Critical factors for the recovery of marine mammals

{en} Over the past decades, much research has focused on understanding the critical factors for marine extinctions with the aim of preventing further species losses in the oceans. Although conservation and management strategies are enabling several species and populations to recover, others remain at low abundance levels or continue to decline. To understand these discrepancies, we used a published database on abundance trends of 137 populations of marine mammals worldwide and compiled data on 28 potentially critical factors for recovery. We then applied random forests and additive mixed models to determine which intrinsic and extrinsic factors are critical for the recovery of marine mammals. A mix of life‐history characteristics, ecological traits, phylogenetic relatedness, population size, geographic range, human impacts, and management efforts explained why populations recovered or not. Consistently, species with lower age at maturity and intermediate habitat area were more likely to recover, which is consistent with life‐history and ecological theory. Body size, trophic level, social interactions, dominant habitat, ocean basin, and habitat disturbance also explained some differences in recovery patterns. Overall, a variety of intrinsic and extrinsic factors were important for species’ recovery, pointing to cumulative effects. Our results provide insight for improving conservation and management strategies to enhance recoveries in the future.     

Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

A mathematical model of PCB bioaccumulation in plankton

In batch experiments exposing individual plankton constituents to Aroclor 1254 PCB, the rate at which the organism approaches partitioning equilibrium appears to be partly size-dependent while the extent of PCB accumulation is species-specific. The sorptive desorptive kinetics of PCB in these experiments can be described mathematically by a first-order expression. Employing this expression in a model plankton food web permits examination of the role of feeding and sorptive processes in determining PCB body burden under various environmental conditions. When ingestion rates exceed desorption and excretion rates, a consuming organism accumulates PCB above levels predicted by equilibrium partitioning relationships. Feeding-induced oscillations in PCB body burden could thus obscure the reduction of soluble PCB concentration which determine a “baseline” PCB body burden. Unless referenced to a specific set of biological and environmental conditions, the importance of direct partitioning from water vs. food uptake appears to be a moot topic.     

A chance-constrained goal programming model to evaluate response resources for marine pollution disasters

This paper develops a model to aid Coast Guard managers in formulating appropriate policies with respect to planning for various types of equipment required to contain major pollution incidents. The model is elaborated in terms of three primary stages of response: offloading, containment, and removal. The zero order rule of chance constrained programming is used to obtain a deterministic equivalent of the original chance constrained model. This is then replaced by a goal programming formulation to allow for plans that come “as close as possible” to desired quality and risk levels for each pertinent region and type of incident. Numerical examples illustrate potential uses of the model with special emphasis on its value for budgetary (equipment) planning by central management that extends to evaluation of risk and performance quality levels, as well as the usual dual evaluator approaches for evaluating initially prescribed levels for equipment and their efficiency coefficients.     

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.     

A Bayesian mixture model for missing data in marine mammal growth analysis

Much of what is known about bottle nose dolphin (Tursiops truncatus) anatomy and physiology is based on necropsies from stranding events. Measurements of total body length, total body mass, and age are used to estimate growth. It is more feasible to retrieve and transport smaller animals for total body mass measurement than larger animals, introducing a systematic bias in sampling. Adverse weather events, volunteer availability, and other unforeseen circumstances also contribute to incomplete measurement. We have developed a Bayesian mixture model to describe growth in detected stranded animals using data from both those that are fully measured and those not fully measured. Our approach uses a shared random effect to link the missingness mechanism (i.e. full/partial measurement) to distinct growth curves in the fully and partially measured populations, thereby enabling drawing of strength for estimation. We use simulation to compare our model to complete case analysis and two common multiple imputation methods according to model mean square error. Results indicate that our mixture model provides better fit both when the two populations are present and when they are not. The feasibility and utility of our new method is demonstrated by application to South Carolina strandings data.     

