Estimation of some parameters of large fish populations by capture-recapture experiments

A certain number of tagged fish is liberated and assumed to be distributed randomly among a natural fish population. The fish are subjected to a number of fishing experiments within relatively short periods, and lie between equal intervening periods of durationT. Untagged fish are retained, while tagged fish are released during the fishing experiments. Denoting the catchability of untagged fish byq _u and that for tagged fish byq _t , it is assumed that they are related by the equation “q _u =cq _t ” wherec is a constant. Denoting the survival rates of tagged fish and the effective fishing effort of commerical fisheries per unit time from the (k-1)^th to thek ^th experiments by _t S _k andf _k , respectively, it is assumed that they vary from period to period. Assuming that during thek ^th experiment, the number of untagged fish captured and the experimental fishing rate of tagged fish are denoted by _u X _k and _t P _k , respectively, then $$\begin{array}{*{20}c} {\frac{{(_u X_k )^2 }}{{[_u X_{(k - 1)} ][_u X_{(k + 1)} ]}} = \frac{{_t S_k }}{{_t S_{(k + 1)} }} \cdot \frac{{e^{ - (1 - c)q_t f_{(k + 1)} T} }}{{e^{ - (1 - c)q_t f_k T} }} \cdot } \\ {\frac{{1 - c_t P_{(k - 1)} }}{{1 - c_t P_k }} \cdot \frac{{(_t P_k )^2 }}{{[_t P_{(k - 1)} ][_t P_{(k + 1)} ]}}.} \\ \end{array}$$ The above equation containsc as a single unknown, while all other terms are supplied by the capture-recapture experiments, exceptf _k andf _(k+1) which may be obtained from fisheries statistics. A number of the above equations are obtained from several experiments and can be combined into a single equation to obtain an overall estimate forc which can be used to derive estimates for experimental fishing rates, abundance, and instantaneous natural and fishing mortality rates for natural fish populations. These estimates are free from type (A) tagging errors, and have the advantage of taking into consideration the probable different behaviour of tagged and untagged fish.     

Estimation of abundance of fish populations by capture-recapture experiments

A natural fish population is assumed to be subjected to a number of fishing experiments, during which the captured fish are released after tagging all untagged fish. Assuming that, in thek ^th sampling survey, the tagged and untagged fish are subjected toP _tk andP _uk fishing rates, andS _tk andS _uk survival rates, respectively, the population size can be estimated from an estimate ofP _uk $$\begin{gathered} i.e., 1 - P_{uk} = \hfill \\ P_{tk \cdot } \frac{{S_t (k + 1)}}{{S_u (k + 1)}}\frac{\begin{gathered} Number of fish captured for the first time in \hfill \\ the (k + 1)^{th} survey plus fish belonging to this \hfill \\ group captured later \hfill \\ \end{gathered} }{\begin{gathered} Number of fish captured for the first time in \hfill \\ the k^{th} and captured again in the (k + 1)^{th} \hfill \\ \exp eriment plus fish belonging to this group \hfill \\ which are captured later \hfill \\ \end{gathered} } \hfill \\ \end{gathered}$$ All terms of the right hand side of the above equation are provided by capture-recapture experiments, except the survival rate of untagged fish which may be obtained by other information. This estimate of fishing rate of untagged fish is free from type (A) errors.     

Interactions between cod, herring and sprat in the changing environment of the Baltic Sea: A dynamic model analysis

The interactions between cod (Gadus morhua), herring (Clupea harengus) and sprat (Sprattus sprattus) in the Central Baltic Sea were examined with a simple dynamic model, an alternative to more complicated and data-demanding multispecies and ecosystem models. The main aims of the study were to compare the effect of alternative structures on the model output and examine the control relationships in the fish assemblage under different environmental conditions. The effect of environmental conditions was modelled using a stock-recruitment equation for cod incorporating an environmental index. The model output was especially sensitive to the functional response in predation by cod on herring and sprat. The type II functional response led to a collapse of the clupeid stocks when cod was abundant, while the type III response produced more realistic stock dynamics. According to the simulations, an abundant cod stock was able to keep the sprat stock at a low level, while the herring stock was less affected and benefited from the decreased density of sprat. Simulation of different fishing scenarios indicated that reducing fishing mortality to the level currently advised by ICES would allow the recovery of the cod stock even in unfavourable environmental conditions.     

