peelii peelii population as a test case, and the model incorporated density-dependent recruitment processes for both hatchery and wild fish. We estimated the spawning potential ratio (SPR) and fishery metrics (e.g., angler catch) across a range of minimum length limits and stocking rates. Model estimates showed that increased minimum length limits were much more effective than stock enhancement for increasing SPR and angler catches in exploited populations, but length limits resulted in reduced harvest. Stocking was predicted to significantly increase total recruitment, population sustainability, and fishery metrics only in systems where natural reproduction had been greatly reduced via habitat loss, fishing mortality was high, or both. If angler fishing effort increased with increased fish abundance from stocking efforts, fishing mortality was predicted to increase and reduce the benefits realized from stocking. The model also indicated that benefits from stock enhancement would be reduced if reproductive efficiency of hatchery-origin fish was compromised. The simulations indicated that stock enhancement was a less effective method to improve fishery sustainability than measures designed to reduce fishing mortality (e.g., length limits).     

Wide-spread genetic variability and the paradox of effective population size in the gag,Mycteroperca microlepis,along the West Florida Shelf

Wide-ranging marine species are often described as having a low effective population size (N _e) to census size (N) ratio. This genetic phenomenon is typically attributed to large variation among individuals in reproductive success because of the high mortality rates and unpredictable environments associated with larval dispersal. In this study, we examined patterns of genetic variation in gag (Mycteroperca microlepis) on the West Florida Shelf across year classes of post-settlement juveniles and spawning adults. With no significant genetic differentiation among year classes despite varying recruitment dynamics, little evidence for chaotic genetic patchiness, and no truncation of adult genetic diversity in subsequent juvenile cohorts, there was little support for large variation among individual in reproductive success contributing to a low N _e/N ratio. In fact, the consistent lack of significant differences in annual recruitment classes indicated that reproductive success among individuals was resistant to skewing. Among the various evolutionary forces that may be affecting N _e, changes to demography due to fishing pressure are posited as a likely mechanism affecting current levels of genetic variation.     

Diamondback Terrapin (Malaclemys terrapin) Mortality in Crab Pots

The entrapment of diamondback terrapins (  Malaclemys terrapin) in crab pots frequently results in drowning and death of the trapped turtles. We determined the rate of capture, size, sex, and age of terrapins captured, and the potential impact crab pot mortality has on local terrapin populations. We estimated terrapin capture rates of 0.17 terrapins/pot/day in shallow water areas of Chesapeake Bay (Maryland, USA). The sex ratio of terrapins caught in crab pots was 3:2 male biased because female terrapins become too large to enter crab pots by the time they reach 8 years of age. Males, however, remain vulnerable to entrapment throughout their life. Our estimates of capture rates and local population size suggest that 15–78% of a local population may be captured in a single year. As a consequence, crab pots may be the major reason terrapins are extirpated in coastal, shallow water areas with heavy crab pot fisheries. Additionally, the selective removal of males may also contribute to female-biased sex ratios observed in this diamondback terrapin population. We developed and tested a modified crab pot that increases terrapin survival and does not reduce the number of crabs caught. Our modified crab pot maintained permanent access to air and prevented the drowning of terrapins. Additionally, our modified crab pot caught more crabs than standard commercial crab pots, suggesting that the modified crab pot could be a viable alternative to standard traps that result in terrapin mortality.     

Persistent Unequal Sex Ratio in a Population of Grayling (Salmonidae) and Possible Role of Temperature Increase