Ecological energetics of the seaweed zone in a marine bay on the Atlantic coast of Canada. I. Zonation and biomass of seaweeds

An intertidal and underwater survey of the zonation of seaweed in St. Margaret's Bay, NovaScotia, Canada showed 8 major zones as one proceeded away from shore: (1) Fucus and Ascophyllum; (2) Chorda with filamentous browns; (3) Chondrus crispus; (4) Zostera marina; (5) Laminaria digitata with L. longicrusis; (6) Laminaria longicruris; (7) L. longicruris with Agarum cribrosum; (8) Agarum cribrosum with Ptilota serrata. Zostera occurred at the same level as O. crispus but replaced it in sheltered water. Ascophyllum was more abundant in sheltered water. L. digitata was confined to the more exposed, steeper shore. The average horizontal extent of the seaweed zone was 369 m, and the greatest depth of significant amounts of seaweed 20 to 30 m. Laminaria longicruris contributed 36% of the total biomass, and Laminaria spp. and Agarum together constituted 83%, while intertidal seaweeds contributed less than 10% of the biomass. The estimated average total biomass per m of shore line was 1,481 kg fresh weight, 326 kg dry weight, 98 kg carbon, or 980x10³ kcal. When averaged over the whole area of the bay, the corresponding figures were 1.38 kg/m² fresh weight, 0.30 kg/m² dry weight, 91 g/m² carbon or 912 kcal/m². Dry matter of Laminaria was 15 to 27% of fresh weight in blades, 10 to 12% in stipes. The dry matter content of blades was least in spring and highest in autumn, but carbon content and calorific value of dry matter showed little difference with species or season.Contribution to the International Biological Programme CCIBP 108.Bedford Institute Contribution BI 249.     

Ecological energetics of the sea-weed zone in a marine bay on the Atlantic coast of Canada. II. Productivity of the seaweeds

The growth of the seaweeds Laminaria longicruris, L. digitata and Agarum cribrosum were followed by SCUBA divers for two years, by punching holes 10 cm from the junction of stipe and blade, and recording at intervals the distance the holes had moved. As the holes approached the tip of the blade, new holes were punched at the base. It was found that the blades behaved like moving belts of tissue, eroding at the tips while growing at the bases, so that a total year's growth amounted to 1 to 5 times the initial length. Larger, older plants had wider and thicker blades, and the biomassincrease was roughly proportional to the square of the lengthincrease. Growth was most rapid in January to April, slow in July to October. As a conservative estimate, the summer biomass of the various species of seaweeds had a turnover of 4 to 10 times in the course of 1 year. Applying these figures to an earlier survey of biomass, it is estimated that annual production of seaweeds in St. Margaret's Bay, Nova Scotia, Canada, amounts to 603 gC/m² averaged over the whole bay, 1750 gC/m² averaged over the sea-weed zone, or 648.000 gC/m of shore line. Approximate calorific equivalents are 6030 kcal/m² over the whole bay, or 6480×10³ kcal/m of shore line.Contribution to the International Biological Program CCIBP No. 130, and Bedford Institute Contribution.     

Tracking bioaccumulation in aquatic organisms: A dynamic model integrating life history characteristics and environmental change

We have developed a dynamic model to track the evolution of contaminant concentration in an aquatic organism as a function of season and ontogeny throughout its life cycle. We have focused our analysis on the round goby (Apollonia melanostoma), a globally distributed invasive forage fish. By integrating bioenergetics with a bioaccumulation model, we illustrate how life history characteristics interact to influence contaminant accumulation. We use uncertainty and sensitivity analyses to assess how the model output is affected by uncertainty and variability in model parameters. We then demonstrate the influence of important physiological characteristics on contaminant accumulation with two scenarios. First, we probe the influence of sexual dimorphism by comparing gender-specific accumulation of a standard polychlorinated biphenyl congener, PCB153, in male and female round gobies. We hypothesize that lipid loss in female gobies during spawning season leads to a decrease in the PCB body burden compared to male gobies. Second, we compare PCB accumulation in the round goby and in the mottled sculpin (Cottus bairdi), the native forage fish that the round goby displaced in southern Lake Michigan, to determine whether the invasive species has an intrinsically different bioaccumulation potential than the native one. Our non-intuitive findings from these simulations illustrate how the interaction of growth rate with other life history characteristics lead to unexpected bioaccumulation patterns. The model we present is a flexible tool that integrates complex and dynamic interactions among environmental parameters, thus providing a means to better assess the potential for chemical accumulation in human and wildlife populations, and aiding the development of ecological forecasts.     