Isotopic evidences for microbiologically mediated and direct C input to soil compounds from three different leaf litters during their decomposition

We show the potentiality of coupling together different compound-specific isotopic analyses in a laboratory experiment, where ¹³C-depleted leaf litter was incubated on a ¹³C-enriched soil. The aim of our study was to identify the soil compounds where the C derived from three different litter species is retained. Three ¹³C-depleted leaf litter (Liquidambar styraciflua L., Cercis canadensis L. and Pinus taeda L., δ¹³C_vsPDB ≈ ?43‰), differing in their degradability, were incubated on a C4 soil (δ¹³C_vsPDB ≈ ?18‰) under laboratory-controlled conditions for 8 months. At harvest, compound-specific isotope analyses were performed on different classes of soil compounds [i.e. phospholipids fatty acids (PLFAs), n-alkanes and soil pyrolysis products]. Linoleic acid (PLFA 18:2ω6,9) was found to be very depleted in ¹³C (δ¹³C_vsPDB ≈ from ?38 to ?42‰) compared to all other PLFAs (δ¹³C_vsPDB ≈ from ?14 to ?35‰). Because of this, fungi were identified as the first among microbes to use the litter as source of C. Among n-alkanes, long-chain (C27–C31) n-alkanes were the only to have a depleted δ¹³C. This is an indication that not all of the C derived from litter in the soil was transformed by microbes. The depletion in ¹³C was also found in different classes of pyrolysis products, suggesting that the litter-derived C is incorporated in less or more chemically stable compounds, even only after 8 months decomposition.     

Parameter estimation for distributed delay based population models from laboratory data: egg hatching of Oulema duftschmidi Redthenbacher (Coleoptera,Chrysomelidae) as an example

The cohort development of poikilotherms under favorable temperature conditions may be described by the time-invariant distributed delay models that require three parameters, i.e. the thermal threshold T₀, the thermal constant W and the variability parameter H representing distributed maturation times, also known as time distributed or stochastic development. Here, a parsimonious method is developed that uses stage-frequency matrices obtained under controlled conditions. The analysis of these matrices permits the estimation of the three parameters, while the incorporation of both developmental threshold and thermal constant into the time-invariant distributed delay model permits the representation of stochastic cohort development under constant temperatures. The evaluated parameters can also be used in time-varying distributed delay models that are particularly useful for a wide range of fluctuating temperature conditions.We consider stage-frequency matrices obtained from cohorts of eggs and larvae of Oulema duftschmidi Redthenbacher at different constant temperatures. A visual inspection of the matrices clearly shows the temperature dependent as well as the time distributed passage from the egg to the larval stage. Under the current range of temperatures, a linear model for developmental rate satisfactorily describes egg development, and estimates the thermal threshold (T₀=11.2 °C), the thermal constant (W=81.3 day-degrees) and the delay order parameter (H=70). The resulting model can be used to represent the development of cohorts of O. duftschmidi eggs in a favorable temperature range.     

Tolerance of Hyas araneus zoea I larvae to elevated seawater PCO2 despite elevated metabolic costs

Early life stages of marine crustaceans respond sensitively to elevated seawater PCO₂. However, the underlying physiological mechanisms have not been studied well. We therefore investigated the effects of elevated seawater PCO₂ on oxygen consumption, dry weight, elemental composition, median developmental time (MDT) and mortality in zoea I larvae of the spider crab Hyas araneus (Svalbard 79°N/11°E; collection, May 2009; hatch, December 2009). At the time of moulting, oxygen consumption rate had reached a steady state level under control conditions. In contrast, elevated seawater PCO₂ caused the metabolic rate to rise continuously leading to a maximum 1.5-fold increase beyond control level a few days before moulting into the second stage (zoea II), followed by a pronounced decrease. Dry weight of larvae reared under high CO₂ conditions was lower than in control larvae at the beginning of the moult cycle, yet this difference had disappeared at the time of moulting. MDT of zoea I varied between 45 ± 1 days under control conditions and 42 ± 2 days under the highest seawater CO₂ concentration. The present study indicates that larval development under elevated seawater PCO₂ levels results in higher metabolic costs during premoulting events in zoea I. However, H. araneus zoea I larvae seem to be able to compensate for higher metabolic costs as larval MDT and survival was not affected by elevated PCO₂ levels.     

