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      Long-term historical and projected herbivore population dynamics in Ngorongoro crater, Tanzania

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          Abstract

          The Ngorongoro Crater is an intact caldera with an area of approximately 310 km 2 located within the Ngorongoro Conservation Area (NCA) in northern Tanzania. It is known for the abundance and diversity of its wildlife and is a UNESCO World Heritage Site and an International Biosphere Reserve. Long term records (1963–2012) on herbivore populations, vegetation and rainfall made it possible to analyze historic and project future herbivore population dynamics. NCA was established as a multiple use area in 1959. In 1974 there was a perturbation in that resident Maasai and their livestock were removed from the Ngorongoro Crater. Thus, their pasture management that was a combination of livestock grazing and fire was also removed and 'burning' stopped being a regular occurrence until it was resumed in 2001 by NCA management. The Maasai pasture management would have selected for shorter grasses and more palatable species. Vegetation mapping in 1966–1967 recorded predominately short grasslands. Subsequent vegetation mapping in the crater in 1995 determined that the grassland structure had changed such that mid and tall grasses were dominant. After removal of the Maasai pastoralists from the Ngorongoro Crater in 1974, there were significant changes in population trends for some herbivore species. Buffalo, elephant and ostrich numbers increased significantly during 1974–2012. The zebra population was stable from 1963 to 2012 whereas population numbers of five species declined substantially between 1974 and 2012 relative to their peak numbers during 1974–1976. Grant’s and Thomson’s gazelles, eland, kongoni, and waterbuck (wet season only) declined significantly in the Crater in both seasons after 1974. In addition, some herbivore species were consistently more abundant inside the Crater during the wet than the dry season. This pattern was most evident for the large herbivore species requiring bulk forage, i.e., buffalo, eland, and elephant. Even with a change in grassland structure, total herbivore biomass remained relatively stable from 1963 to 2012, implying that the crater has a stable carrying capacity. Analyses of rainfall indicated that there was a persistent cycle of 4.83 years for the annual component. Herbivore population size was correlated with rainfall in both the wet and dry seasons. The relationships established between the time series of historic animal counts in the wet and dry seasons and lagged wet and dry season rainfall series were used to forecast the likely future trajectories of the wet and dry season population size for each species under three alternative climate change scenarios.

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          RCP 8.5—A scenario of comparatively high greenhouse gas emissions

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            Small sample inference for fixed effects from restricted maximum likelihood.

            Restricted maximum likelihood (REML) is now well established as a method for estimating the parameters of the general Gaussian linear model with a structured covariance matrix, in particular for mixed linear models. Conventionally, estimates of precision and inference for fixed effects are based on their asymptotic distribution, which is known to be inadequate for some small-sample problems. In this paper, we present a scaled Wald statistic, together with an F approximation to its sampling distribution, that is shown to perform well in a range of small sample settings. The statistic uses an adjusted estimator of the covariance matrix that has reduced small sample bias. This approach has the advantage that it reproduces both the statistics and F distributions in those settings where the latter is exact, namely for Hotelling T2 type statistics and for analysis of variance F-ratios. The performance of the modified statistics is assessed through simulation studies of four different REML analyses and the methods are illustrated using three examples.
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              Implications of limiting CO2 concentrations for land use and energy.

              Limiting atmospheric carbon dioxide (CO2) concentrations to low levels requires strategies to manage anthropogenic carbon emissions from terrestrial systems as well as fossil fuel and industrial sources. We explore the implications of fully integrating terrestrial systems and the energy system into a comprehensive mitigation regime that limits atmospheric CO2 concentrations. We find that this comprehensive approach lowers the cost of meeting environmental goals but also carries with it profound implications for agriculture: Unmanaged ecosystems and forests expand, and food crop and livestock prices rise. Finally, we find that future improvement in food crop productivity directly affects land-use change emissions, making the technology for growing crops potentially important for limiting atmospheric CO2 concentrations.
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                Author and article information

