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      Extraterritorial forays by great tits are associated with dawn song in unexpected ways

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          Abstract

          Conspicuous male signals often play an important role in both attracting mates and deterring rivals. In territorial species with extrapair mating, female and male forays to other territories may be an important component underlying female choice and male mating success and might be influenced by male advertisement signals. Yet, whether off-territory foraying is associated with male signals is still not well understood. Here, we tested how female and male forays are associated with short-range visual and long-range acoustic signals (dawn song). We used an automated radio tracking system to follow the movements of wild great tits ( Parus major) to other territories in relation to male dawn song, plumage ornaments, and extrapair paternity. We show that both sexes frequently forayed into others’ territories throughout the breeding period. Movements of both males and females were associated with male song but not with plumage ornaments. Contrary to our expectations, females stayed away from territories where males sang elaborately, whereas males were attracted to those territories. Moreover, neither female nor male forays were associated with the occurrence of extrapair offspring. Our results, thus, suggest that, although forays into other territories are associated with male dawn song, females may not be attracted and males not repelled by dawn song. This sheds a different light on the sex-specific effects of male advertisement signals, expanding the view on the selection pressures shaping such communication systems.

          Abstract

          Conspicuous signals of many animals, like song in birds, may attract mates and repel rivals. We tracked the movements of wild great tits ( Parus major) during an entire breeding season and show the opposite; females stayed away from territories of males with elaborate singing, whereas males were attracted. This suggests that the dawn song of male birds can have the opposite effect than commonly shown based on immediate responses, indicating a more complex role of song in communication systems.

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          Fitting Linear Mixed-Effects Models Using lme4

          Maximum likelihood or restricted maximum likelihood (REML) estimates of the parameters in linear mixed-effects models can be determined using the lmer function in the lme4 package for R. As for most model-fitting functions in R, the model is described in an lmer call by a formula, in this case including both fixed- and random-effects terms. The formula and data together determine a numerical representation of the model from which the profiled deviance or the profiled REML criterion can be evaluated as a function of some of the model parameters. The appropriate criterion is optimized, using one of the constrained optimization functions in R, to provide the parameter estimates. We describe the structure of the model, the steps in evaluating the profiled deviance or REML criterion, and the structure of classes or types that represents such a model. Sufficient detail is included to allow specialization of these structures by users who wish to write functions to fit specialized linear mixed models, such as models incorporating pedigrees or smoothing splines, that are not easily expressible in the formula language used by lmer. Journal of Statistical Software, 67 (1) ISSN:1548-7660
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            Statistical confidence for likelihood-based paternity inference in natural populations.

            Paternity inference using highly polymorphic codominant markers is becoming common in the study of natural populations. However, multiple males are often found to be genetically compatible with each offspring tested, even when the probability of excluding an unrelated male is high. While various methods exist for evaluating the likelihood of paternity of each nonexcluded male, interpreting these likelihoods has hitherto been difficult, and no method takes account of the incomplete sampling and error-prone genetic data typical of large-scale studies of natural systems. We derive likelihood ratios for paternity inference with codominant markers taking account of typing error, and define a statistic delta for resolving paternity. Using allele frequencies from the study population in question, a simulation program generates criteria for delta that permit assignment of paternity to the most likely male with a known level of statistical confidence. The simulation takes account of the number of candidate males, the proportion of males that are sampled and gaps and errors in genetic data. We explore the potentially confounding effect of relatives and show that the method is robust to their presence under commonly encountered conditions. The method is demonstrated using genetic data from the intensively studied red deer (Cervus elaphus) population on the island of Rum, Scotland. The Windows-based computer program, CERVUS, described in this study is available from the authors. CERVUS can be used to calculate allele frequencies, run simulations and perform parentage analysis using data from all types of codominant markers.
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              Complex signal function: developing a framework of testable hypotheses

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                Author and article information

                Contributors
                Role: Editor
                Journal
                Behav Ecol
                Behav. Ecol
                beheco
                Behavioral Ecology
                Oxford University Press (UK )
                1045-2249
                1465-7279
                Jul-Aug 2020
                20 May 2020
                20 May 2020
                : 31
                : 4
                : 873-883
                Affiliations
                [1 ] Behavioural Ecology Group, Wageningen University and Research , Wageningen, The Netherlands
                [2 ] Department of Animal Ecology, Netherlands Institute of Ecology (NIOO-KNAW) , Wageningen, The Netherlands
                [3 ] Behavioural Ecology Research Group, Department of Biology, Anglia Ruskin University , Cambridgeshire, UK
                Author notes
                 Address correspondence to N. Bircher. E-mail: ninabircher123@ 123456gmail.com
                Author information
                http://orcid.org/0000-0003-4963-9617
                http://orcid.org/0000-0001-6984-906X
                http://orcid.org/0000-0001-9376-4023
                http://orcid.org/0000-0003-0494-4888
                Article
                araa040
                10.1093/beheco/araa040
                7390995
                32760175
                fdee4cb1-1dfc-4a4a-b48c-8e1216b8d88d
                © The Author(s) 2020. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 02 September 2019
                : 16 January 2020
                : 29 January 2020
                : 14 April 2020
                Page count
                Pages: 11
                Funding
                Funded by: Dutch Research Council;
                Award ID: 824.15.012
                Categories
                Editor’s Choice
                Editor's Choice
                AcademicSubjects/SCI01330

                Ecology
                advertisement signals,dawn song,color ornaments,extrapair paternity,extraterritorial forays,radio tracking

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