Using demography and movement behavior to predict range expansion of the southern sea otter

In addition to forecasting population growth, basic demographic data combined with movement data provide a means for predicting rates of range expansion. Quantitative models of range expansion have rarely been applied to large vertebrates, although such tools could be useful for restoration and management of many threatened but recovering populations. Using the southern sea otter (Enhydra lutris nereis) as a case study, we utilized integro-difference equations in combination with a stage-structured projection matrix that incorporated spatial variation in dispersal and demography to make forecasts of population recovery and range recolonization. In addition to these basic predictions, we emphasize how to make these modeling predictions useful in a management context through the inclusion of parameter uncertainty and sensitivity analysis. Our models resulted in hind-cast (1989-2003) predictions of net population growth and range expansion that closely matched observed patterns. We next made projections of future range expansion and population growth, incorporating uncertainty in all model parameters, and explored the sensitivity of model predictions to variation in spatially explicit survival and dispersal rates. The predicted rate of southward range expansion (median = 5.2 km/yr) was sensitive to both dispersal and survival rates; elasticity analysis indicated that changes in adult survival would have the greatest potential effect on the rate of range expansion, while perturbation analysis showed that variation in subadult dispersal contributed most to variance in model predictions. Variation in survival and dispersal of females at the south end of the range contributed most of the variance in predicted southward range expansion. Our approach provides guidance for the acquisition of further data and a means of forecasting the consequence of specific management actions. Similar methods could aid in the management of other recovering populations.     

Dispersal of fish populations in dams: modelling and simulation

A mathematical model is introduced for the analysis of changes in the habitat of certain species and consequent migration processes. The weak formulation for the resulting partial differential non-linear equation is presented, as well as a linearization method. Existence and uniqueness results are analyzed. In addition, a numerical method is suggested. Finally, results from numerical simulations with empirical parameters are shown and discussed from an ecological point of view.     

A simulation model for the management of vicuña populations

The population dynamics of the vicuña was modelled based on field data from the Central Galeras Sector, Perú. Rains were simulated in order to obtain net primary productivity and grass availability which act upon density-dependent fecundity and mortality. The model produces a population growth curve that tends to stabilize at densities around 100 vicuñas per km². Harvest and shearing processes were simulated as part of the vicuña population management model. Harvest was based upon a fixed threshold density D_e below which no harvest occurs; for densities above D_e harvesting is applied at a rate proportional to the difference between D_e and the current population density (fixed escapement or ‘bang-bang’ harvest rule). Management optimization was analyzed by determining the optimal escapement density D_e, which maximized either net profits or the number of animals harvested. Analyzing the results as cumulative totals over 20 years of simulation, an optimum harvest production was obtained for a D_e of about 40 vicuñas per km², and maximum profit was obtained for a D_e around 70 vicuñas per km². Analyzing the results of 20 years of simulation as annual averages with an original population density of 40 vicuñas per km², the harvest production and profits were maximized for a D_e of 40 and 60 vicuñas per km², respectively.The model was validated using data from the Reserva Nacional San Guillermo, Argentina, where vicuña and guanaco populations coexist (although their interaction was not modelled). The sensitivity analysis was performed with three different techniques: (a) stepwise multiple linear regression, (b) visual graphic analysis based on a polar coordinates system, and (c) direct evaluation of the effect on management decisions. The curve shape parameters of the fecundity and mortality functions proved to be the most important ones in determining the outcome of the model.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