In some fishes, water chemistry or temperature affects sex determination or creates sex‐specific selection pressures. The resulting population sex ratios are hard to predict from laboratory studies if the environmental triggers interact with other factors, whereas in field studies, singular observations of unusual sex ratios may be particularly prone to selective reporting. Long‐term monitoring largely avoids these problems. We studied a population of grayling (Thymallus thymallus) in Lake Thun, Switzerland, that has been monitored since 1948. Samples of spawning fish have been caught about 3 times/week around spawning season, and water temperature at the spawning site has been continuously recorded since 1970. We used scale samples collected in different years to determine the average age of spawners (for life‐stage specific analyses) and to identify the cohort born in 2003 (an extraordinarily warm year). Recent tissue samples were genotyped on microsatellite markers to test for genetic bottlenecks in the past and to estimate the genetically effective population size (N_e). Operational sex ratios changed from approximately 65% males before 1993 to approximately 85% males from 1993 to 2011. Sex ratios correlated with the water temperatures the fish experienced in their first year of life. Sex ratios were best explained by the average temperature juvenile fish experienced during their first summer. Grayling abundance is declining, but we found no evidence of a strong genetic bottleneck that would explain the apparent lack of evolutionary response to the unequal sex ratio. Results of other studies show no evidence of endocrine disruptors in the study area. Our findings suggest temperature affects population sex ratio and thereby contributes to population decline. Persistencia de Proporción de Sexos Desigual en una Población de Tímalos (Salmonidae) y el Posible Papel del Incremento de la Temperatura     

The role of life histories and trophic interactions in population recovery

Factors affecting population recovery from depletion are at the focus of wildlife management. Particularly, it has been debated how life‐history characteristics might affect population recovery ability and productivity. Many exploited fish stocks have shown temporal changes towards earlier maturation and reduced adult body size, potentially owing to evolutionary responses to fishing. Whereas such life‐history changes have been widely documented, their potential role on stock's ability to recover from exploitation often remains ignored by traditional fisheries management. We used a marine ecosystem model parameterized for Southeastern Australian ecosystem to explore how changes towards “faster” life histories might affect population per capita growth rate r. We show that for most species changes towards earlier maturation during fishing have a negative effect (3–40% decrease) on r during the recovery phase. Faster juvenile growth and earlier maturation were beneficial early in life, but smaller adult body sizes reduced the lifetime reproductive output and increased adult natural mortality. However, both at intra‐ and inter‐specific level natural mortality and trophic position of the species were as important in determining r as species longevity and age of maturation, suggesting that r cannot be predicted from life‐history traits alone. Our study highlights that factors affecting population recovery ability and productivity should be explored in a multi‐species context, where both age‐specific fecundity and survival schedules are addressed simultaneously. It also suggests that contemporary life‐history changes in harvested species are unlikely to increase their resilience and recovery ability.     

A Contemporary,Sex‐Limited Change in Body Size of an Estuarine Turtle in Response to Commercial Fishing

Abstract: Juvenile growth rate and adult body size are important components of life‐history strategies because of their direct impact on fitness. The diamondback terrapin (Malaclemys terrapin) is a sexually dimorphic, long‐lived turtle inhabiting brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of its range, terrapins face anthropogenically imposed mortality: juveniles of both sexes inadvertently enter commercial crab traps and drown. For adult females, the carapace eventually grows large enough that they cannot enter traps, whereas males almost never reach that critical size. We compared age structure, carapace dimensions, growth curves, and indices of sexual dimorphism for a Chesapeake Bay population of terrapins (where mortality of turtles is high due to crab traps) with contemporary terrapins from Long Island Sound and museum specimens from Chesapeake Bay (neither group subject to commercial crab traps). We also calculated the allochronic and synchronic rates of evolutionary change (haldanes) for males and females to measure the rate of trait change in a population or between populations, respectively. We found a dramatic shift to a younger male age structure, a decrease in the length of time to terminal female carapace size, a 15% increase in female carapace width, and an increase in sexual dimorphism in Chesapeake Bay. In a new twist, our results implicate a fishery in the selective increase in size of a reptilian bycatch species. These sex‐specific changes in life history and demography have implications for population viability that need to be considered when addressing conservation of this threatened turtle.     

Density-dependent habitat selection and performance by a large mobile reef fish.

