Generation time and the maximum growth rate for populations with age-specific fecundities and unknown juvenile survival

In age-classified population models where all parameters are known, the generation time and growth rate are calculated in a straightforward manner. For many populations, some parameters, such as juvenile survival, are difficult to estimate accurately. In a simplified population model where fecundity and survival are constant from the onset of breeding, it is known that generation time may be calculated given only adult survival, age at first reproduction, and the population growth rate. However, the assumption of constant fecundity from the onset of breeding does not hold for many populations. An extended population model allows calculation of generation time with the additional knowledge of the ratio of age-specific fecundities compared to a maximum fecundity rate. When these relative fecundities are unknown, an ad hoc adjustment to the simplified model performs well.When the study population is in an ideal environment, the optimal generation time and maximum growth rate are linked, and both may be approximated knowing only adult survival, age at first reproduction, and the relative fecundities. The maximum growth rate has important conservation implications, and calculating it correctly is therefore important. Improper use of the simplified population model to calculate the maximum growth rate, combined with a simple decision rule, leads to an average overharvest of 36%, and >60% for three of six bird species studied, compared to the full population model. By comparison, using the approximation from the extended or adjusted models results in average overharvests of only 8% (extended model) and 5% (adjusted model), and <50% for all six species (either model).     

The onset of natural convection in lakes and reservoirs due to night time cooling

A theoretical model, based on linear stability analysis, is proposed to predict the onset of natural convection in lakes and reservoirs due to night time cooling. To such purpose, the system was modelled as a initially quiescent deep Boussinesq fluid reservoir, whose upper boundary temperature changes sinusoidally. From scaling analysis, it is found that critical onset times for convection are proportional to R ^−2/7, where R is a Rayleigh number including fluid properties and forcing frequency. The proportionality constant was found, from the solution of an eigenvalue problem, as a function of the Prandtl number. The onset time for convection was easily observed from experiments and quantitatively detected as a rapid increase of the RMS of the computed velocity field obtained using PIV. In this controlled conditions, differences close to 10% between predicted and observed times for the start of the convective flow was found. It is apparent from the present set of results that predictions are reasonable.     

Microsatellite length polymorphisms associated with dispersal-related agonistic onset in male wild house mice (Mus musculus domesticus)

Dispersal propensity, reflecting one of the most decisive mammalian life history traits, has been suggested to vary heritably and to locally adapt to prevailing dispersal conditions in wild house mouse populations. Because individual dispersal propensity highly significantly covaries with the developmental timing of the onset of agonistic interactions between littermate brothers, we used agonistic onset as an endophenotype to explore the potential genetic basis of dispersal-related behavioral variation in male house mice. We found significant covariation of microsatellite marker compositions with the probability of fraternal pairs to exhibit agonistic relationships before the age of 2 months. In particular, the presence of two alleles associated with a serotonin transporter protein gene (Slc6a4) and a testosterone dehydrogenase gene (Cyp3a11), respectively, strongly covaried with the probability of early agonistic onset. These results are congruent with recent findings of microsatellite length polymorphisms marking regulatory variation of gene expression that is relevant for social behavior, including dispersal propensity development, in other mammals. Genetic variability for ontogenetic timing of agonistic onset would be in agreement with genotypic differentiation of the dispersive behavioral syndrome in natural populations that could lead to local adaptation.     

