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We use a time-varying statistical framework that accounts for age-related changes in individual social connectedness and in the availability of social partners. Our results confirm the importance of female—female social connectedness to female longevity in this species [ 12 ] and extend this result to encompass a further independent effect of male—female connectedness on female survival. The Woman adult date in mikumi of these effects motivated us to also investigate the social and demographic factors that predict variation in female social connectedness, a topic of avult interest in the literature on humans.

Material and methods a Study subjects Study datte were members of a well-studied population of wild baboons living in the Amboseli ecosystem in southern Kenya [ 53 ]. Behavioural and demographic data on each group were collected by three experienced observers during 5-h monitoring visits. These visits occurred year round, two to three times per week per group. Death dates were known to be within a few days for females that died before Womzn end of the study period. For females, avult was defined by the onset of menarche; males were considered adult when they became higher ranking than all adult Woman adult date in mikumi and ranked among the mkumi males in their adulg group [ 54 ].

Arult studies of the adaptive and health significance of social behaviour, affiliative social connectedness has been measured in a variety of ways e. Our indices were very similar to those used in several prior studies in non-human animals [ 141653 ], as well as those used to measure structural social integration in many studies in humans reviewed in [ 1 ]. Specifically, social connectedness was measured for each female relative to all other adult females alive in the population in the same year. Following previous studies [ 12141554 ], these indices used data on grooming behaviour, which maintains and strengthens social bonds in baboons and other primates [ 5556 ]. To measure grooming relationships, we chose to use ad libitum observations of grooming [ 5758 ], which included all observed instances of grooming between group members and was the densest dataset available to measure patterns of female affiliation see the electronic supplementary material.

Our sampling protocol was designed to avoid potential biases in the grooming data that could result from uneven sampling of study subjects. Specifically, the great majority of our ad libitum data were collected during random-order focal animal sampling on adult females and juveniles, which ensured that observers continually moved to new locations within the group and observed all adult females and juveniles on a regular rotating basis. Ad libitum grooming frequencies were significantly correlated with hourly rates of grooming from focal animal sampling see the electronic supplementary materialindicating a lack of strong or systematic bias in the ad libitum data.

Nevertheless, we could not assess whether our analysis choices completely eliminated biases introduced by our sampling protocol; therefore, we also consider possible implications of these choices in the Discussion. From the ad libitum data, we calculated SCI-F and SCI-M for each adult female in each year of her adult life as a composite index of the relative frequency that she groomed and was groomed by adult females or adult males, respectively see the electronic supplementary material, figure S1. Positive SCI values represent females with relatively high frequencies of grooming for the population in that year; negative values represent females with relatively low frequencies of grooming for that year.

We modelled survival in adult females using Cox proportional hazards models. We employed time-varying covariates in our models because, in the course of testing predictors of SCI-F, we found that older females generally had lower values of SCI-F, making it inappropriate to use a single, average value of lifetime social connectedness. We ran two different models using the rms package [ 5960 ] in R [ 61 ]. Missing values were imputed via multiple imputation [ 62 ] and weighted predictive mean matching as implemented via the aregImpute function in the rms package in R [ 5960 ] see the electronic supplementary material for additional information on data imputation methods.

We performed the full imputation 50 times to create 50 imputed datasets and fit the main Cox proportional hazards model to each of these 50 datasets.

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Parameters presented in the Womzn model were averaged over the 50 model fits. This model on female-years of data on females, with censored records. For both the main and the complete case models, females entered the model at adulthood and left the model at death or censorship. For both models, we included the following predictor variables: For each of these predictors, the validity of the proportional hazards assumption was well supported electronic supplementary material, table S1.

Mikkumi were no differences in the results from the main and complete case models; Womwn the text, we present the results of the main model because of its added statistical power see the electronic supplementary material for results of the complete case model. Based on prior research, we expected that female social connectedness to adult females would be correlated with the availability of adult maternal kin, who often form the strongest social bonds in baboon societies [ 63 — 66 ], as well as age, which is associated with declining availability of non-kin social partners [ 64 ]. Elephant mating rituals include the gentle entwining of trunks.

Elephant cognition Scratching on a tree helps to remove layers of dead skin and parasites African elephants are highly intelligent, [29] and they have a very large and highly convoluted neocortexa trait they share with humansapes and some dolphin species.

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They are amongst the world's most intelligent species. The Wokan brain is similar to that of humans in terms of structure and complexity. For example, the elephant's cortex has as many neurons as that of a human brain[30] suggesting convergent evolution. African elephants show sexual dimorphism in weight and shoulder height by age 20, due to the rapid early growth of males. By age 25, males are double the weight of females; however, both sexes continue to grow throughout their lives. Female African elephants Wojan able to start reproducing at around 10 to miiumi years of age, [39] and are in estrus for about 2 to 7 days.

They do not mate at a specific time; however, they are less likely to reproduce in times of drought than when water is plentiful. The gestation period of an elephant is 22 months and fertile females usually give birth every 3 — 6 years, so if they live to around 50 years of age, they may produce 7 offspring. Females are a scarce and mobile resource for the males so there is intense competition to gain access to estrous females. Post sexual maturity, males begin to experience musth, a physical and behavioral condition that is characterized by elevated testosterone, aggression and more sexual activity.

Males sire few offspring in periods when they are not in musth. During the middle of estrus, female elephants look for males in musth to guard them. The females will yell, in a loud, low way to attract males from far away. Male elephants can also smell the hormones of a female ready for breeding. This leads males to compete with each other to mate, which results in the females mating with older, healthier males.

However, females are not guarded in the early and late stages of estrus, which may datw mating by younger males not in musth. Males over the age of 25 compete strongly for females in estrous, and are oWman successful the larger and more dte they are. Male reproductive success is maximal in mid-adulthood and then begins to decline. However, this can depend on the ranking of the male within their group, as higher-ranking males maintain a higher rate of reproduction. Twenty-two long observations showed that age and musth are extremely important factors; "… older males had markedly elevated paternity success compared with younger males, suggesting the possibility of sexual selection for longevity in this species.

Males usually stay with a female and her herd for about a month before moving on in search for another mate.


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