Many exploited reef fish are vulnerable to overfishing because they concentrate over hard-bottom patchy habitats. How mobile reef fish use patchy habitat, and the potential consequences on demographic parameters, must be known for spatially explicit population dynamics modeling, for discriminating essential fish habitat (EFH), and for effectively planning conservation measures (e.g., marine protected areas, stock enhancement, and artificial reefs). Gag, Mycteroperca microlepis, is an ecologically and economically important warm-temperate grouper in the southeastern United States, with behavioral and life history traits conducive to large-scale field experiments. The Suwannee Regional Reef System (SRRS) was built of standard habitat units (SHUs) in 1991-1993 to manipulate and control habitat patchiness and intrinsic habitat quality, and thereby test predictions from habitat selection theory. Colonization of the SRRS by gag over the first six years showed significant interactions of SHU size, spacing, and reef age; with trajectories modeled using a quadratic function for closely spaced SHUs (25 m) and a linear model for widely spaced SHUs (225 m), with larger SHUs (16 standardized cubes) accumulating significantly more gag faster than smaller 4-cube SHUs (mean = 72.5 gag/16-cube SHU at 225-m spacing by year 6, compared to 24.2 gag/4-cube SHU for same spacing and reef age). Residency times (mean = 9.8 mo), indicative of choice and measured by ultrasonic telemetry (1995-1998), showed significant interaction of SHU size and spacing consistent with colonization trajectories. Average relative weight (W(r)) and incremental growth were greater on smaller than larger SHUs (mean W(r) = 104.2 vs. 97.7; incremental growth differed by 15%), contrary to patterns of abundance and residency. Experimental manipulation of shelter on a subset of SRRS sites (2000-2001) confirmed our hypothesis that shelter limits local densities of gag, which, in turn, regulates their growth and condition. Density-dependent habitat selection for shelter and individual growth dynamics were therefore interdependent ecological processes that help to explain how patchy reef habitat sustains gag production. Moreover, gag selected shelter at the expense of maximizing their growth. Thus, mobile reef fishes could experience density-dependent effects on growth, survival, and/or reproduction (i.e., demographic parameters) despite reduced stock sizes as a consequence of fishing.     

Management and Recovery Options for Ural River Beluga Sturgeon

Abstract: Management of declining fisheries of anadromous species sometimes relies heavily on supplementation of populations with captive breeding, despite evidence that captive breeding can have negative consequences and may not address the root cause of decline. The beluga sturgeon (Huso huso), a species threatened by the market for black caviar and reductions in habitat quality, is managed through harvest control and hatchery supplementation, with an emphasis on the latter. We used yield per recruit and elasticity analyses to evaluate the population status and current levels of fishing and to identify the life‐history stages that are the best targets for conservation of beluga of the Ural River. Harvest rates in recent years were four to five times higher than rates that would sustain population abundance. Sustainable rates of fishing mortality are similar to those for other long‐lived marine species such as sharks and mammals. Yield per recruit, which is maximized if fish are first harvested at age 31 years, would be greatly enhanced by raising minimum size limits or reducing illegal take of subadults. Improving the survival of subadult and adult females would increase population productivity by 10 times that achieved by improving fecundity and survival from egg to age 1 year (i.e., hatchery supplementation). These results suggest that reducing mortality of subadults and adult wild fish is a more effective conservation strategy than hatchery supplementation. Because genetics is not factored into hatchery management practices, supplementation may even reduce the viability of the beluga sturgeon.     