Brood temperature, task division and colony survival in honeybees: A model

One of the mechanisms by which honeybees regulate division of labour among their colony members is age polyethism. Here the younger bees perform in-hive tasks such as heating and the older ones carry out tasks outside the hive such as foraging. Recently it has been shown that the higher developmental temperatures of the brood, which occur in the centre of the brood nest, reduce the age at which individuals start to forage once they are adult. It is unknown whether this effect has an impact on the survival of the colony. The aim of this paper is to study the consequences of the temperature gradient on the colony survival in a model on the basis of empirical data.We created a deterministic simulation of a honeybee colony (Apis mellifera) which we tuned to our empirical data. In the model in-hive bees regulate the temperature of the brood nest by their heating activities. These temperatures determine the age of first foraging in the newly emerging bees and thus the number of in-hive bees present in the colony. The results of the model show that variation in the onset of foraging due to the different developmental temperatures has little impact on the population dynamics and on the absolute number of bees heating the nest unless we increase this effect by several times to unrealistic values, where individuals start foraging up to 10 days earlier or later. Rather than on variation in the onset of foraging due to the temperature gradient it appears that the survival of the colony depends on a minimal number of bees available for heating at the beginning of the simulation.     

Frailty in state-space models: application to actuarial senescence in the Dipper

Senescence, a decrease in life history traits with age, is a within-individual process. The lack of suitable methods to deal with individual heterogeneity has long impeded progress in exploring senescence in wild populations. Analyses of survival senescence are additionally complicated by the often neglected issue of imperfect detectability. To deal with both these issues, we developed state-space models to analyze capture-mark-recapture data while accounting for individual heterogeneity by incorporating random effects. We illustrated our approach by applying it to 29 years of data on breeding females in a Dipper (Cinclus cinclus) population. We highlighted patterns of age-related variation in annual survival by statistical comparisons of piecewise linear, quadratic, Gompertz, and Weibull survival models. The Gompertz model was ranked first in our set. It provided strong evidence for actuarial senescence with an onset of senescence estimated at about 2.3 years. The probability for this model to involve a frailty was 0.15, and the probability to involve an individual latent effect in detection was about 0.4. The estimated mean age at first reproduction was 1.2 years. The general case model described here in detail should encourage the reanalysis of actuarial senescence in cases where imperfect detection or individual heterogeneity is suspected.     

Hatching asynchrony is constrained by parental nest attendance during laying

Hatching asynchrony is widespread amongst animals, but no consensus has yet emerged as to why asynchronous hatching has evolved. It is generally thought to have adaptive benefits during the raising of dependent young. However, here, we considered an alternative view of hatching asynchrony in birds as a consequence of factors acting at the onset of incubation. We recorded parental nest attendance behaviour during laying using continuous records of nest temperature in herring gulls, Larus argentatus. We tested whether nest attendance during laying was related to individual factors (clutch size and diet) and whether it had consequences on fitness outcomes (hatching spread, incubation period, hatching success and chick survival). Low nest attendance was associated with small clutch size, and independent of clutch size, pairs on a more marine diet had lower nest attendance than pairs on a lower trophic level terrestrial diet, possibly due to higher foraging effort for marine food. Broods hatched more asynchronous where pairs had a lower nest attendance during laying or took longer to complete a clutch and where the last egg took longer to hatch. Low nest attendance was also related to shorter incubation periods, possibly representing a strategy of birds in poor condition to reduce the demand of incubation by reducing the length of incubation. We found that low nest attendance during laying and increasing hatching asynchrony had detrimental effects on hatching success for small eggs laid early in the laying sequence. Increasing hatching asynchrony also had a detrimental effect on the survival of the youngest sibling. In our study population, hatching asynchrony was influenced by a more complex set of factors than simply onset of incubation and appears to be constrained by circumstances at the onset of incubation rather than to be an adaptive strategy. Thus, factors acting both during offspring rearing and at the onset of incubation need to be considered for a better understanding of hatching asynchrony.     