A general bioeconomic simulation model for annual-crop marine fisheries

A generalized bioeconomic simulation model of annual-crop marine fisheries is described and its use in marine fisheries management is demonstrated. The biological submodel represents the recruitment of new organisms into the fishery, the movement of organisms from one fishing area to another and from one depth to another, the growth of organisms and the mortality of organisms resulting both from natural causes and from fishing. The economic submodel represents the fishing effort exerted on each resource species, the monetary costs of fishing, the value of the harvest and the rent (or excess profits) to the fishery.Basic dynamics of the model results from changes in the number of organisms in the fishery over time, which can be summarized as a set of difference equations of the general form ΔN/Δt = R + I ? E ? M ? F where ΔN/Δt is the net change in number of organisms in the fishery over time, R is recruitment, I is immigration, E is emigration, M is natural mortality and F is fishing mortality. R is a driving variable, whereas I, E, M and F are functions of the state of the system at any given point in time. The model can be run in a deterministic or stochastic mode. Values for parameters affecting rates of recruitment, movement, growth, natural mortality and fishing mortality can be selected from uniform, triangular or normal distributions.Use of the model within a fisheries-management framework is demonstrated by evaluating several management alternatives for the pink shrimp (Penaeus duorarum) fishery on the Tortugas grounds in the Gulf of Mexico. Steps involved in use of the model, including parameterization, validation, sensitivity analysis and stochastic simulations of management policies, are explained.     

Modelling N mineralization from green manure and farmyard manure from a laboratory incubation study

Predicting N mineralization from organic manures like farmyard manure (FYM) is more difficult than from fresh organic materials like crop residues, as the manures vary greatly in composition. A laboratory incubation experiment was carried out for 98 days at 30 °C under aerobic conditions to study the effects on N dynamics of Gliricidia (Gliricidia sepium, Jacquin) and FYM application to soil at 5 and 10 g kg⁻¹. Application of Gliricidia induced N mineralization from the start of incubation period, with the amount of N mineralized increasing with rate of application. In contrast, application of FYM resulted in immobilization of mineral N in soil, irrespective of the rate of application. The initial net immobilization from FYM was limited by availability of N in the soil for the higher rate of application.We used the APSIM SoilN module to simulate these contrasting patterns of mineralization of N from Gliricidia and from FYM. The prediction of N mineralized from Gliricidia was better than FYM. The default model parameters specify that the fresh organic matter pools (FPOOL1, FPOOL2 and FPOOL3) have the same C:N ratio and this assumption was ineffective in predicting N mineralized from FYM. The predictive ability of the model improved when this default assumption was modified based on the size of the individual pools (FPOOL1, FPOOL2 and FPOOL3), and the pool's C:N ratios. The modelling efficiency, a measure of goodness of fit between the simulated and observed data, improved markedly for the modified model. The discrepancy between the modelled and observed data was a tendency for the model to underestimate the rate of re-mineralization at the lower rate of application of FYM in the later part of incubation. Unfortunately the appropriate modification to the size and C:N ratios of the FPOOLs could not be determined on the basis of chemical analysis alone. Thus, a true predictive application of the model to a new FYM material is not yet possible.     

Dynamics of a chemostat with three competitive hydrogen oxidizing denitrifying microbial populations and their efficiency for denitrification

In this work, competition for two nitrogen resources (nitrate-, nitrite-nitrogen) between three hydrogen oxidizing denitrifying populations (Acidovorax sp. strain Ic3 (X₁), Paracoccus sp. strain Ic1 (X₂), and Acinetobacter sp. strain Ic2 (X₃)) was examined. The dynamics of three systems of microbial populations (system I: X₁ − X₃, system II: X₂ − X₃, and system III: X₁ − X₂ − X₃), grown in a chemostat, was studied using bifurcation analysis. The chemostat is the most common type of biological reactor used for the study of microbial growth under controlled conditions. The effect of the operating parameters (i.e., dilution rate and feed nitrate nitrogen concentration) on the long-term behavior of the systems showed that X₃ was the predominant population for a wide range of combinations of dilution rate and feed nitrate nitrogen concentration. Also, coexistence of two populations (X₂X₃, X₁X₃) was observed. The results of the bifurcation analysis were also used to determine the denitrification rate and the nitrite nitrogen accumulation for each of the three systems as a function of the dilution rate (up to 0.17 h⁻¹) and the feed nitrate nitrogen concentration (up to 300 mg/L). The highest denitrification rate was achieved by system I (28 mg/Lh). A comparison between the three systems showed that the nitrite nitrogen concentration in system I was less than the one in system III, while the two systems gave similar denitrification rates. The second system had the greatest accumulation of nitrites with the lowest denitrification rate.     