Abundance of the New Zealand subantarctic southern right whale population estimated from photo-identification and genotype mark-recapture

The abundance of New Zealand subantarctic southern right whales (Eubalaena australis) was estimated for the first time using mark-recapture methods based on photo-identification and microsatellite genotyping (13 loci). Individual identification photographs of 383 whales and microsatellite genotypes of 235 whales were collected during annual austral winter field surveys from 1995 to 1998. Given the 4-year survey period and lack of geographic and demographic closure, we estimated super-population abundance using the POPAN Jolly-Seber model implemented in the software programme MARK. Models with constant survivorship but time-varying capture probability and probability of entry into the population were the most suitable due to the survey design. This provided estimates of abundance in 1998 of 908 non-calf whales (95% C.L. = 755, 1,123) for the photo-identification and 910 non-calf whales (95% C.L. = 641, 1,354) for the microsatellite genotype data sets. The current estimate of 900 whales may represent less than 5% of the pre-whaling abundance in New Zealand waters.     

A goal interval programming model for resource allocation in a marine environmental protection program

A multidimensional “goal programming” model is developed to aid resource allocation decisions in the U. S. Coast Guard's Marine Environmental Protection (MEP) program. It is then extended to a model of “goal interval programming” (GIP ) type where exact values for the indicated goals, as in ordinary goal programming, are replaced by ranges. Deviations outside these ranges are also accommodated by piecewise linear functions with slopes that vary with distance from the goal intervals. Uses and generalizations are discussed in the context of applications to allocating manhours and planning the activities of the Coast Guard's MEP program.     

A proteomic approach to the study of the marine mussels Mytilus edulis and M. galloprovincialis

In this study, a proteomic approach was applied for the generation of reference maps and subsequently to detect, quantify and compare the global protein expression between two related species of marine mussels, Mytilus edulis and Mytilus galloprovincialis, growing in their own geographical habitats. A comparative study of the protein profiles generated from analytical two-dimensional electrophoresis gels was performed, and changes in protein expression were analysed quantitatively by computer analysis. An average of 1,278 spots per gel was detected in 16 individuals (8 M. edulis and 8 M. galloprovincialis); however, not all spots were included in the study. Expression of 420 spots was compared, and significant differences in the intensity levels were detected in 37 protein spots (8.8%). Fifteen proteins showed higher expression in M. edulis, and 22 proteins, in M. galloprovincialis. The technique of peptide mass fingerprinting using MALDI-TOF (matrix-assisted laser desorption ionisation/time-of-flight) and/or nanoelectrospray double subfragmentation mass spectrometry enabled the unambiguous identification of 15 of these 37 differentially expressed proteins. Most of the identified proteins can be grouped basically into four broad functional classes: cytoskeletal and myofibrillar proteins, proteins associated with stress response, proteins associated with the storage or production of energy, and proteins related to rearrangement in the synthesis of native structures. These results expand our understanding of the molecular differentiation of the two mussel taxa and serve as a useful base for future ecological, physiological and genetic studies.     