Modelling the effect of fibropapilloma disease on the somatic growth dynamics of Hawaiian green sea turtles

The effect of the tumour-forming disease, fibropapillomatosis, on the somatic growth dynamics of green turtles resident in the Pala’au foraging grounds (Moloka’i, Hawai’i) was evaluated using a Bayesian generalised additive mixed modelling approach. This regression model enabled us to account for fixed effects (fibropapilloma tumour severity), nonlinear covariate functional form (carapace size, sampling year) as well as random effects due to individual heterogeneity and correlation between repeated growth measurements on some turtles. Somatic growth rates were found to be nonlinear functions of carapace size and sampling year but were not a function of low-to-moderate tumour severity. On the other hand, growth rates were significantly lower for turtles with advanced fibropapillomatosis, which suggests a limited or threshold-specific disease effect. However, tumour severity was an increasing function of carapace size—larger turtles tended to have higher tumour severity scores, presumably due to longer exposure of larger (older) turtles to the factors that cause the disease. Hence turtles with advanced fibropapillomatosis tended to be the larger turtles, which confounds size and tumour severity in this study. But somatic growth rates for the Pala’au population have also declined since the mid-1980s (sampling year effect) while disease prevalence and severity increased from the mid-1980s before levelling off by the mid-1990s. It is unlikely that this decline was related to the increasing tumour severity because growth rates have also declined over the last 10–20 years for other green turtle populations resident in Hawaiian waters that have low or no disease prevalence. The declining somatic growth rate trends evident in the Hawaiian stock are more likely a density-dependent effect caused by a dramatic increase in abundance by this once-seriously-depleted stock since the mid-1980s. So despite increasing fibropapillomatosis risk over the last 20 years, only a limited effect on somatic growth dynamics was apparent and the Hawaiian green turtle stock continues to increase in abundance.     

A model for the larval migration of the Japanese eel: roles of the trade winds and salinity front

A model that helps explain the mysterious long-distance migration of the Japanese eel (Anguilla japonica) is presented, based on oceanographic observations, satellite buoy drift experiments, and samplings of eel larvae taken in 1991. The trajectory of a 150 m depth buoy relased in the spawning area strongly suggests that A. japonica larvae spawned just south of the salinity front are transported westward by the North Equatorial Current (NEC). The larvae are then thought to be entrained into the Mindanao Current flowing southward along the Philippine Islands where A. japonica juveniles are scarcely distributed. These controversial results lead to the assumption that eel larvae are transferred from the NEC to the northward flowing Kuroshio, which distributes the eel larvae to the growth habitats of eastern Asia. In this eel larvae transfer model, a northward Ekman transport caused by trade winds plays an important role in explaining the wind-induced northward shift of the larvae together with the onset of diel vertical migration. Assuming that leptocephali greater than 20 mm initiate the vertical migration, a westward wind velocity greater than 5 to 10 m s^-1 should be high enough to diminish the southward current velocity. When the physical and geophysical conditions — such as the salinity front for spawning activity, the water tunnel for westward larval transport, the Ekman transport by the trade wind for transfer of the larvae from the NEC to the Kuroshio, and the strong velocity of the Kuroshio for rapid transport to growth habitats — are well matched with the timing of the onset of vertical migration, large-scale eel migration could result.     

The Early Development of Forest Fragmentation Effects on Birds

The early development of forest fragmentation effects on forest organisms is poorly understood partly because most fragmentation studies have been done in agricultural or suburban landscapes, long after the onset of fragmentation. We develop a temporal model of forest fragmentation effects on densities of forest-breeding birds and provide data from an active industrial forest landscape to test the model. The model and our empirical data indicate that densities of several forest-dwelling bird species can increase within a forest stand soon after the onset of fragmentation as a result of displaced individuals packing into remaining habitat. Along with higher densities in the newly formed fragments, pairing success in one species, the Ovenbird ( Seiurus aurocapillus ), was lower in fragments than nonfragments, possibly due to behavioral dysfunction resulting from high densities. Thus, density was inversely related to productivity. The duration and extent of increased densities following onset of fragmentation depends on many factors, including the sensitivity of a species to edge and area effects, the duration and rate of habitat loss and fragmentation, and the proximity of a forest stand to the disturbance. Incipient forest fragmentation may affect populations differently from later stages of fragmentation when the geometry of the landscape has reached a more stable configuration. Our model and data indicate, for reasons unrelated to traditional edge effects, that large tracts of forest can be important because they are relatively free from the variety of plant and animal population dynamics that might take place near new edges, including the encroachment of individuals displaced by habitat loss.     