The interplay of sex ratio, male success and density-independent mortality affects population dynamics

Environmental constraints can limit a population to a certain size, which is usually called the carrying capacity of a habitat. Besides to this ‘external’ factor, which is mainly determined by the limitation of resources, we investigate here another set of population-intrinsic factors that can limit a population size significantly below the maximum sustainable size. Firstly, density-independent mortality is a prominent factor in all organisms that show age-related and/or accidental death. Secondly, in sexually reproducing organisms the sex ratio and the success of pairing is important for finding reproductive partners. Using a simple model, we demonstrate how sex ratio, mating success and gender-specific mortality can strongly affect the speed of population growth and the maximum population size. In addition, we demonstrate that density-independent mortality, which is often neglected in population models, adds a very important feature to the system: it strongly enhances the negative influence of unbiased sex ratios and inefficient pairing to the maximum sustainable population size. A decrease of the maximum population size significantly affects a population's survival chance in inter-specific competition. Thus, we conclude that the inclusion of density-independent mortality is crucial, especially for models of species that reproduce sexually. We show that density-independent mortality, together with biased sex ratios, can significantly lower the abilities of a population to survive in conditions of strong inter-specific competition and due to the Allee effect. We emphasize that population models should incorporate the sex ratio, male success and density-independent mortality to make plausible predictions of the population dynamics in a gender-structured population. We show that the population size is limited by these intrinsic factors. This is of high ecological significance, because it means that there will always be resources available in any habitat that allows other species (e.g., invaders) to use these resources and settle successfully, if they are sufficiently adapted.     

Development and drift of northern shrimp larvae (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) at West Greenland

This is the first study of the West Greenland offshore population of Pandalus borealis in recent history that covers all larval stages. Shrimp larvae were sampled on the fishing banks off the west coast of Greenland from 63.5°N to 67°N in May, June and July. Abundances decreased during the summer as did cumulated mortality rates [0.06 day^–1 (ZI) to 0.04 day^–1 (ZVI)]. Modelling stage development time as a function of temperature alone by means of the Blehrádek function gave decreasing stage durations from 22.7 to 16.7 days. Drift buoys showed a northbound current with an average velocity of 0.06 m s^–1. Potential spawning grounds of shrimp larvae were located from back-calculation by coupling development times and mortality rates with current velocity. This showed larval transport of up to 500 km. The adult female shrimp abundances were estimated to 0.12–96 females per 100 m², and locations of the estimated spawning stock agreed with observations from trawl surveys.Communicated by L. Hagerman, Helsingør     

Sex ratio and optimal harvest of canvasback ducks,A model

The North American population of canvasback ducks (Aythya valisineria) exhibits extreme distortion of the sex ratio in favor of males. This paper describes a model which accounts for this pattern by relatively heavier female mortality in both the breeding and nonbreeding seasons. The density-dependence of winter mortality leads to the conclusion that the observed sex ratio depresses total population numbers. Variation in nesting success is shown to influence sex ratios and strongly depress population numbers. Because a standard harvest scheme can be demonstrated to severely depress the numbers of ducks, an alternative graduated or weighted harvest procedure is recommended.     

Possible Extinction Vortex for a Population of Iberian Lynx on the Verge of Extirpation

Abstract: Theory suggests that demographic and genetic traits deteriorate (i.e., fitness and genetic diversity decrease) when populations become small, and that such deterioration could precipitate positive feedback loops called extinction vortices. We examined whether demographic attributes and genetic traits have changed over time in one of the 2 remaining small populations of the highly endangered Iberian lynx (Lynx pardinus) in Doñana, Spain. From 1983 to 2008, we recorded nontraumatic mortality rates, litter size, offspring survival, age at territory acquisition, and sex ratio. We combined these demographic attributes with measures of inbreeding and genetic diversity at neutral loci (microsatellites) and genes subjected to selection (major histocompatibility complex). Data on demographic traits were obtained through capture and radio tracking, checking dens during breeding, track surveys, and camera trapping. For genetic analyses, we obtained blood or tissue samples from captured or necropsied individuals or from museum specimens. Over time a female‐biased sex ratio developed, age of territory acquisition decreased, mean litter size decreased, and rates of nontraumatic mortality increased, but there were no significant changes in overall mortality rates, standardized individual heterozygosity declined steadily, and allelic diversity of exon 2 of class II major histocompatibility complex DRB genes remained constant (2 allelic variants present in all individuals analyzed). Changes in sex ratio and age of territory acquisition may have resulted from demographic stochasticity, whereas changes in litter size and nontraumatic mortality may be related to observed increases in inbreeding. Concomitant deterioration of both demographic attributes and genetic traits is consistent with an extinction vortex. The co‐occurrence, with or without interaction, of demographic and genetic deterioration may explain the lack of success of conservation efforts with the Doñana population of Iberian lynx.     