A simulation analysis of continental shelf food webs

Energy flow through continental shelf food webs was examined using a simulation model. The model structure expands the two traditional marine food chains of phytoplankton-zooplankton-pelagic fish and benthos-demersal fish into a complex web which includes detritus, dissolved organic matter (DOM), bacteria, protozoa, and mucus net feeders. Simulation of energy flux for different shelf systems using the expanded web revealed that heterotrophic microorganisms and their predators account for a significant component of the energy flux in the continental shelf ecosystem. Contrary to previous models, where all phytoplankton were considered to be grazed by zooplankton, our simulation results indicate that only slightly more than 50% of the annual net primary production is grazed. A substantial quantity of the phytoplankton production directly becomes detritus. Bacteria mineralize detritus and DOM produced by phytoplankton and other components of the food web, converting these to biomass with high efficiency. Consequently, the model predicts that planktonic bacterial production is equivalent to zooplankton production. Exclusion of the bacteria requires the assumption that all DOM is either exported from the system or consumed by another component of the food web. Neither of these assumptions can be supported by present knowledge of the dynamics of DOM in the sea. Model simulations were also employed to test the hypothesis that production exceeds consumption on continental shelves, resulting in exports of 50% of the annual primary production. Simulations of shelves with high rates of primary production resulted in a particulate export of 27% and realistic estimates of secondary production. Results of other simulations suggest that shelves with lower primary production cannot export production and still maintain the macrobenthos and their predators. General properties about continental shelves can also be inferred from the model. From simulations of shelves of differing primary production, nanoplankton are predicted to account for a greater proportion of the primary production in nutrient limited systems. Benthic production appears to be related to both the quantity of primary production and the sinking rates of the phytoplankton. The model indicates that zooplankton fecal inputs to the shelf benthos are only a small portion of the total detrital flux, leading to the prediction that fecal pellets are of little significance in determining benthic production. Finally, the model generates production efficiencies that are highly variable depending on the type of system and kind of populations involved. We argue that the assumed ecological efficiency of 10% should be abandoned for continental shelves and other ecosystems.     

Temperature adaptation in sea anemones: Physiological and biochemical variability in geographically separate populations of Metridium senile

Latitudinally separate populations of the sea anemone Metridium senile (L.) are very similar genetically by electrophoretic criteria, yet respond differently to temperature. Anemones from southern and northern California (USA) have different oxygen consumption patterns in response to acclimatory and acute changes in temperature. Northern anemones show a pronounced increase in Q₁₀ at temperatures just above the normal environmental range, but southern anemones do not. The two populations also differed in the extent of metabolic compensation to temperature following several weeks of acclimation. This acclimation regime resulted in changes in the activities of several enzymes of intermediary metabolism, yet the extent and direction of these changes did not display a consistent trend with regard to acclimation temperature or population. The biochemical concomitants of acclimatory and acute temperature effects were studied further by measuring the concentrations of adenylates (ATP, ADP, and AMP) in anemones from the two populations exposed to different temperature regimes. During cold acclimation for several weeks, total adenylate concentrations (A_T) increased in both the southern and northern populations, possibly due to metabolic rate compensation, since A_T is positively correlated with tissue metabolic rate in many species. Moreover, the extremely low weight-specific oxygen consumption rates of M. senile are probably related to its very low A_T values. Acute temperature decreases had no effect on adenylate concentrations and adenylate energy charge (AEC); in contrast, acute temperature increases led to large changes in adenylate concentrations. The effects of starvation on adenylate concentrations are pronounced, and the effect is temperature-dependent. In starved individuals held at 20°C, AEC values fell to 50% of normal values after 8 d, while those held at 10°C maintained normal AEC values.     

Short-term stability of genetic structure in populations of the sea anemone Metridium senile