Identification of calanoid copepod prey species via molecular detection of carbon fixation genes

Zooplankton and their phytoplankton prey form the basis of the marine food web, yet historically it has been difficult to discern species-specific trophic interactions. Molecular techniques provide opportunities to obtain taxonomic data where the traditional methodologies for gut content analysis lack resolution. The large subunit gene of RubisC/O, rbcL, was utilized as a molecular marker for the identification of prey species in calanoid copepods. Clone libraries were generated from DNA extracted from seawater and whole copepods during a transect cruise on the northern Gulf of Mexico shelf. Sequence data analysis provided evidence of diatoms, nanoplankton-sized chlorophytes, and cyanobacteria in DNA extracted from whole copepods. These data demonstrate that rbcL can be a useful marker for the identification of copepod phytoplankton prey. Combining the described approach with quantitative techniques such as quantitative PCR will provide opportunities for the assessment of species-specific predator–prey interactions.     

Transition zone in otoliths of orange roughy (Hoplostethus atlanticus) and its relationship to the onset of maturity

A transition zone in the otoliths of orange roughy (Hoplostethus atlanticus) is described where annulus width decreases markedly. For fish sampled at spawning time, the relationship between gonad stage and the presence or absence of a transition zone is consistent with the hypothesis that this zone marks the onset of maturity, if allowance is made for some error in gonad staging and identification of the zone. Another data set from a non-spawning area after the spawning season is not consistent with the hypothesis. However, these latter data are considered less reliable because of the small size of the fish in the sample and difficulties in determining, outside the spawning season, whether a fish has been reproductively active. This hypothesis was used to provide estimates of the median age (30 yr) and length (31 cm) at the onset of maturity for the Chatham Rise, New Zealand population. These estimates are unaffected by the biases associated with the usual ogive method of estimation. Received: 27 June 1997 / Accepted: 1 July 1997     

Humic substances. Part III. sorptive interactions with environmental chemicals†

The formation of covalent binding to DNA of a carcinogen is now widely accepted to represent a classical mechanism of tumour induction in mammals. This mechanism does not operate with metals since no covalent binding of these agents to DNA does occur. Nevertheless, somatic mutations as typical consequences of DNA‐damage have been reported to be induced by metals in various model systems. Beside DNA‐alkylation such damages can be caused by changes in the conformation of DNA or in the fidelity of DNA‐repair. The activity of the repair enzymes DNA‐polymerases is indeed impaired by many metal ions at least in vitro. It is not yet established whether these mechanisms are also important in the intact mammalian organism. Much evidence has accumulated during the last years that a disturbance of the balance of cations and especially metal ions represents another possible mechanism of tumour induction. The tumours found with high doses of chelating agents such as nitrilo‐triacetic acid (NTA) have to be discussed in this context. Since most—if not—all of the speculative mechanisms of metal carcinogenesis resemble classical pharmacological reactions the existence of a threshold level is likely. So metal carcinogenesis will not be a problem of the environmental contamination at trace levels but a problem of occupational medicine.     

Patch dynamics based on Prisoner's Dilemma game: superiority of golden rule

There has been much literature on ecological model of Prisoner's Dilemma (PD) game. This game illustrates that cooperation can evolve in situations where individuals tend to look after themselves. In order to explain some behaviors of altruism in animal societies, the strategy All Cooperate (AC), often called the Golden Rule, is more appropriate than other strategies. However, very little is known about the superiority of AC. In the present article, we study patch dynamics based on non-iterated PD game, applying two different methods: island and lattice models. Each patch is assumed to be either vacant or composed of a population of AC or All Defect (AD), where AD means a selfish strategy. Both models exhibit a phase transition between a phase where both AC and AD survive, and a phase where AD is extinct. The latter phase means that AC beats AD completely. In the case of lattice model, the extinction of AD easily occurs and the abundance of AC takes a larger value, compared with the island model. Our models can be also extended to general iterated PD game; we describe the reason why AC can outperform any other strategy.     