Population Consequences of Environmental Sex Reversal

Abstract:  When sex determination in a species is predominantly genetic but environmentally reversible, exposure to (anthropogenic) changes in the environment can lead to shifts in a population's sex ratio. Such scenarios may be common in many fishes and amphibians, yet their ramifications remain largely unexplored. We used a simple model to study the (short-term) population consequences of environmental sex reversal (ESR). We examined the effects on sex ratios, sex chromosome frequencies, and population growth and persistence after exposure to environmental forces with feminizing or masculinizing tendencies. When environmental feminization was strong, X chromosomes were driven to extinction. Analogously, extinction of normally male-linked genetic factors (e.g., Y chromosomes) was caused by continuous environmental masculinization. Although moderate feminization was beneficial for population growth in the absence of large viability effects, our results suggest that the consequences of ESR are generally negative in terms of population size and the persistence of sex chromosomes. Extreme sex ratios resulting from high rates of ESR also reduced effective population sizes considerably. This may limit any evolutionary response to the deleterious effects of ESR. Our findings suggest that ESR changes population growth and sex ratios in some counter-intuitive ways and can change the predominant factor in sex determination from genetic to fully environmental, often within only a few tens of generations. Populations that lose genetic sex determination may quickly go extinct if the environmental forces that cause sex reversal cease.     

Effects of spatial variability and colony size on the reproductive output and gonadal development cycle of the Mediterranean red coral (Corallium rubrum L.)

Red coral (Corallium rubrum, L. 1758) is an over-exploited Mediterranean gorgonian. The gonadal development cycle of this gorgonian is examined at the Costa Brava (NW Mediterranean) taking into account for the first time colony size, depth and spatial horizontal variability. This study compares the gonad development and fertility in two colony size classes (colonies <6-cm height, and >10-cm height, both at 40–45-m depth), and two populations at different depths (16–18-m depth, and 40–45-m depth, both consisting of <6-cm high colonies) in a 15-month period. The fertility of seven size classes (<2 cm to >12 cm high colonies, in 2 cm intervals) was examined in the deep population, where large colonies were present. Furthermore, reproductive output was compared in 6 populations (distributed along more than 70-km coastline) one month before spawning (June). Red coral was found to be dioecious and gonochoric with a sex ratio of 1:1, which differs from other NW Mediterranean populations. On the other hand, fertility of different size classes indicates that small colonies of 2-cm height already produce gonads, which is in line with previous studies. Female and male polyp fertility and sperm sac size increase significantly with colony size [sperm sac diameter: 476±144 μm (mean±SD) and 305±150 μm in the >10-cm and <6-cm height colonies, respectively), whereas no significant effect on oocyte diameter was found (oocyte diameter: 373.7±18.7 μm). Depth staggered spawning, that is, an earlier release of gonads in the shallow populations, was observed in summer 2003, coinciding with the highest temperature gradient between shallow and deep water during the study period. Colonies of <6-cm height were significantly less fertile than colonies >12 cm, thus the recommendation of this study is that a minimum height should be incorporated into fishing regulations. The six studied populations at the Costa Brava showed a comparable reproductive potential, which demonstrates little variability within the homogenous population structure and range of size classes (due to overharvesting) found at the Costa Brava. The study of reproductive output is an important tool for ecosystem management, and this work recommends basing specific exploitation laws for distinctive populations on colony size, which is found to have a larger effect on reproductive potential than mesoscale variability. An erratum to this article can be found at     

Climate and predation dominate juvenile and adult recruitment in a turtle with temperature-dependent sex determination