Introducing effects of temperature and CO2 elevation on tree growth into a statistical growth and yield model

Impacts of elevated temperature and CO₂ on tree growth were introduced into a statistical growth and yield model for Finnish conditions based on corresponding predictions obtained from a physiological growth model. This one-way link between models was made by means of species-specific transfer functions describing the increase in stem volume growth of trees as a function of elevated temperature and CO₂, stand density and the tree's competition status in a stand of Scots pine (Pinus sylvestris), silver birch (Betula pendula) and Norway spruce (Picea abies). This method allows the inner dynamics of the statistical model to be followed when the impacts of temperature and CO₂ elevation on tree growth are introduced into the calculation of volume growth and further allocated between diameter and height growth. In this way compatibility with previous predictions of tree growth by means of statistical models and related model systems under current climatic conditions could be retained.The performance of the statistical model with species-specific transfer functions was evaluated by comparing its predictions with corresponding predictions given by a physiological model under conditions of elevated temperature and CO₂. These calculations revealed that the growth response of individual trees to elevated temperature and CO₂ can be introduced into the statistical model from a physiological growth model with an outcome that results in fairly satisfactory growth responses at the stand level as well.     

Different proxies for the reactivity of aquatic sediments towards oxygen: A model assessment

Information on benthic carbon mineralization rates is often derived from the analysis of oxygen microprofiles in sediments. To enable a direct comparison of different sediment environments, it is often desirable to characterize sediments by a single proxy that expresses their “reactivity” towards oxygen. For this, there are three commonly used proxies: the oxygen penetration depth (OPD), the oxygen flux at the sediment-water interface (DOU), and the maximum volumetric oxygen consumption rate (R_max). The OPD can be directly determined from the oxygen depth profile, while the DOU is usually obtained by a linear fit to the oxygen gradient either in diffusive boundary layer. The oxygen consumption rate R_max requires the fitting of a reactive-transport model to the data profile. This article shows that the OPD alone is a suboptimal proxy, because it shows a strong dependence on the half-saturation constant K_s, and secondly, because it is sensitive to the particular re-oxidation conditions right above the oxic-anoxic interface. Similarly, the volumetric oxygen consumption rate R_max is rather strongly dependent on the kinetic model formulation employed. To show this we fitted three different (Bouldin, Blackman and Monod) kinetics to the same oxygen data profiles. When fitting these models, the R_max values obtained differed by 20% for exactly the same oxygen profile. Accordingly, if one reports R_max values, it is crucial to specify the kinetic model alongside. Overall, DOU emerges as sediment reactivity proxy which is the least model dependent.     

Interaction and independence on methane oxidation of landfill cover soil among three impact factors: water, oxygen and ammonium

To understand the influence patterns and interactions of three important environmental factors, i.e. soil water content, oxygen concentration, and ammonium addition, on methane oxidation, the soils from landfill cover layers were incubated under full factorial parameter settings. In addition to the methane oxidation rate, the quantities and community structures of methanotrophs were analyzed to determine the methane oxidation capacity of the soils. Canonical correspondence analysis was utilized to distinguish the important impact factors. Water content was found to be the most important factor influencing the methane oxidation rate and Type II methanotrophs, and the optimum value was 15% (w/w), which induced methane oxidation rates 10- and 6- times greater than those observed at 5% (w/w) and 20% (w/w), respectively. Ambient oxygen conditions were more suitable for methane oxidation than 3% oxygen. The addition of 100 mg-N·kg _drysoil ^?1 of ammonium induced different effects on methane oxidation capacity when conducted at low or high water content. With regard to the methanotrophs, Type II was sensitive to the changes of water content, while Type I was influenced by oxygen content. Furthermore, the methanotrophic acidophile, Verrucomicrobia, was detected in soils with a pH of 4.9, which extended their known living environments.     

The role of male harassment on female fitness for the dengue vector mosquito Aedes aegypti

Sexual harassment studies in insects suggest that females can incur several kinds of costs from male harassment and mating. Here, we examined direct and indirect costs of male harassment on components of female fitness in the predominantly monandrous mosquito Aedes aegypti. To disentangle the costs of harassment versus the costs of mating, we held females at a low or high density with males whose claspers were modified to prevent insemination and compared these to females held with normal males and to those held with females or alone. A reduced longevity was observed when females were held under high-density conditions with males or females, regardless if male claspers had been modified. There was no consistent effect of harassment on female fecundity. Net reproductive rate (R ₀) was higher in females held at low density with normal males compared to females held with males in the other treatments, even though only a small number of females showed direct evidence of remating. Indirect costs and benefits that were not due to harassment alone were observed. Daughters of females held with normal males at high density had reduced longevity compared to daughters from females held without conspecifics. However, their fitness (R ₀) was higher compared to females in all other treatments. Overall, our results indicate that A. aegypti females do not suffer a fitness cost from harassment of males when kept at moderate densities, and they suggest the potential for benefits obtained from ejaculate components.     