Accounting for large-scale factors in the study of understory vegetation using a conditional logistic model

Local-scale and large-scale factors can affect the presence of a species of understory vegetation in the forest. Local-scale factors may be the influence of surrounding trees, while climate and latitude are typically considered large-scale factors. A model for the presence of a species needs to take into account both scales. A conditional logistic model is proposed for those studies where only the local-scale factors are of interest and that avoids estimating the large-scale parameters. Conditioning is carried out by the number of quadrats in the plot where the vegetation is found. As the latter is a sufficient statistic for the large-scale factors, a model free from these parameters is obtained. Data gathered in the permanent sample plots of the 1985–1986 National Forest Inventory of Finland is used for illustration, where the local-scale factor of interest is the influence of the trees, quantified by an index based on the size and location of the trees. The model fitted to Vaccinium vitis-idaea showed a significant and positive influence of Scots pine on the presence of this species, while for Calamagrostis arundinacea, a decrease in the odds ratio was observed due to the influence of Norway spruce.     

Evaluating predictive errors of a complex environmental model using a general linear model and least square means

A general linear model (GLM) was used to evaluate the deviation of predicted values from expected values for a complex environmental model. For this demonstration, we used the default level interface of the regional mercury cycling model (R-MCM) to simulate epilimnetic total mercury concentrations in Vermont and New Hampshire lakes based on data gathered through the EPAs Regional Environmental Monitoring and Assessment Program (REMAP). The response variable for the GLM was defined as R-MCMs predictive error: the difference between observed mercury concentrations and modeled mercury concentrations in each lake. Least square means of the response variable are used as an estimate of the magnitude and significance of bias, i.e., a statistically discernable trend in predictive errors for a given lake type, e.g., acidic, stratified, or oligotrophic. Using our approach, we determined lake types where significant over-prediction and under-prediction of epilimnetic total mercury concentration was occurring, i.e., regions in parameter space where the model demonstrated significant bias was distinguished from regions where no significant bias existed. This technique is most effective for finding regions of parameter space where bias is significant. Drawing conclusions concerning regions that show no significant bias can be misleading. The significant interaction terms in the GLM demonstrated that addressing this problem using univariate statistical techniques would lead to a loss of important information.     

Population differences in predation on Batesian mimics in allopatry with their model: selection against mimics is strongest when they are common

Batesian mimicry evolves when a palatable species, the “mimic,” resembles a dangerous species, the “model,” because both receive protection from predation. Yet, this protection should break down where the model is absent, because predators in such areas would not be under selection to avoid the model. Here, we test this prediction in a coral snake mimicry complex. We exposed plasticine replicas of milk snakes that closely mimic coral snakes to natural predators to determine if good mimetic milk snakes are preferentially attacked in allopatry with their model. Moreover, we evaluated whether attack rates on these replicas varied among three different allopatric regions that differed in the type of mimic found locally (i.e., good mimic, poor mimic, or no mimic). When all three regions were considered together, mimics were not preferentially attacked. When regions were analyzed separately, however, attacks on mimics were significantly greater than randomness only where good mimics were found. These variable levels of predation on good mimics might reflect frequency-dependent (i.e., apostatic) predation. In allopatric regions where good mimics are present, predators might have learned or evolved preferences for conspicuous, palatable prey that they encounter frequently. By contrast, in allopatric regions where good mimics are absent, predators might not have learned or evolved preferences for novel phenotypes. Thus, when predation is frequency-dependent, as long as good mimics are rare, they might not experience elevated levels of predation in allopatry with their model as predicted by the Batesian mimicry hypothesis.     