A dynamic model of mating behavior in digger wasps: the energetics of male-male competition mimic size-dependent thermal constraints

I develop a state-based dynamic model of behavior to demonstrate that size-dependent differences in temperature tolerances are not necessary to account for the activity of small male digger wasps late in the day. In the model, males defend or patrol the nesting area, wait near nests, or feed away from the nesting area depending on time of day, energy reserves and size rank. I assume a large male competitive advantage, so mating opportunities decrease with size rank for territorial or patrolling males and are rare for all waiting males; the costs of patrolling or defense are higher than the costs of waiting. If energy reserves of all males are initially small, all males alternate feeding and territorial or patrolling behavior. If energy reserves are initially large, large males patrol or maintain territories until they risk starvation and leave the area to feed. At this time, smaller males that have conserved their resources by waiting and feeding may defend territories or patrol. I simulate the behavior of three populations representing two species of Microbembex by assuming large initial energy reserves for populations in which males were territorial and small initial reserves for populations in which males patrolled, and then convert the predicted time of activity to temperature using local regressions from field studies. Temporal patterns in the activity of large and small males were similar to those actually observed, and relationships between size and temperature predicted by the model corresponded to most observations and were sometimes positive. Thus, the delayed activity of smaller males does not correspond to activity at higher temperatures and is probably not attributable to size-dependent thermal tolerances, but may represent a temporal displacement of mating activity due to intra-sexual competition and mediated by energetics. The model makes testable predictions on the timing of feeding and depletion of energy reserves in relation to size and initial energy state, and suggests how differences among species may influence the temporal and spatial organization of male mating behavior. Received: 27 February 1997 / Accepted after revision: 26 July 1997     

A mathematical model of competition and colonization in a community of marine benthic algae

A mathematical model has been constructed for the algal community on the rocky shores of a Norwegian fjord. We report here on the studies of competition and colonization along a vertical transect from the upper intertidal to the sublittoral habiats. Results on species abundance and distribution (patterns of zonation) and time to reach maturity have been compared to observations both in the fjord area and in other rocky shore areas.Competition coefficients for the algae were inferred from plant morphology and shown to be in agreement with observations of algal abundance and their zone-forming ability. Competition restricts the distribution of the species, especially at the lower elevations, but does not alter their relative position. However, increasing uniform competition prolongs the time in which zone-forming can occur, and it also decreases the overall biomass which an area can sustain. Colonization by a single species may create transient stages in community development of the same order of magnitude as algae longevity, and probably also alters the zonation pattern to some degree.The simulation results indicate that the large-scale algal distribution pattern in the Hardangerfjord area results from global stability of the rocky shore community.     

Stable isotopes in southern right whale (Eubalaena australis) baleen as indicators of seasonal movements,feeding and growth

Ratios of stable carbon and nitrogen isotopes were examined in the baleen of 11 southern right whales originating from South Africa, including one neonate, six juveniles and four adults. Oscillations in carbon isotope ratios were marked, and indicated feeding north of or at the Subtropical Convergence (STC) alternating with feeding south of the STC. There was an inverse relationship in juveniles between the periodicity of the oscillations and the length of the baleen plate, indicating a reduction in baleen growth with age. The size of the periodicity predicted for the smallest juvenile plate was equivalent to the length of the baleen at 1 yr of age as estimated from the rate of baleen growth in calves, suggesting that the oscillations in carbon isotope ratios were annual events. On this assumption, the timing of the formation of the most recent carbon enrichment peak could be calculated for each individual, given the date of death and the rate of baleen growth in the preceding year; formation occurred over a period of 100 d from January to April (mean in February). A similar analysis indicated that valleys in the nitrogen isotope ratios were formed between January and June (mean in April), and enrichment peaks between August and May (mean in December). These patterns were not inconsistent with previous scenarios of southern right whale migration, if the enrichment peaks in carbon isotopes were taken to represent feeding just north of the STC, the subsequent decline in enrichment levels to represent feeding south of the STC in autumn, and the persistence of an isotopic signature characteristic of high latitude plankton throughout the winter and early spring to indicate that feeding essentially ceased when the northern migration began, and did not resume until the southern migration was under way. The oscillations in nitrogen isotope ratios would support this interpretation, if they were assumed to represent cycles of starvation and recovery. A comparison with baleen growth rates for bowhead whales allowed the ages of the six juveniles to be assessed. Their size at age, when compared to the lengths and growth rates of calves measured photogrammetrically, suggested that growth in body length of southern right whales slows markedly between weaning and 1 yr, and may be almost negligible from 1 to 4 yr of age.     