Integrating soil carbon cycling with that of nitrogen and phosphorus in the watershed model SWAT: Theory and model testing

In this paper we describe and test a sub-model that integrates the cycling of carbon (C), nitrogen (N) and phosphorus (P) in the Soil Water Assessment Tool (SWAT) watershed model. The core of the sub-model is a multi-layer, one-pool soil organic carbon (S_C) algorithm, in which the decomposition rate of S_C and input rate to S_C (through decomposition and humification of residues) depend on the current size of S_C. The organic N and P fluxes are coupled to that of C and depend on the available mineral N and P, and the C:N and N:P ratios of the decomposing pools. Tillage explicitly affects the soil organic matter turnover rate through tool-specific coefficients. Unlike most models, the turnover of soil organic matter does not follow first order kinetics. Each soil layer has a specific maximum capacity to accumulate C or C saturation (S_x) that depends on texture and controls the turnover rate. It is shown in an analytical solution that S_x is a parameter with major influence in the model C dynamics. Testing with a 65-yr data set from the dryland wheat growing region in Oregon shows that the model adequately simulates the S_C dynamics in the topsoil (top 0.3 m) for three different treatments. Three key model parameters, the optimal decomposition and humification rates and a factor controlling the effect of soil moisture and temperature on the decomposition rate, showed low uncertainty as determined by generalized likelihood uncertainty estimation. Nonetheless, the parameter set that provided accurate simulations in the topsoil tended to overestimate S_C in the subsoil, suggesting that a mechanism that expresses at depth might not be represented in the current sub-model structure. The explicit integration of C, N, and P fluxes allows for a more cohesive simulation of nutrient cycling in the SWAT model. The sub-model has to be tested in forestland and rangeland in addition to agricultural land, and in diverse soils with extreme properties such high or low pH, an organic horizon, or volcanic soils.     

Kinetics of the oxidation of endocrine disruptor nonylphenol by ferrate(VI)

The kinetics of the oxidation of endocrine disruptor nonylphenol (NP) by potassium ferrate(VI) (K₂FeO₄) in water as a function of pH 8.0–10.9 at 25°C is presented. The observed second-order rate constants, k _obs, decrease with an increase in pH 269–32 M^?1 s^?1. The speciation of Fe(VI) (HFeO ₄ ^? and FeO ₄ ^2? ) and NP (NP–OH and NP–O^?) species was used to explain the pH dependence of the k _obs values. At a dose of 10 mg L^?1 (50 μM) K₂FeO₄, the half-life for the removal of NP by Fe(VI), under water treatment conditions, is less than 1 min.     

Effects of feeding and light intensity on the response of the coral Porites rus to ocean acidification

Recently, it has been suggested that there are conditions under which some coral species appear to be resistant to the effects of ocean acidification. To test if such resistance can be explained by environmental factors such as light and food availability, the present study investigated the effect of 3 feeding regimes crossed with 2 light levels on the response of the coral Porites rus to 2 levels of pCO₂ at 28 °C. After 1, 2, and 3 weeks of incubation under experimental conditions, none of the factors—including pCO₂—significantly affected area-normalized calcification and biomass-normalized calcification. Biomass also was unaffected during the first 2 weeks, but after 3 weeks, corals that were fed had more biomass per unit area than starved corals. These results suggest that P. rus is resistant to short-term exposure to high pCO₂, regardless of food availability and light intensity. P. rus might therefore represent a model system for exploring the genetic basis of tolerance to OA.     