Populations of the sea anemone Metridium senile (L.) were sampled from several locations in eastern North America in two series, one collected from 1977–1979 and the other from 1981–1985. Fourteen populations were sampled twice at one- to six-year intervals. Samples were analyzed for temporal differences in genetic composition at both the single locus and multiple locus levels. Overall patterns of geographic variation in allele frequency did not change between series. Regressions describing clines did not differ significantly, and loci not showing clinal variation in the first series remained similar in the second. Analysis of populations sampled twice produced no systematic evidence of change in allele frequency with time for any of four polymorphic loci. Comparisons with computer simulations of repeated sampling of multiple locus genotypes from panmictic populations with free recombination also revealed little temporal change at this level. One population showed possible evidence of recruitment from a different gene pool. Other significant departures from expectations reflected more reproducibility of genotype distributions between samples than expected for sexual populations. This excess stability likely results jointly from clonal reproduction and little sexual recruitment. Despite these indications of genetic stability in adult populations, newly settled juveniles were genetically different from resistant adults in one population, demonstrating the potential for genetic change by immigration. Successful sexual recruitment seems to be rare, even though larvae regularly settle from the plankton. Although interpretation of these results is somewhat limited by lack of knowledge of longevities and generation times, to the extent that they reflect longer-term trends, they suggest that at least some of the observed patterns of geographic variation in allele frequency probably result from natural selection.     

Structure of monogenean populations,ectoparasites of the gilthead sea bream Sparus aurata

In the Western Mediterranean, the gilthead sea bream Sparus aurata Linnaeus, 1758 is parasitized by two gill monogeneans, Furnestinia echeneis (Wagener, 1857) Euzet and Audouin, 1959 (Monopisthocotylea: Diplectanidae) and Sparicotyle chrysophrii (van Beneden and Hesse, 1863) Mamaev, 1984 (Polyopisthocotylea: Microcotylidae). The spatial structure of these parasitic populations were studied on the French coast in the Gulf of Lions in 1984 and 1985. The relations between the two species forming this guild have been demographically analyses. Their distribution profiles are very similar and display some stability with time, although some increasing trends are perceptible in the spring. These two monogeneans are globally frequent, but always present in only moderate intensities in infrapopulations. Recruitment of F. echeneis is independent of that of S. chrysophrii. This mode of contamination limits massive and simultaneous infestations of a single host, and interspecific competition risks are consequently considerably reduced. Our results suggest these parasites can affect the size of a natural gilthead population to only a limited degree.     

Climate change threatens polar bear populations: a stochastic demographic analysis

The polar bear (Ursus maritimus) depends on sea ice for feeding, breeding, and movement. Significant reductions in Arctic sea ice are forecast to continue because of climate warming. We evaluated the impacts of climate change on polar bears in the southern Beaufort Sea by means of a demographic analysis, combining deterministic, stochastic, environment-dependent matrix population models with forecasts of future sea ice conditions from IPCC general circulation models (GCMs). The matrix population models classified individuals by age and breeding status; mothers and dependent cubs were treated as units. Parameter estimates were obtained from a capture-recapture study conducted from 2001 to 2006. Candidate statistical models allowed vital rates to vary with time and as functions of a sea ice covariate. Model averaging was used to produce the vital rate estimates, and a parametric bootstrap procedure was used to quantify model selection and parameter estimation uncertainty. Deterministic models projected population growth in years with more extensive ice coverage (2001-2003) and population decline in years with less ice coverage (2004-2005). LTRE (life table response experiment) analysis showed that the reduction in lambda in years with low sea ice was due primarily to reduced adult female survival, and secondarily to reduced breeding. A stochastic model with two environmental states, good and poor sea ice conditions, projected a declining stochastic growth rate, log lambdas, as the frequency of poor ice years increased. The observed frequency of poor ice years since 1979 would imply log lambdas approximately - 0.01, which agrees with available (albeit crude) observations of population size. The stochastic model was linked to a set of 10 GCMs compiled by the IPCC; the models were chosen for their ability to reproduce historical observations of sea ice and were forced with "business as usual" (A1B) greenhouse gas emissions. The resulting stochastic population projections showed drastic declines in the polar bear population by the end of the 21st century. These projections were instrumental in the decision to list the polar bear as a threatened species under the U.S. Endangered Species Act.     