                Contributors
                Role: ConceptualizationRole: Data curationRole: Funding acquisitionRole: InvestigationRole: Project administrationRole: ResourcesRole: SupervisionRole: ValidationRole: Writing – original draftRole: Writing – review & editing
                Role: ConceptualizationRole: Data curationRole: Formal analysisRole: Funding acquisitionRole: InvestigationRole: MethodologyRole: Project administrationRole: ResourcesRole: SoftwareRole: SupervisionRole: ValidationRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: InvestigationRole: MethodologyRole: ResourcesRole: SoftwareRole: Writing – review & editing
                Role: Data curationRole: InvestigationRole: Writing – review & editing
                Role: InvestigationRole: MethodologyRole: Writing – review & editing
                Role: InvestigationRole: MethodologyRole: VisualizationRole: Writing – review & editing
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                10 March 2020
                2020
                : 15
                : 3
                : e0212530
                Affiliations
                [1 ] IUCN/SSC Equid Specialist Group, Arusha, Arusha, Tanzania
                [2 ] Institute for Crop Science-340, University of Hohenheim, Stuttgart, Germany
                [3 ] Environmental Reliance Consultants Limited, Arusha, Tanzania
                [4 ] Department of Ecosystem Science and Sustainability, Colorado State University, Fort Collins, Colorado, United States of America
                [5 ] Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, Colorado, United States of America
                Texas State University, UNITED STATES
                Author notes

                Competing Interests: The authors have declared that no competing interests exist

                ‡ PDM and JOO are joint first authors on this work.

                Author information
                http://orcid.org/0000-0002-7379-0387
                Article
                PONE-D-19-03269
                10.1371/journal.pone.0212530
                7064247
                32155150
                1ba016b5-589c-4f22-bc1c-a67117d8e54f
                © 2020 Moehlman et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 2 February 2019
                : 10 February 2020
                Page count
                Figures: 11, Tables: 1, Pages: 39
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/501100001659, Deutsche Forschungsgemeinschaft;
                Award ID: OG 83/1-2
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/501100007601, Horizon 2020;
                Award ID: 641918
                Award Recipient :
                JOO was supported by a grant from the German Research Foundation (Grant No. OG 83/1-1). JOO was also supported by The Planning for Resilience in East Africa through Policy, Adaptation, Research, and Economic Development (USAID PREPARED) project. This project has received funding from the European Union’s Horizon 2020 research and innovation programme under Grant Agreement No. 641918.
                Categories
                Research Article
                Biology and Life Sciences
                Ecology
                Plant Ecology
                Plant-Animal Interactions
                Herbivory
                Ecology and Environmental Sciences
                Ecology
                Plant Ecology
                Plant-Animal Interactions
                Herbivory
                Biology and Life Sciences
                Plant Science
                Plant Ecology
                Plant-Animal Interactions
                Herbivory
                Biology and Life Sciences
                Ecology
                Community Ecology
                Trophic Interactions
                Herbivory
                Ecology and Environmental Sciences
                Ecology
                Community Ecology
                Trophic Interactions
                Herbivory
                Earth Sciences
                Seasons
                Biology and Life Sciences
                Organisms
                Eukaryota
                Animals
                Vertebrates
                Amniotes
                Mammals
                Wildebeest
                Biology and Life Sciences
                Organisms
                Eukaryota
                Animals
                Vertebrates
                Amniotes
                Mammals
                Equines
                Zebras
                Biology and Life Sciences
                Population Biology
                Population Metrics
                Population Size
                Biology and Life Sciences
                Population Biology
                Population Dynamics
                Biology and Life Sciences
                Organisms
                Eukaryota
                Animals
                Vertebrates
                Amniotes
                Mammals
                Elephants
                Biology and Life Sciences
                Agriculture
                Livestock
                Custom metadata
                All relevant data are within the manuscript and its Supporting Information files.

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