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      Sixteen years later: Occurrence, group size, and habitat use of humpback dolphins (Sousa plumbea) in Algoa Bay, South Africa

      1 , 2 , 3 , 4
      Marine Mammal Science
      Wiley

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          Decline in relative abundance of bottlenose dolphins exposed to long-term disturbance.

          Studies evaluating effects of human activity on wildlife typically emphasize short-term behavioral responses from which it is difficult to infer biological significance or formulate plans to mitigate harmful impacts. Based on decades of detailed behavioral records, we evaluated long-term impacts of vessel activity on bottlenose dolphins (Tursiops sp.) in Shark Bay, Australia. We compared dolphin abundance within adjacent 36-km2 tourism and control sites, over three consecutive 4.5-year periods wherein research activity was relatively constant but tourism levels increased from zero, to one, to two dolphin-watching operators. A nonlinear logistic model demonstrated that there was no difference in dolphin abundance between periods with no tourism and periods in which one operator offered tours. As the number of tour operators increased to two, there was a significant average decline in dolphin abundance (14.9%; 95% CI=-20.8 to -8.23), approximating a decline of one per seven individuals. Concurrently, within the control site, the average increase in dolphin abundance was not significant (8.5%; 95% CI=-4.0 to +16.7). Given the substantially greater presence and proximity of tour vessels to dolphins relative to research vessels, tour-vessel activity contributed more to declining dolphin numbers within the tourism site than research vessels. Although this trend may not jeopardize the large, genetically diverse dolphin population of Shark Bay, the decline is unlikely to be sustainable for local dolphin tourism. A similar decline would be devastating for small, closed, resident, or endangered cetacean populations. The substantial effect of tour vessels on dolphin abundance in a region of low-level tourism calls into question the presumption that dolphin-watching tourism is benign.
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            Estimating equations for association structures.

            This paper investigates generalized estimating equations for association parameters, which are frequently of interest in family studies, with emphasis on covariance estimation. Separate link functions are used to connect the mean, the scale, and the correlation to linear predictors involving possibly different sets of covariates, and separate estimating equations are proposed for the three sets of parameters. Simulations show that the robust 'sandwich' variance estimator and the jackknife variance estimator for the correlation parameters are generally close to the empirical variance for the sample size of 50 clusters. The results contradict Ziegler et al. and Kastner and Ziegler, where the 'sandwich' estimator obtained from the software MAREG was shown to be unsuitable for practical usage. The problem appears to arise because the MAREG variance estimator does not account for variability in estimation of the scale parameters, but may be valid with fixed scale. We also find that the formula for the approximate jackknife variance estimator in Ziegler et al. is deficient, resulting in systematic deviations from the fully iterated jackknife variance estimator. A general jackknife formula is provided and performs well in numerical studies. Data from a study on the genetics of alcoholism is used to illustrate the importance of reliable variance estimation in biomedical applications. Copyright 2004 John Wiley & Sons, Ltd.
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              The social structure and strategies of delphinids: predictions based on an ecological framework.

              Dolphins live in complex social groupings with a wide variety of social strategies. In this chapter we investigate the role that differing habitats and ecological conditions have played in the evolution of delphinid social strategies. We propose a conceptual framework for understanding natural patterns of delphinid social structure in which the spatial and temporal predictability of resources influences the ranging patterns of individuals and communities. The framework predicts that when resources are spatially and temporally predictable, dolphins should remain resident in relatively small areas. Predictable resources are often found in complex inshore environments where dolphins may hide from predators or avoid areas with high predator density. Additionally, available food resources may limit group size. Thus, we predict that there are few benefits to forming large groups and potentially many benefits to being solitary or in small groups. Males may be able to sequester solitary females, controlling mating opportunities. Observations of inshore populations of bottlenose dolphins (Tursiops sp.) and island-associated spinner dolphins (Stenella longirostris) seem to fit this pattern well, along with forest-dwelling African antelope and primates such as vervets (Cercopithicus aethiops), baboons (Papio sp.), macaques (Macaca sp.) and chimpanzees (Pan troglodytes). In contrast, the framework predicts that when resources such as food are unpredictable, individuals must range further to find the necessary resources. Forming groups may be the only strategy available to avoid predation, especially in the open ocean. Larger home ranges are likely to support a greater number of individuals; however, prey is often sparsely distributed, which may act to reduce foraging competition. Cooperative foraging and herding of prey schools may be advantageous, potentially facilitating the formation of long-term bonds. Alternately, individuals may display many short-term affiliations. These large groups make it difficult for a male or a small group of males to sequester a female, and polygynandry is the most likely mating strategy. While it is difficult to study wide-ranging delphinids to examine these predictions, this ranging and behavioural pattern has been suggested for dusky dolphins (Lagenorhynchus obscurus), coastal bottlenose dolphins (Tursiops sp.) and mixed species of dolphins in the Eastern Tropical Pacific. These patterns also resemble the ranging and social strategies of open savannah African antelopes and desert-dwelling macropods. Resource availability exists in a range of complex distributions and we predict that delphinid ranging patterns will also vary. At intermediate-ranging patterns, the framework predicts that individuals should form mid-sized groups balancing intra-group competition with predation protection. Humpback dolphins (Sousa sp.) appear to fit this pattern, with some site fidelity over relatively large ranges. They display fluid associations with other individuals. Predation pressure is not sufficiently high to cause large groups to form, and individuals probably reduce predation pressure more by hiding whenever possible. This pattern is likely to prevent the formation of long-term complex bonds. In contrast, killer whales (Orcinus orca) also display intermediate-ranging patterns, but have extremely strong social bonds within familial groups. Cooperative and altruistic behaviour in killer whales facilitate the formation of life-long bonds, similar to those observations in sperm whales (Physeter macrocephalus) and elephants (Loxodonta africana). This conceptual framework remains largely untested, and for many species it is not currently possible to describe ranging behaviours, anti-predator tactics or social behaviour in sufficient detail for appropriate examination of these ideas. Few studies on dolphins have been conducted to explicitly test this type of framework; however, existing observations of delphinid social strategies and communities are used throughout this chapter to examine this framework. Additionally, we anticipate that the present framework may provide a starting point to test hypotheses regarding the evolution of social strategies of delphinids.
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                Author and article information

                Journal
                Marine Mammal Science
                Mar Mam Sci
                Wiley
                08240469
                April 2016
                April 2016
                October 23 2015
                : 32
                : 2
                : 490-507
                Affiliations
                [1 ]University of Groningen; Department of Ocean Ecosystems; PO Box 11103, 9700 CC Groningen The Netherlands
                [2 ]The Swire Institute of Marine Science and School of Biological Sciences; The University of Hong Kong; Cape d'Aguilar Shek O Hong Kong
                [3 ]Nelson Mandela Metropolitan University; Coastal and Marine Research Institute; PO Box 77000 Port Elizabeth 6031 South Africa
                [4 ]South African Institute for Aquatic Biodiversity (SAIAB) and South African Environmental Observation Network (SAEON); Grahamstown 6140 South Africa
                Article
                10.1111/mms.12279
                fe09f08c-81f7-4479-ae63-2d4f4c328d92
                © 2015

                http://doi.wiley.com/10.1002/tdm_license_1.1

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