Physiological energetics of cultivated mussel (Mytilus edulis) populations in two Scottish west coast sea lochs

Physiological responses of suspended cultured mussels, Mytilus edulis L., in two Scottish sea lochs (Lochs Etive and Leven) were investigated in their native and transplanted environments, after 15 d, 4.5 mo and 1 yr acclimatization, during the main growing season of May to September 1992. These measurements were integrated by means of the balanced energy equation, and scope for growth was calculated to assess the performance of each stock. Transplanted mussels showed clear signs of stress during the first 15 d after transfer, with low clearance rates and energy retention and high rates of respiration and nitrogen excretion. There were significant differences in some of the physiological responses (clearance, respiration and excretion rates), scope for growth and growth efficiency between the native populations, with the responses of mussels in Loch Etive being more favourable than those in Loch Leven. With newly transplanted mussels after 15 d acclimatization, almost all these measurements also differed significantly from native mussels in their host site and the original stocks but, with the exception of ammonia excretion rates, all the variables of cross-transplanted mussels after 4.5 mo acclimatization were the same as those of the native stock at the host site. Good agreement between observed long-term growth rates and estimated scope for growth suggests that, like growth rate, differences in physiological responses are mainly controlled by environmental factors; i.e., stock or origin had no significant influence on variations in physiological response (except ammonia excretion). The results further indicate that scope for growth estimated during the main growing season can be used to assess the actual growth rate and to compare sites for on-growing.     

Genetic divergence between colour morphs in populations of the common intertidal sea anemones Actinia equina and A. prasina (Anthozoa: Actiniaria) in the Isle of Man

A distinctive morph ofActinia equina (L.) is found at low frequency among populations on intertidal hard substrata on some shores on the Isle of Man. This morph is red with rows of green oval spots or elongate markings running longitudinally down the column. Starch gel electrophoresis of 21 allozyme loci was used to compare samples of this morph from two localities with other sympatricActinia spp. Collections were made in early summer 1986. For one sampling site the comparison was with redA. equina and greenA. prasina Gosse, whilst at the other site with redA. equina only. At both sites significant genetic differences in allele frequencies at several loci were found between the new morph and sympatricA. equina andA. prasina. The results also confirm the reproductive isolation ofA. prasina. As with much previous work on sea anemones, levels of mean heterozygosity per locus (H) were found to be high (H=0.157 to 0.342). A surprising feature of the results, although bearing out earlier unpublished data, is that high levels of genetic differentiation are found between populations of what appear to be the same morphs collected from shores only a few km apart. Among theActinia spp. studied, differentiation between populations of the same morph on different shores was of the same order as between sympatric, reproductively isolated populations.     

Utility maximization and catasphore aversion: A simulation test

Economists have proved that the exclusion of the effects of irreversibility from a decision-making framework is a specification error which leads to sub-optimal solutions. Consequently, in any cost-benefit framework, the effects of an irreversibility are included in the form of an option value. Unfortunately, in the real world, there exist certain potential irreversibilities that jeopardize the very existence of the decision-making entity, were they to occur. We demonstrate that if the effects of such irreversibilities are modeled in terms of a finite option value (in cost-benefit analysis) then the optimal solutions derived therefrom may be unable to avert a potential catastrophe. In theory, an infinite option value should lead to catastrophe-averting decisions in such situations. However, in practice, such mathematical formulations tend to be intractable. In many situations, irreversible catastrophes proceed from a positive reinforcement of fluctuations in system behavior, primarily due to sympathetic responses in sub-systems. Engineers have designed relatively low-cost methods to dampen similar oscillations in physical systems. We demonstrate how such engineering type analysis can be used successfully to identify catastrophe-averting policies for a model ecosystem that faces irreversible famine that must be avoided as a precondition for survival.     

