Editing Behavioral ecology (section)

==Mating systems==
{{Main|Mating systems}}

Various types of mating systems include [[Monogamy in animals|monogamy]], [[Polygyny in animals|polygyny]], [[Polyandry in nature|polyandry]], and [[Promiscuity#Other animals|promiscuity]]. Each is differentiated by the sexual behavior between mates, such as which males mate with certain females. An influential paper by Stephen Emlen and Lewis Oring (1977)<ref>{{cite journal | last1 = Emlen | first1 = S. T. | last2 = Oring | first2 = L. W. | s2cid = 16786432 | year = 1977 | title = Ecology, sexual selection, and the evolution of mating systems | journal = Science | volume = 197 | issue = 4300| pages = 214–223 | doi=10.1126/science.327542 | pmid=327542|bibcode = 1977Sci...197..215E }}</ref> argued that two main factors of animal behavior influence the diversity of mating systems: the relative accessibility that each sex has to mates, and the parental desertion by either sex.

===Mating systems with no male parental care===
In a system that does not have male parental care, [[resource]] [[Biological dispersal|dispersion]], [[predation]], and the effects of [[Social animal|social living]] primarily influence female dispersion, which in turn influences male dispersion. Since males' primary concern is female acquisition, the males either indirectly or directly compete for the females. In direct [[competition]], the males are directly focused on the females.<ref name="Davies et al., (2012) pp. 254-263">Davies, N.B., Krebs, J.R. and West., S.A., (2012). ''An Introduction to Behavioural Ecology.'' 4th ed. John Wiley & Sons, pp. 254–263{{ISBN?}}</ref> [[Thalassoma bifasciatum|Blue-headed wrasse]] demonstrate the behavior in which females follow resources—such as good [[nest]] sites—and males follow the females.<ref name="Davies et al., (2012) pp. 254-263" /><ref>{{cite journal | last1 = Warner | first1 = R. R. | year = 1987 | title = Female choice of sites versus mates in a coral reef fish Thalassoma bifasciatum | journal = Animal Behaviour | volume = 35 | issue = 5| pages = 1470–1478 | doi=10.1016/s0003-3472(87)80019-2| s2cid = 53177154 }}</ref> Conversely, species with males that exemplify indirectly competitive behavior tend towards the males' anticipation of the resources desired by females and their subsequent effort to control or acquire these resources, which helps them to achieve success with females.<ref name="Davies et al., (2012) pp. 254-263" /> Grey-sided voles demonstrate indirect male competition for females. The males were experimentally observed to home in on the sites with the best food in anticipation of females settling in these areas.<ref name="Davies et al., (2012) pp. 254-263" /><ref>{{cite journal | last1 = Ims | first1 = R.A. | year = 1987 | title = Responses in spatial organization and behavior to manipulations of the food resource in the vole Clethrionomys rufocanus | journal = Journal of Animal Ecology | volume = 56 | issue = 2| pages = 585–596 | doi=10.2307/5070| jstor = 5070 }}</ref> Males of ''[[Euglossa imperialis]]'', a non-social bee species, also demonstrate indirect competitive behavior by forming aggregations of territories, which can be considered leks, to defend fragrant-rich primary territories. The purpose of these aggregations is largely only facultative, since the more suitable fragrant-rich sites there are, the more habitable territories there are to inhabit, giving females of this species a large selection of males with whom to potentially mate.<ref>{{cite journal | last1 = Kimsey | first1 = Lynn Siri | year = 1980 | title = The behaviour of male orchid bees (Apidae, Hymenoptera, Insecta) and the question of leks | journal = Animal Behaviour | volume = 28 | issue = 4| pages = 996–1004 | doi=10.1016/s0003-3472(80)80088-1| s2cid = 53161684 }}</ref> [[Lek mating|Leks]] and choruses have also been deemed another behavior among the phenomena of male competition for females. Due to the resource-poor nature of the territories that lekking males often defend, it is difficult to categorize them as indirect competitors. For example, the [[ghost moth]] males display in leks to attract a female mate. Additionally, it is difficult to classify them as direct competitors seeing as they put a great deal of effort into their defense of their territories before females arrive, and upon female arrival they put for the great mating displays to attract the females to their individual sites. These observations make it difficult to determine whether female or resource dispersion primarily influences male aggregation, especially in lieu of the apparent difficulty that males may have defending resources and females in such densely populated areas.<ref name="Davies et al., (2012) pp. 254-263" /> Because the reason for male aggregation into leks is unclear, five hypotheses have been proposed. These postulates propose the following as reasons for male lekking: hotspot, [[predation]] reduction, increased female attraction, hotshot males, facilitation of female choice.<ref name="Davies et al., (2012) pp. 254-263" /><ref>Bradbury, J. E. and Gibson, R. M. (1983) Leks and mate choice. In: ''Mate Choice'' (ed. P. Bateson). pp. 109–138. Cambridge University Press. Cambridge{{ISBN?}}</ref> With all of the mating behaviors discussed, the primary factors influencing differences within and between species are [[ecology]], social conflicts, and life history differences.<ref name="Davies et al., (2012) pp. 254-263" />