Conditions experienced early in life can influence phenotypes in ecologically important ways, as exemplified by organisms with environmental sex determination. For organisms with temperature-dependent sex determination (TSD), variation in nest temperatures induces phenotypic variation that could impact population growth rates. In environments that vary over space and time, how does this variation influence key demographic parameters (cohort sex ratio and hatchling recruitment) in early life stages of populations exhibiting TSD? We leverage a 17-year data set on a population of painted turtles, Chrysemys picta, to investigate how spatial variation in nest vegetation cover and temporal variation in climate influence early life-history demography. We found that spatial variation in nest cover strongly influenced nest temperature and sex ratio, but was not correlated with clutch size, nest predation, total nest failure, or hatching success. Temporal variation in climate influenced percentage of total nest failure and cohort sex ratio, but not depredation rate, mean clutch size, or mean hatching success. Total hatchling recruitment in a year was influenced primarily by temporal variation in climate-independent factors, number of nests constructed, and depredation rate. Recruitment of female hatchlings was determined by stochastic variation in nest depredation and annual climate and also by the total nest production. Overall population demography depends more strongly on annual variation in climate and predation than it does on the intricacies of nest-specific biology. Finally, we demonstrate that recruitment of female hatchlings translates into recruitment of breeding females into the population, thus linking climate (and other) effects on early life stages to adult demographics.     

A General Model for Analyzing Data from Mark-recapture Experiments with an Application to the Pacific Halibut

A general model is developed to examine the patterns of the regional movement of tagged and released fish from mark-recapture experiments. It is a stochastic model that incorporates fishing mortality, natural mortality, fish movement, tag-shedding, and different rates of reporting. A likelihood function is constructed for estimating its parameters. We used this model to analyze data on the Pacific halibut from mark-recapture experiments conducted by the International Pacific Halibut Commission (IPHC), with a total of 36,058 releases from 1982 to 1986 and 5,826 recoveries from 1982 to 2000. We estimated their rates of movement among IPHC management areas, along with their instantaneous rates of natural and fishing mortalities. Our analysis revealed that fish movement was not significant among areas, with a resident probability of > 0.92. This lends support to the IPHC catch-at-age stock assessment model (which has no built-in movement components). The estimated instantaneous rate of natural mortality (0.198 year⁻¹) lies between that assumed in all IPHC stock assessments before 1998 (0.20 year⁻¹) and that from 1999 onwards (0.15 year⁻¹). The estimates of the instantaneous rates of fishing mortality were consistent with those from the IPHC stock assessment model. Received: April 2003 / Revised: May 2005     

Invasion dynamics of the varnish clam (Nuttallia obscurata): a matrix demographic modeling approach

Marine invaders have become a significant threat to native biodiversity and ecosystem function. In this study, the invasion of the varnish clam (Nuttallia obscurata) in British Columbia, Canada, is investigated using a matrix modeling approach to identify the life history characteristics most crucial for population growth and to investigate population differences. Mark-recapture analyses and field collections from 2003 to 2004 were used to determine individual growth, survival rates, and fecundity for two sites. A multi-state matrix model was used to determine population growth rates and to conduct sensitivity and elasticity analyses. A life table response experiment was also used to determine what life history stage contributed most to observed differences in population growth rates. Population survey data were used in conjunction with the matrix model to determine plausible recruitment levels and to investigate recruitment scenarios. Both populations are currently declining but are likely sustainable because of the pulsed nature of large recruitment events. Survival of larger clams (>40 mm) is the most important for population growth based on elasticity and sensitivity analyses. Adult survival also had the largest influence on observed differences between site-specific population growth rates. The two populations studied differed in recruitment dynamics; one experiencing annual recruitment with higher post-settlement mortality and the other, episodic recruitment and lower post-settlement mortality. The most influential factor for the successful invasion of the varnish clam appears to be survival of the larger size classes. Therefore, any process that decreases adult survival (e.g., predation, commercial harvest) will have the greatest impact on population growth.