Changes in size and trends of North American sea duck populations associated with North Pacific oceanic regime shifts

Broad-scale multi-species declines in populations of North American sea ducks for unknown reasons is cause for management concern. Oceanic regime shifts have been associated with rapid changes in ecosystem structure of the North Pacific and Bering Sea. However, relatively little is known about potential effects of these changes in oceanic conditions on marine bird populations at broad scales. I examined changes in North American breeding populations of sea ducks from 1957 to 2011 in relation to potential oceanic regime shifts in the North Pacific in 1977, 1989, and 1998. There was strong support for population-level effects of regime shifts in 1977 and 1989, but little support for an effect of the 1998 shift. The continental-level effects of these regime shifts differed across species groups and time. Based on patterns of sea duck population dynamics associated with regime shifts, it is unclear if the mechanism of change relates to survival or reproduction. Results of this analysis support the hypothesis that population size and trends of North American sea ducks are strongly influenced by oceanic conditions. The perceived population declines appear to have halted >20 years ago, and populations have been relatively stable or increasing since that time. Given these results, we should reasonably expect dramatic changes in sea duck population status and trends with future oceanic regime shifts.     

Increases in surgeonfish populations after mass mortality of the sea urchinDiadema antillarum in Panamá indicate food limitation

In 1983/1984,Diadema antillarum suffered mass mortalities throughout its West Atlantic range. Its populations were reduced by 95% and subsequently have failed to recover. These die-offs led to sustained increases in the abundance of soft algae, including types eaten by herbivorous reef fishes. I monitored adult populations of three herbivorous surgeonfishes (Acanthurus coeruleus, A. chirurugus andA. bahianus) between 1978 and 1990, and the recruitment of their pelagic juveniles between 1979 and 1989, on six patch reefs in Panamá. Adult populations ofA. coeruleus andA. chirurgus, which largely restrict their feeding to reef substrata, increased by averages of 250 and 160%, respectively, after the die-off ofD. antillarum in 1983. No increases occurred in the adult populations ofA. bahianus, which often feeds in off-reef habitats unaffected byD. antillarum. Average annual levels of juvenile recruitment of all three surgeonfishes did not differ before and after the die-off. These results support the hypothesis that adult populations of two herbivorous fishes that are strongly reliant on reef algae for food previously were limited by competition withD. antillarum.     

On interacting bee/mite populations: a stochastic model with analysis using cumulant truncation

Parasitism by the Varroa mite has had recent drastic impact on both managed and feral bee colonies. This paper proposes a stochastic population dynamics model for interacting African bee colony and Varroa mite populations. Cumulant truncation procedures are used to obtain approximate transient cumulant functions, unconstrained by the usual assumption of bivariate Normality, for an assumed large-scale model. The apparent size of the variance and skewness functions suggest the importance of the proposed truncation procedure which retains some higher-order cumulants, but determining the accuracy of the approximations is problematical. A smaller-scale bee/Varroa mite model is hence proposed and investigated. The accuracy for the means is exceptional, for the second-order cumulants is moderate, and for some third-order cumulants is poor. Notwithstanding the poor accuracy of a skewness approximation, the saddlepoint approximations for the marginal transient population size distributions are excellent. The cumulant truncation methodology is very general, and research is continuing in its application to this new class of host-parasite models.     

Potential effects of a resuspension event on the vertical distribution of copepod eggs in the sea bed: a laboratory simulation

Adult female Acartia tonsa and Labidocera aestiva were obtained from inshore marine waters near Turkey Point, Florida (29°54.5N;84°31W) during April and May 1991. For each species, eggs spawned overnight in the laboratory were collected, mixed with sediment, and added to a plastic tube (26 to 29 cm longx5 cm diam) to yield a 2 cm-thick layer. This tube was connected to a 245 cm-long tube, seawater was added and the contents were thoroughly mixed by turning the assembly end-over-end several times. The vertical distribution of eggs in the sediments following resuspension varied according to the grain size of the sediments. For sediments of 125 m particle-diameter, 68 to 72%, and 70 to 73% of the eggs of A. tonsa and L. aestiva, respectively, occurred in the upper 4 mm.For sediments 63 to <125 m particle-diameter, 34 to 36% of the eggs of A. tonsa and L. aestiva, respectively, occurred in the upper 4 mm. For sediments of <63 m particle-diameter, egg distribution was uniform for both species throughout the 2 cm layer. The results indicate that the vertical distribution of eggs in the sea bed following a resuspension event should be highly dependent upon the grain-size composition of the sediments.     