Landscape history, fragmentation, and patch occupancy: models for a forest bird with limited dispersal.

We developed and tested patch occupancy models for an endemic understory bird with limited dispersal ability, the Chucao Tapaculo (Scelorchilus rubecula), in two South American temperate rain forest landscapes that differed in levels and duration of forest loss. We assessed cover changes since 1961 in each landscape and surveyed patches for Chucao Tapaculo occupancy. We then developed incidence-based predictive models independently for each landscape and tested each model reciprocally in the alternative study area. We thereby assessed the domain of model applicability and identified those predictor variables with general effects and those that varied between the two landscapes. The two models were consistent regarding variable selection, and predictive accuracy of each model was high in the landscape where training data were collected. However, the models differed substantially in the magnitudes of effects related to patch size, with larger unoccupied patches observed in the landscape with the more advanced stage of fragmentation. Due to this discrepancy, each model performed poorly when applied to the alternative landscape, potentially reflecting the contrasting stages of habitat loss. Although it was impossible to dissociate effects of level and duration of forest loss, we viewed the landscapes as representing two extremes along a continuum of fragmentation, providing insights into potential trajectories for portions of the biome where deforestation is occurring. Further, our data suggest that static equilibrium models developed from distribution patterns in recently fragmented landscapes may overestimate persistence when used as a forecasting tool, or when extrapolated to alternative landscapes where fragmentation is more advanced. Thus, we suggest that landscapes used as standards for model building should be selected with caution. We recommend that distribution patterns be obtained from landscapes where fragmentation is advanced, preferably with histories of fragmentation long enough that time-delayed extinctions already would have occurred.     

Mating behaviour of the tiger blue butterfly (Tarucus theophrastus): competitive mate-searching when not all females are captured

Summary Females of the tiger blue butterfly (Tarucus theophrastus) fly upwind in search of the larval hostplant. Males perch or patrol the downwind edges, searching for incoming females or those already on the bush. A model of competitive mate-searching is developed for the case where not all receptive females are contacted by searching males: the model differs from the earlier ones of Parker (1970, 1974), particularly when few males are involved. Observed distributions of males upon bushes of different sizes agree better with the predictions of the game theory model than with a random distribution of males. The described model may be widely applicable to populations where females remate, but frequently evade detection by searching males.     

Fish guts as chemical reactors: a model of the alimentary canals of marine herbivorous fishes

The alimentary canals of marine herbivorous fishes are variously composed of a few basic structures strung together in series. We model the structures where digestive processes occur as chemical reactors following the model of Penry and Jumars: the stomach as a batch or continuous-flow stirred-tank (CSTR), the intestine as a plug-flow reactor (PFR), and the hindgut caecum as a CSTR. Other structures, where food is mechanically processed-gill rakers, pharyngeal mills, and muscular stomachs-are classified as gates. The optimality criterion for the model is the digestion of the most nutrient in the least amount of time. With the model we are able to predict gut configuration as a function of nutrient concentration and hypothesize that the guts of herbivorous fishes always have a PFR component and may or may not have a CSTR component. The Penry-Jumars model appears to provide a consistent theoretical framework for four main types of digestive mechanisms in marine herbivorous fishes and offers specific testable hypotheses on the feeding ecology and digestive physiology of four representative species of fish as well as other, still unstudied, herbivorous fishes.Contribution No. 68 from the Ocean Studies Institute, Long Beach, California 90840, USA     

The optimal use of non-renewable resources: The theory of extraction

The traditional theory of exhaustion is revised to allow for long run capital mobility. This model is extended to include the impact of cumulative environmental damages on the optimal path of resource use. Models and optimal markets are then analyzed for cases where minerals are available over a continuum of quality and where recycling ameliorates inevitable exhaustion of non-renewable resources. Finally, the impact of technological change on long run trends in mineral prices is examined.