A stochastic simulation model of brown shrimp (Penaeus aztecus ives) growth,movement, and survival in Galveston Bay,Texas

A stochastic simulation model of brown shrimp (Penaeus aztecus Ives) population dynamics in Galveston Bay, Texas, is described, validated, and used to evaluate the effects of management alternatives and changing environmental conditions on shrimp dynamics. The model is composed of submodels representing: (1) recruitment, (2) growth, (3) natural mortality, (4) fishing mortality, and (5) emigration of brown shrimp. The model predicts significant changes in total annual harvest from the food shrimp, bait, and recreational fisheries resulting from (1) closure of the bay system to all fishing except during the spring and fall open seasons, (2) two-week postponement of the opening and closing of the open seasons for the food shrimp fishery, (3) a 2.5°C increase and (4) a 2.5°C decrease in mean water temperature, (5) an 80% increase and (6) an 80% decrease in fishing effort. No significant change in the total annual harvest is predicted when the food shrimp fishing season is extended from May 15 through December 15. Sensitivity analysis suggests that field experimentation designed specifically to test the hypothesis of a 60-day time lag between brown shrimp recruitment into the bays and exposure to the fishery should receive high priority. Simulation results are discussed within a management framework.     

Testing for relationships between individual crab behavior and metabolic rate across ecological contexts

Two classes of explanations for covariation between activity level and metabolic rate among conspecifics have been proposed. First, individual-level variation in activity exhibited during the measurement of metabolic rate should covary with routine metabolic rate because movement increases respiration (a methodological relationship). Second, energetic-based hypotheses posit relationships between individual activity measured under more natural conditions and baseline measures of metabolic rate, among other behavioral, physiological, and life-history traits (functional relationships). Here, we examined these potential relationships between individual behavior and metabolic rate in the mud crab (Panopeus herbstii). Specifically, we tested for (1) an effect of crab activity in metabolic chambers (activity_chamber) on routine metabolic rate (RMR), and (2) an effect of crab activity in mesocosms that mimicked field conditions (activity_mesocosm) on standard metabolic rate (SMR). To test for context dependence, we assessed both activity-metabolic rate relationships in the absence and presence of predation threat from toadfish (Opsanus tau) in the form of waterborne chemical cues. Individual variation in activity_mesocosm and RMR was repeatable over time. In support of a methodological relationship, individual differences in RMR were partially explained by crab activity_chamber. After accounting for this methodological relationship, individual SMR was inversely related to activity_mesocosm, supporting an allocation model that predicts behavior and baseline metabolic rate compete for finite energy reserves. We found no evidence of context dependence in either activity-metabolic rate relationship. Thus, our study emphasizes the importance of considering methodological artifacts in elucidating functional relationships between individual behavior and energetics.     

Calculation and use of selectivity coefficients of feeding: Zooplankton grazing

A straightforward method for calculating selectivity coefficients (W_ij) of predation from raw data, mortality rates of prey, filtering rates, feeding rates and electivity indices is derived. Results from a comparison of selectivity coefficients for the copepod Diaptomus oregonensis grazing under a number of experimental conditions suggest that W_ij's for size-selective feeding are invariant, a conclusion also supported by the leaky-sieve model. Recommendations are made on how to use W_ij's in linear and nonlinear feeding constructs for zooplankton and other animals.     

Demographic variation within spatially structured reef fish populations: when are larger-bodied subpopulations more important?

Environmental heterogeneity frequently induces spatial variability in somatic growth, which can cause inter-population differences in reproductive output among organisms for which fecundity is dependent upon body size. Mean asymptotic body size, L_∞, varies among populations of several reef fish species. Deterministic models suggest L_∞ has little effect on population growth, so subpopulations with larger L_∞ may not have disproportionate effects in sustaining an open system. We used a stochastic simulation model to examine the potential role of a larger L_∞ subpopulation in aspects of population dynamics beyond population growth under a range of assumptions about the prevailing recruitment relationships. We compared dynamics of a demographically homogeneous system with a system that included one subpopulation with 20% larger L_∞. Despite the magnitude of the increase in L_∞, mean population size and average time at large population sizes differed little between the homogenous system and that with the larger L_∞ subpopulation. However, including the larger L_∞ subpopulation did result in less time spent at very small population sizes, which could reduce extinction risks. Effects of the larger L_∞ subpopulation were most pronounced when a deterministic recruitment cycle was imposed in combination with high stochastic variability in recruitment. This was due to regular series of poor recruitment years shifting the population structure toward older cohorts where differences in body size (and reproductive output) between the larger L_∞ subpopulation and the other subpopulations were greatest. Differences were also greater when recruitment variability was regionally correlated. When recruitment variability was locally independent, the probability of system-wide declines was reduced because declines of individual populations at one time were replenished by unaffected neighbors in subsequent years. Our study suggests that variation in L_∞ within a network of interconnected subpopulations may not be an important determinant of population behavior under certain conditions, but might be important in coping with periods of persistent, system-wide recruitment failure.