In some other instances, neither direct nor indirect competition is seen. Instead, in species like the [[Edith's checkerspot]] butterfly, males' efforts are directed at acquisition of females and they exhibit indiscriminate mate location behavior, where, given the low cost of mistakes, they blindly attempt to mate both correctly with females and incorrectly with other objects.<ref name=five>{{cite journal|last=Moore|first=Sandra D.|title=Male-Biased Mortality in the Butterfly ''Euphydryas editha'': a Novel Cost of Mate Acquisition|journal=The American Naturalist |year=1987 |volume=130 |issue=2 |pages=306–309 |doi=10.1086/284711|s2cid=84989304}}</ref>

===Mating systems with male parental care===

====Monogamy====
Monogamy is the mating system in 90% of birds, possibly because each male and female has a greater number of offspring if they share in raising a brood.<ref>Lack, D. (1968) Ecological Adaptations for Breeding in Birds. Methuen, London.</ref> In obligate monogamy, males feed females on the nest, or share in incubation and chick-feeding. In some species, males and females form lifelong pair bonds. Monogamy may also arise from limited opportunities for polygamy, due to strong competition among males for mates, females suffering from loss of male help, and female–female aggression.<ref name="Davies et al., (2012) pp. 266">Davies, N. B., Krebs, J. R and West, S. A., (2012). ''An Introduction to Behavioral Ecology.'' West Sussex, UK: Wiley-Blackwell. p. 266. {{ISBN|978-1-4051-1416-5}}.</ref>

====Polygyny====
In birds, polygyny occurs when males indirectly monopolize females by controlling resources. In species where males normally do not contribute much to parental care, females suffer relatively little or not at all.<ref>{{cite journal | last1 = Lightbody | first1 = J.P. | last2 = Weatherhead | first2 = P.J. | year = 1988 | title = Female settling patterns and polygyny: tests of a neutral-mate-choice hypothesis | journal = American Naturalist | volume = 132 | pages = 20–33 | doi=10.1086/284835| s2cid = 84147769 }}</ref> In other species, however, females suffer through the loss of male contribution, and the cost of having to share resources that the male controls, such as nest sites or food. In some cases, a polygynous male may control a high-quality territory so for the female, the benefits of polygyny may outweigh the costs.<ref>{{cite journal | last1 = Verner | first1 = J. | last2 = Wilson | first2 = M.F. | year = 1966 | title = The influence of habitats on mating systems of North American passerine birds | journal = Ecology | volume = 47 | issue = 1| pages = 143–147 | doi=10.2307/1935753| jstor = 1935753 }}</ref>

====Polyandry threshold====
There also seems to be a "polyandry threshold" where males may do better by agreeing to share a female instead of maintaining a monogamous mating system.<ref>{{cite journal | last1 = Gowaty | first1 = P.A. | year = 1981 | title = An extension of the Orians-Verner-Willson model to account for mating systems besides polygyny | journal = American Naturalist | volume = 118 | issue = 6| pages = 851–859 | doi=10.1086/283875| s2cid = 83991131 }}</ref> Situations that may lead to cooperation among males include when food is scarce, and when there is intense competition for territories or females. For example, male [[lion]]s sometimes form coalitions to gain control of a pride of females. In some populations of [[Galapagos hawks]], groups of males would cooperate to defend one breeding territory. The males would share matings with the female and share paternity with the offspring.<ref>{{cite journal | last1 = Faaborg | first1 = J. | last2 = Parker | first2 = P.G. | last3 = DeLay | first3 = L. | year = 1995 | title = Confirmation of cooperative polyandry in the Galapagos hawk Buteo galapagoensis | journal = Behavioral Ecology and Sociobiology | volume = 36 | issue = 2| pages = 83–90 | doi=10.1007/bf00170712| s2cid = 3334592 |display-authors=etal}}</ref>

====Female desertion and sex role reversal====
In birds, desertion often happens when food is abundant, so the remaining partner is better able to raise the young unaided. Desertion also occurs if there is a great chance of a parent to gain another mate, which depends on environmental and populational factors.<ref>{{cite journal | last1 = Beissinger | first1 = S. R. | last2 = Snyder | first2 = N. F. R. | year = 1987 | title = Mate desertion in the snail kite | journal = Animal Behaviour | volume = 35 | issue = 2| pages = 477–487 | doi=10.1016/s0003-3472(87)80273-7| s2cid = 53192930 }}</ref> Some birds, such as the phalaropes, have reversed sex roles where the female is larger and more brightly colored, and compete for males to incubate their clutches.<ref>(Reynolds)</ref> In jacanas, the female is larger than the male and her territory could overlap the multiple territories of up to four males.<ref>{{cite journal | last1 = Butchart | first1 = S. H. M. | last2 = Seddon | first2 = N. | last3 = Ekstrom | first3 = J. M. M. | year = 1999b | title = Yelling for sex: harem males compete for female access in bronze-winged jacanas | journal = Animal Behaviour | volume = 57 | issue = 3| pages = 637–646 | doi=10.1006/anbe.1998.0985| pmid = 10196054 | s2cid = 24253395 }}</ref> In the frog species ''[[Bibron's toadlet|P. bibronii]],'' the female is fertilizes multiple nests, and the male is left to tend to each nest while the female moves on.