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      Modeling the demography of species providing extended parental care: A capture–recapture multievent model with a case study on polar bears ( Ursus maritimus)

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          Abstract

          1. In species providing extended parental care, one or both parents care for altricial young over a period including more than one breeding season. We expect large parental investment and long‐term dependency within family units to cause high variability in life trajectories among individuals with complex consequences at the population level. So far, models for estimating demographic parameters in free‐ranging animal populations mostly ignore extended parental care, thereby limiting our understanding of its consequences on parents and offspring life histories.

          2. We designed a capture–recapture multievent model for studying the demography of species providing extended parental care. It handles statistical multiple‐year dependency among individual demographic parameters grouped within family units, variable litter size, and uncertainty on the timing at offspring independence. It allows for the evaluation of trade‐offs among demographic parameters, the influence of past reproductive history on the caring parent's survival status, breeding probability, and litter size probability, while accounting for imperfect detection of family units. We assess the model performance using simulated data and illustrate its use with a long‐term dataset collected on the Svalbard polar bears ( Ursus maritimus).

          3. Our model performed well in terms of bias and mean square error and in estimating demographic parameters in all simulated scenarios, both when offspring departure probability from the family unit occurred at a constant rate or varied during the field season depending on the date of capture. For the polar bear case study, we provide estimates of adult and dependent offspring survival rates, breeding probability, and litter size probability. Results showed that the outcome of the previous reproduction influenced breeding probability.

          4. Overall, our results show the importance of accounting for i) the multiple‐year statistical dependency within family units, ii) uncertainty on the timing at offspring independence, and iii) past reproductive history of the caring parent. If ignored, estimates obtained for breeding probability, litter size, and survival can be biased. This is of interest in terms of conservation because species providing extended parental care are often long‐living mammals vulnerable or threatened with extinction.

          Abstract

          We developed a capture–recapture model for studying the demography of species providing extended parental care. Our model handles statistical dependency among individual demographic parameters and life histories within family units until offspring independence, and include variable litter size, age at offspring independence and breeding frequency. It also includes the influence of past reproductive history on the caring parent's survival status and current reproduction, while accounting for imperfect detection of family units. We assess the model performances using simulated data, and illustrate its use with a long‐term dataset collected on the Svalbard polar bears ( Ursus maritimus).

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          Most cited references48

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          Multievent: an extension of multistate capture-recapture models to uncertain states.

          Capture-recapture models were originally developed to account for encounter probabilities that are less than 1 in free-ranging animal populations. Nowadays, these models can deal with the movement of animals between different locations and are also used to study transitions between different states. However, their use to estimate transitions between states does not account for uncertainty in state assignment. I present the extension of multievent models, which does incorporate this uncertainty. Multievent models belong to the family of hidden Markov models. I also show in this article that the memory model, in which the next state or location is influenced by the previous state occupied, can be fully treated within the framework of multievent models.
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            Natural Selection, the Costs of Reproduction, and a Refinement of Lack's Principle

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              Immobilization of polar bears (Ursus maritimus) with Telazol in the Canadian Arctic.

              In 1986, 213 polar bears (Ursus maritimus) were immobilized with Telazol on the sea ice of the eastern Beaufort Sea during April and May, and 106 along the western coast of Hudson Bay near Churchill, Manitoba (Canada) in September. No animals died from handling. The efficacy of this drug at different seasons and the physiological responses of the immobilized bears were compared. A single injection of 8 to 9 mg of Telazol per kg of body weight gave a rapid full immobilization with satisfactory analgesia, and faster recovery than other drugs for which there is no antagonist. The reactions of the bears could be reliably and easily interpreted from a safe distance before the animal was approached. There was a wide range of tolerance to high dosages and bears appeared able to thermoregulate while immobilized. The mortality rate due to handling was lower than with any other drug used to date.
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                Author and article information

                Contributors
                sarah.cubaynes@ephe.psl.eu
                Journal
                Ecol Evol
                Ecol Evol
                10.1002/(ISSN)2045-7758
                ECE3
                Ecology and Evolution
                John Wiley and Sons Inc. (Hoboken )
                2045-7758
                10 March 2021
                April 2021
                : 11
                : 7 ( doiID: 10.1002/ece3.v11.7 )
                : 3380-3392
                Affiliations
                [ 1 ] CEFE Univ Montpellier CNRS EPHE‐PSL University IRD Univ Paul Valéry Montpellier 3 Montpellier France
                [ 2 ] Norwegian Polar Institute FRAM Centre Tromsø Norway
                [ 3 ] Department of Arctic and Marine Biology UiT The Arctic University of Norway Tromsø Norway
                [ 4 ] Natural History Museum University of Oslo Oslo Norway
                Author notes
                [*] [* ] Correspondence

                Sarah Cubaynes, CEFE, Univ Montpellier, CNRS, EPHE‐PSL University, IRD, Univ Paul Valéry Montpellier 3, Montpellier, France.

                Email: sarah.cubaynes@ 123456ephe.psl.eu

                Author information
                https://orcid.org/0000-0002-3935-9825
                https://orcid.org/0000-0002-0574-6712
                https://orcid.org/0000-0003-2192-1039
                https://orcid.org/0000-0002-2684-9251
                https://orcid.org/0000-0003-0395-5251
                https://orcid.org/0000-0001-7001-5142
                Article
                ECE37296
                10.1002/ece3.7296
                8019049
                33841791
                d2e99f07-d97b-4982-ba0a-16c0887f815f
                © 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

                This is an open access article under the terms of the http://creativecommons.org/licenses/by/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 07 January 2021
                : 24 November 2020
                : 25 January 2021
                Page count
                Figures: 3, Tables: 2, Pages: 13, Words: 9473
                Funding
                Funded by: Agence Nationale de la Recherche , open-funder-registry 10.13039/501100001665;
                Award ID: ANR‐18‐CE02‐0011
                Funded by: World Wildlife Fund , open-funder-registry 10.13039/100001399;
                Categories
                Original Research
                Original Research
                Custom metadata
                2.0
                April 2021
                Converter:WILEY_ML3GV2_TO_JATSPMC version:6.0.1 mode:remove_FC converted:03.04.2021

                Evolutionary Biology
                apex predator,arctic ecosystem,bayesian modeling,capture–recapture,dependency among individuals,family structure,parental care,state uncertainty,timing at independence

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