Societies

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Hierarchy

Social Structure

The social structures and dynamics seen within the spotted hyena species are extremely rare among large mammalian carnivores. They're most similar to the social structures seen in primate groups like babboons and macaques. These structures are characterized by having strict linear dominance patterns1) ). The higher the rank you are, the more access to resources you have . These rankings demonstrate how complex spotted hyena's social dynamics truly are. Spotted hyenas' social structures are also characterized by being matriarchal in which the females are more dominant to males <sup>1) </sup> . This is due to females evolving to be masculine. This evolution allowed for morphological and behavioral differences between male and female spotted hyenas2) . Examples of behavioral differences include females having control over mating and more access to food2) . Examples of morphological differences include the female development of a pseudopenis and females having larger body sizes. These differences are attributable to the matriarchial systems within spotted hyena clans. Another characteristic of hyena clans is that they are matrilineal2) . This indicates that lineages within the clans are tracable maternally. This is due to males dispersing from natal groups after reaching sexual maturity1) . Multiple matrilineal lineages are present within a hyena clan1) . Because of this, clans contain many genetically unrelated individuals

Rank Acquisition

Mother hyenas' ranks are inherited by their offspring 1) . This indicates that hyena mothers' fitnesses directly affect their offsprings' fitnesses. Rank inheritance is said to be a learning process in which young hyenas will either observe or partake in interactions with other caln members3) . Most of these interactions occur in intense feedings3) . Hyena feedings are very violent situations in which clan members try to eat as much food as they possibly can<sup>1) </sup> . Young hyenas will challenge clan members ranked below their mothers3) . The mother hyena may interneve in these challenges3) . By observing this intervention, young hyenas learn what their rank is compared to other clan members3) . Male hyenas will lose their rank when they disperse to a new clan1) .

Males entering new clans are considered to be the lowest ranking members1) . They'll need to behave submissively to all females as well as their cubs4) . If these males fail to do this, they'll be rejected from the clan4) . These submissive behaviors include retreat, ears back, tail between legs, and head bobbing7) . As with cubs, immigrant males learn of their rank through interactions with other group members1) . Males are said to disperse to new clans in order to breed with other females1) . This is because females would rather mate with immigrant males than males within their clan1) . Immigrant males will then contribute at least some parental care to their offspring4) .

Competition

Feeding Competition

Spotted hyenas are said to prey on ungulates including wildebeest, cape buffalo, gazelles, impalas, and many others1) . Usually, hyenas hunt alone1) . After a prey has been taken down, clan members will aggregate and compete to feed1) . As stated prior, mother hyenas will interfere if their young are competing with other clan members. In addition to this, mothers and their young form coalitions to compete with other clan members for access to the prey1) . These social ranks will determine other clan members; access to the food.

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Clan Competition

As a clan is feeding on its prey, neighboring clans may attempt to steal the prey. The clan will then form a coalition in order to defend their kill1) . Clans will also form coalitions in order to protect their territories. This occurs when a kill has been made near clan borders. Since females are ranked higher, they will lead coalitions1) . Hyenas have been observed to patrol their territory borders1) . Doing this, the hyenas will scent mark to reiterate to other clans where their territory boundaries are1) .

Interspecific Competition

As well as defending their kills from other clans, hyenas need to defend against lions. These two species are known to be intense competitors. This is due to them sharing similar diets as well as occupying the same habitats5) . Since lions are much larger than hyenas, they are usually successful in stealing food. This will occur if there aren't enough hyenas within the clan to defend their prey. Lions are also known to have a huge impact on hyena mortality5) . Hyenas deaths typically occur at communal dens where the lions are actively trying to hunt hyenas6) . Through this intense interspecific competition, hyenas have developed certain strategies to outcompete lions.

Fitness of Gregariousness

Prey Defense

Typically, animals aggregate together in order to either reduce predation or to enhance the exploitation of food resources. However, none of these explain spotted hyenas' gregarious behaviors. Hyenas aggregate together in order to better defend their food resources1) . As stated before, hyena clans face intense competition from other clans as well as from lions1) . Clans that have more individuals will be able to better defend their prey. This being said, natural selection will favor clans with higher densities of individuals1) . Hyenas not only defend their carcasses but they're also known to defend territories containing prey animals1) . This allows them to ensure that they'll have a reliable food supply for future usage1) .

Female Dominance

Masculinization

As stated before, female hyenas have evolved to be masculne compared to male hyenas. These masculine features include a large body and also an enlarged clitoris known as a pseudopenis. Several theories attempt to explain why this phenomenon occurred. These theories state that female masculinization occurred only as an accident. This isbecause elevated androgen levels within female hyenas were favored7) . One theory states elevated androgen levels were favored since larger, aggressive females were better able to compete with fellow clan members7) . These aggressive females had more access to food and mating opportunities compared to females who weren't as aggressive.

Another theory believes that wasn't a by-product of selection for increased androgen levels7) . They believe that the phenotype of having an enlarged clitoris was favored because of its usage in greetings7) . Female hyenas are known to erect their clitoris as a sign of submission to other females who are ranked higher hierarchically8) .

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Maternal Care

Within the spotted hyena species, most of the parental care care delived to offspring is maternal7) . This is because males are known to regularly migrate between clans. Maternal care is classified to be either maternal input or maternal effort. Maternal input is the energy and time that hyena mothers put into taking care of their offspring while maternal effort is the amount of care that's actually received by offspring7) . Maternal care issaid to fluctuate as the abundance of prey decreases or increases. Researchers have determined that maternal effort is low and maternal input is high during periods of high prey abundance7) . This is because mother hyenas don't need to spend as much time looking for food. As a result of this, mother hyenas are able to nurse their cubs more often7) . Researchers also determined that maternal effort is high while maternal care is low during periods of low prey abundance7) . This is because hyenas may travel out of their clan territory in order to find food7) . Because of this, cubs may go 2-3 days without being nursed7) .

1)
Watts, H and Hoelkamp, K (2007). Hyena Societies. Current Biology, 17(16). doi:10.1016/j.cub.2007.06.002 _FCKG_BLANK_TD_
2)
Hoelkamp, K & et al (2012). Society, demography, and genetic structure in the spotted hyena. Molecular Ecology, 12(21). 613-632. https://doi.org/10.1111/j.1365-294X.2011.05240.x
3)
Hoelkamp, K. & Smale. L. Dominance Acquisition During Mammalian Social Development: The Inheritance of Maternal Rank. American Zoologist, 31(2). 306-317, https://doi.org/10.1093/icb/31.2.306
4)
Hoelkamp, K & Smale, L (1998). Behavioral Development in the Spotted Hyena. BioScience, 48(12). (997-1005). https://doi.org/10.2307/1313456
5)
Hoelkamp, K & Watts, H (2009). Ecological Determinants of Survival and Reproduction in the Spotted Hyena. Journal of Mammalogy, 90((Watts, H.E. and Holekamp, K.E. (2008), Interspecific competition influences reproduction in spotted hyenas. Journal of Zoology, 276: 402-410. https://doi.org/10.1111/j.1469-7998.2008.00506.x
6)
Watts, H.E. and Holekamp, K.E. (2008), Interspecific competition influences reproduction in spotted hyenas. Journal of Zoology, 276: 402-410. https://doi.org/10.1111/j.1469-7998.2008.00506.x
7)
East, M & Hofer, H. Conflict and Cooperation in a Female-Dominated Society: A Reassessment of the “Hyperaggressive” Image of Spotted Hyenas. Advances in the Study of Behavior, Vol 31. 1-30. https://doi.org/10.1016/S0065-3454 (02)80004-3
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