Population viability analysis for several populations using multivariate state-space models

The International Union for the Conservation of Nature and Natural Resources (IUCN), the world's largest and most important global conservation network, has listed approximately 16,000 species worldwide as threatened. The most important tool for recognizing and listing species as threatened is population viability analysis (PVA), which estimates the probability of extinction of a population or species over a specified time horizon. The most common PVA approach is to apply it to single time series of population abundance. This approach to population viability analysis ignores covariability of local populations. Covariability can be important because high synchrony of local populations reduces the effective number of local populations and leads to greater extinction risk. Needed is a way of extending PVA to model correlation structure among multiple local populations. Multivariate state-space modeling is applied to this problem and alternative estimation methods are compared. The multivariate state-space technique is applied to endangered populations of pacific salmon, USA. Simulations demonstrated that the correlation structure can strongly influence population viability and is best estimated using restricted maximum likelihood instead of maximum likelihood.     

Genetic structure of Atlantic and Gulf of Mexico populations of sea bass,menhaden, and sturgeon: Influence of zoogeographic factors and life-history patterns

To assess the influence of zoogeographic factors and life-history parameters (effective population size, generation length, and dispersal) on the evolutionary genetic structure of marine fishes in the southeastern USA, phylogeographic patterns of mitochondrial DNA (mtDNA) were compared between disjunct Atlantic and Gulf of Mexico populations in three coastal marine fishes whose juveniles require an estuarine or freshwater habitat for development. Black sea bass (Centropristis striata), menhaden (Brevoortia tyrannus andB. patronus) and sturgeon (Acipenser oxyrhynchus) samples were collected between 1986 and 1988. All species showed significant haplotype frequency differences between the Atlantic and Gulf, but the magnitude and distribution of mtDNA variation differed greatly among these taxa: sea bass showed little within-region mtDNA polymorphism and a clear phylogenetic distinction between the Atlantic and Gulf; menhaden showed extensive within-region polymorphism and a paraphyletic relationship between Atlantic and Gulf populations; and sturgeon exhibited very low mtDNA diversity both within regions and overall. Evolutionary effective sizes of the female populations (N _f (e)) estimated from the mtDNA data ranged fromN _f (e) = 50 (Gulf of Mexico sturgeon) toN _f (e) = 800 000 (Atlantic menhaden), and showed a strong rank-order agreement with the current-day census sizes of these species. The relationship betweenN _f (e) and the estimated times of divergence (t) among mtDNA lineages (from conventional clock calibrations) predicts the observed phylogenetic distinction between Atlantic and Gulf sea bass, as well as the paraphyletic pattern in menhaden, provided the populations have been separated by the same long-standing zoogeographic barriers thought to have influenced other coastal taxa in the southeastern USA. However, vicariant scenarios alone cannot explain other phylogenetic aspects of the menhaden (and sturgeon) mtDNA data and, for these species, recent gene flow between the Atlantic and Gulf coasts is strongly implicated. These data are relevant to management and conservation issues for these species.Please address all requests for reprints to Dr. J. C. Avise     

A simulation model of estuarine subsystem coupling and carbon exchange with the sea. II. North Inlet model structure,output and validation

A first-stage verified model of carbon/energy flux through the North Inlet (South Carolina) marsh—estuarine ecosystem is presented. The time series output for model compartments and overall septem carbon flow are compared with observed data collected over the past five to ten years. Results indicate that the model is stable and can broadly reproduce some of the major trends of a salt marsh—estuarine system. Further avenues of research are suggested.