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      Evolution of ungulate mating systems: Integrating social and environmental factors

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          Abstract

          Ungulates exhibit diverse mating systems that range from monogamous pair territories to highly polygynous leks. We review mating systems and behaviors across ungulates and offer a new approach synthesizing how interacting factors may shape those mating systems. Variability exists in mating systems among and within species of ungulates and likely is affected by predation risk, availability of resources (food and mates), habitat structure, and sociality. Ungulate mating systems may be labile as a consequence of the varying strength of those interacting factors. In addition, degree of polygyny and sexual dimorphism in size are associated with the evolution of mating systems. Neither male–male combat nor paternal care, however, can completely explain differences in sexual size dimorphism for ungulates, a necessary component in understanding the development of some mating systems. Whatever the evolutionary pathway, sexual segregation limits paternal care allowing more intense male–male competition. Selection of habitat structure, because it modifies risk of predation, is a major determinant of sociality for ungulates. Likewise, ruggedness and steepness of terrain limit the types of mating systems that can occur because of limitations in group size and cohesiveness, as well as the ability of males to herd even small groups of females effectively. The quality and defensibility of resources affect mating systems, as does the defensibility of females. Population density of females also may be a critical determinant of the types of mating systems that develop. Size of groups likewise constrains the types of mating tactics that males can employ. Our aim was to use those relationships to create a broad conceptual model that predicts how various environmental and social factors interact to structure mating systems in ungulates. This model provides a useful framework for future tests of the roles of both ecological and social conditions in influencing the social systems of ungulates.

          Abstract

          We provide a new approach to understanding mating systems in ungulates. We integrate both social and environmental factors in doing so. We develop a predictive model to help understand the evolution of these unique behaviors, which should help to direct future studies.

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          Most cited references156

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          Alternative reproductive strategies and tactics: diversity within sexes.

          Mart Gross (1996)
          Not all members of a sex behave in the same way. Frequency- and statusdependent selection have given rise to many alternative reproductive phenotypes within the sexes. The evolution and proximate control of these alternatives are only beginning to be understood. Although game theory has provided a theoretical framework, the concept of the mixed strategy has not been realized in nature, and alternative strategies are very rare. Recent findings suggest that almost all alternative reproductive phenotypes within the sexes are due to alternative tactics within a conditional strategy, and, as such, while the average fitnesses of the alternative phenotypes are unequal, the strategy is favoured in evolution. Proximate mechanisms that underlie alternative phenotypes may have many similarities with those operating between the sexes.
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            Grazing as an Optimization Process: Grass-Ungulate Relationships in the Serengeti

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              Female competition and its evolutionary consequences in mammals.

              Following Darwin's original insights regarding sexual selection, studies of intrasexual competition have mainly focused on male competition for mates; by contrast, female reproductive competition has received less attention. Here, we review evidence that female mammals compete for both resources and mates in order to secure reproductive benefits. We describe how females compete for resources such as food, nest sites, and protection by means of dominance relationships, territoriality and inter-group aggression, and by inhibiting the reproduction of other females. We also describe evidence that female mammals compete for mates and consider the ultimate causes of such behaviour, including competition for access to resources provided by mates, sperm limitation and prevention of future resource competition. Our review reveals female competition to be a potentially widespread and significant evolutionary selection pressure among mammals, particularly competition for resources among social species for which most evidence is currently available. We report that female competition is associated with many diverse adaptations, from overtly aggressive behaviour, weaponry, and conspicuous sexual signals to subtle and often complex social behaviour involving olfactory signalling, alliance formation, altruism and spite, and even cases where individuals appear to inhibit their own reproduction. Overall, despite some obvious parallels with male phenotypic traits favoured under sexual selection, it appears that fundamental differences in the reproductive strategies of the sexes (ultimately related to parental investment) commonly lead to contrasting competitive goals and adaptations. Because female adaptations for intrasexual competition are often less conspicuous than those of males, they are generally more challenging to study. In particular, since females often employ competitive strategies that directly influence not only the number but also the quality (survival and reproductive success) of their own offspring, as well as the relative reproductive success of others, a multigenerational view ideally is required to quantify the full extent of variation in female fitness resulting from intrasexual competition. Nonetheless, current evidence indicates that the reproductive success of female mammals can also be highly variable over shorter time scales, with significant reproductive skew related to competitive ability. Whether we choose to describe the outcome of female reproductive competition (competition for mates, for mates controlling resources, or for resources per se) as sexual selection depends on how sexual selection is defined. Considering sexual selection strictly as resulting from differential mating or fertilisation success, the role of female competition for the sperm of preferred (or competitively successful) males appears particularly worthy of more detailed investigation. Broader definitions of sexual selection have recently been proposed to encompass the impact on reproduction of competition for resources other than mates. Although the merits of such definitions are a matter of ongoing debate, our review highlights that understanding the evolutionary causes and consequences of female reproductive competition indeed requires a broader perspective than has traditionally been assumed. We conclude that future research in this field offers much exciting potential to address new and fundamentally important questions relating to social and mating-system evolution. © 2010 The Authors. Biological Reviews © 2010 Cambridge Philosophical Society.
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                Author and article information

                Contributors
                bowyterr@isu.edu
                Journal
                Ecol Evol
                Ecol Evol
                10.1002/(ISSN)2045-7758
                ECE3
                Ecology and Evolution
                John Wiley and Sons Inc. (Hoboken )
                2045-7758
                15 April 2020
                June 2020
                : 10
                : 11 ( doiID: 10.1002/ece3.v10.11 )
                : 5160-5178
                Affiliations
                [ 1 ] Institute of Arctic Biology University of Alaska Fairbanks Fairbanks AK USA
                [ 2 ] Department of Environmental Science, Policy and Management University of California Berkeley CA USA
                [ 3 ] Department of Fish and Wildlife Sciences University of Idaho Moscow ID USA
                [ 4 ] Laboratory of Wildlife Ecology and Behaviour School of Biology and Environmental Science University College Dublin Dublin Ireland
                [ 5 ] Department of Biology Brigham Young University‐Idaho Rexburg ID USA
                Author notes
                [*] [* ] Correspondence

                R. Terry Bowyer, Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, AK 99775, USA.

                Email: bowyterr@ 123456isu.edu

                Author information
                https://orcid.org/0000-0002-1398-4104
                Article
                ECE36246
                10.1002/ece3.6246
                7297761
                32551090
                96dea830-d687-4d79-8dc9-a4bf33a0aaac
                © 2020 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
                : 11 October 2019
                : 12 March 2020
                : 13 March 2020
                Page count
                Figures: 7, Tables: 0, Pages: 19, Words: 16564
                Categories
                Review
                Review
                Custom metadata
                2.0
                June 2020
                Converter:WILEY_ML3GV2_TO_JATSPMC version:5.8.4 mode:remove_FC converted:16.06.2020

                Evolutionary Biology
                mating,monogamy,polygyny,social system,terrain,territoriality
                Evolutionary Biology
                mating, monogamy, polygyny, social system, terrain, territoriality

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