Cadmium and lead bioaccumulation in male chickens for high food concentrations

Cadmium and lead bioaccumulation in broilers fed on rations containing high concentrations of Cd and Pb was studied. The degree of bioaccumulation was assessed by using klark of concentration (Kc). It was established that the 100 and 200 fold increase of lead and cadmium in the ration resulted in their increase in the muscles, the bones and the liver, but the degree of bioaccumulation was considerably lower than the increase of both of the heavy elements in the ration indicated by the lower Kc. A conclusion was drawn that the organism possesses adaptation mechanisms for limitation of cadmium and lead bioaccumulation through these mechanisms are not efficient enough for high ration doses of both elements. High ration doses resulted in bioaccumulation exceeding the safe for humans concentrations of lead in the bones and of cadmium in the muscles (for doses 200 fold higher than the highest Bulgarian State standard 13426–77 permissible concentrations /HPC/) and in the liver for doses 100 fold higher than HPC.     

Mercury bioaccumulation and trophic transfer in sympatric snapper species from the Gulf of Mexico.

Consumption of marine fish is a major route of toxic methyl mercury (MeHg) exposure to ocean apex predators and human populations. Here we explore the influence of trophic structure on total mercury (Hg) accumulation in red snapper (RS, Lutjanus campechanus) and gray snapper (GS, Lutjanus griseus) from the coastal Louisiana region of the Gulf of Mexico, west of the Mississippi River. The objectives of this investigation were to: (1) determine the effectiveness of the use of offshore recreational fishing charter boats and marinas as sources of fish samples and (2) compare species differences in Hg bioaccumulation, trophic position, and carbon sources. Our data show that length-normalized Hg concentrations (> or = 97% as MeHg in tissue of both species) were 230% greater in GS in comparison to RS collected from the same general area. Stable C and N isotope signatures (delta15N and delta13C) indicate that GS occupy a slightly higher trophic position (approximately 30% of one trophic position higher) on the Gulf food web in comparison to RS and that GS appear to incorporate higher trophic positioned prey, continually and at smaller sizes. Mercury was strongly correlated with combined delta15N and delta13C in pooled species data, arguing that most of the substantial difference in Hg bioaccumulation between RS and GS can be explained by modest differences in their trophic position and, to a lesser degree, carbon sources, which had low variation and high overlap among species. These observations demonstrate that even minor to moderate differences in trophic position and food habits in sympatric species can create relatively large differences in bioaccumulation regimes and underscores the importance of quantitative characterization of trophic structure in marine MeHg bioaccumulation studies.     

Application of the simple charge-discharge metabolic model to transient behavior of experimental marine microcosms

A simple charge-discharge circuit model conventionally used to simulate steady-state system metabolism was extended to simulate metabolic responses to perturbations in energy flow rates in balanced benthic marine laboratory microcosms. Empirical determination of the metabolic transfer coefficients using data collected with diel metabolic experiments indicated that these transfer coefficients are a property of the system and not of the environmental conditions to which the system had been acclimated.Used to simulate steady-state metabolic conditions based on various levels of light-energy input and temperature, the model gave values for integrated production and respiration that closely approximated experimental determinations. Moreover, the diel metabolic patterns produced were similar to those observed in the laboratory systems. The model's dynamic behavior upon loss of energy input was similar to that observed in the experimental systems for several days, but diverged after that period. Daytime respiration values were similar in magnitude but different in overall pattern from those observed in the microcosms.Simulations of a series of energy flow rate perturbation experiments produced response curves which were virtually indentical to final steady-state metabolic levels observed in the experiments, and which exhibited initial metabolic transients of the same pattern that had occurred in the laboratory. However, the transient response of the model was of greater amplitude and more highly damped.