Long-lived species such as elephants, whales and primates exhibit extended post-fertile survival compared to species with shorter lifespans but data on age-related fecundity and survival are limited to few species or populations. We assess relationships between longevity, reproductive onset, reproductive rate and age for 834 longitudinally monitored wild female African elephants in Amboseli, Kenya. The mean known age at first reproduction was 13.8 years; only 5 % commenced reproduction by 10 years. Early reproducers (<12.5 years) had higher age-specific fertility rates than did females who commenced reproduction late (15+ years) with no differences in survival between these groups. Age-specific reproductive rates of females dying before 40 years were reduced by comparison to same-aged survivors, illustrating a mortality filter and reproductive advantages of a long life. Overall, 95 % of fertility was completed before 50, and 95 % of mortality experienced by age 65, with a mean life expectancy of 41 years for females who survived to the minimum age at first birth (9 years). Elephant females have a relatively long period (c. 16 years) of viability after 95 % completed fertility, although reproduction does not entirely cease until they are over 65. We found no evidence of increased investment among females aged over 40 in terms of delay to next birth or calf mortality. The presence of a mother reproducing simultaneously with her daughter was associated with higher rates of daughter reproduction suggesting advantages from maternal (and grandmaternal) co-residence during reproduction.
Growth from conception to reproductive onset in African elephants (Loxodonta africana) provides insights into phenotypic plasticity, individual adaptive plastic responses and facultative maternal investment. Using growth for 867 and life histories for 2652 elephants over 40 years, we demonstrate that maternal inexperience plus drought in early life result in reduced growth rates for sons and higher mortality for both sexes. Slow growth during early lactation was associated with smaller adult size, later age at first reproduction, reduced lifetime survival and consequently limited reproductive output. These enduring effects of trading slow early growth against immediate survival were apparent over the very long term; delayed downstream consequences were unexpected for a species with a maximum longevity of 70þ years and unpredictable environmental experiences.
Detailed demographic data on wild Asian elephants have been difficult to collect due to habitat characteristics of much of the species’ remaining range. Such data, however, are critical for understanding and modeling population processes in this endangered species. We present data from six years of an ongoing study of Asian elephants (Elephas maximus) in Uda Walawe National Park, Sri Lanka. This relatively undisturbed population numbering over one thousand elephants is individually monitored, providing cohort-based information on mortality and reproduction. Reproduction was seasonal, such that most births occurred during the long inter-monsoon dry season and peaked in May. During the study, the average age at first reproduction was 13.4 years and the 50th percentile inter-birth interval was approximately 6 years. Birth sex ratios did not deviate significantly from parity. Fecundity was relatively stable throughout the observed reproductive life of an individual (ages 11–60), averaging between 0.13–0.17 female offspring per individual per year. Mortalities and injuries based on carcasses and disappearances showed that males were significantly more likely than females to be killed or injured through anthropogenic activity. Overall, however, most observed injuries did not appear to be fatal. This population exhibits higher fecundity and density relative to published estimates on other Asian elephant populations, possibly enhanced by present range constriction. Understanding the factors responsible for these demographic dynamics can shed insight on the future needs of this elephant population, with probable parallels to other populations in similar settings.
i z a b e t h W e b b e r , T u y S e r e i v a t h a n a , M a t t h e w P . M a l t b y and P h y l l i s C . L e e Abstract Elephants are threatened globally by habitat loss, poaching and accelerating levels of human-elephant conflict. For Elephas maximus in Cambodia, crop raiding underlies this conflict. Understanding the timing of raids and selection of crops can help design locally appropriate mitigation and management strategies. This study, using a 4-year database of events, investigated the most frequently raided crops and patterns of raids, over time and seasons and by location. Damage frequency varied significantly by crop, with rice, banana, cassava, sugar cane and papaya most frequently raided. Considering raid events per unit crop area, banana, sugar cane and pineapple were raided more than would be expected based on their availability. There were differences in both crop-raiding events and crop-damage frequencies over study years and there was a peak raiding season in October-December. Nationally, significant differences were found among provinces but not between years. Rates of damage decreased after mitigation strategies such as observation towers, deterrents and fences were implemented. We suggest further mechanisms to improve human-elephant conflict monitoring in relation to crop choice and availability.
We explore elephant play behaviour since (a) play has been proposed to represent a potential welfare indicator; and (b) play has been associated with long-term survival in the wild. We categorised play into four types, and investigate both social (gentle, escalated-contact) and non-social (lone-locomotor, exploratory-object) play from observations made on wild (Asian N = 101; African N = 130) and captive (Asian N = 8; African N = 7) elephant calves ranging in age from birth to five years. Social play was the most frequent type of play among immature elephants, accounting for an average of 3%–9% of active time. Non-social play accounted for an additional 1%–11% of time. The most time spent in play was seen in captive Asian calves, particularly at the ages of 1–6 months, while wild African calves spent the least time in play overall, even though they had the greatest number and most diverse range of play partners available. We assessed calf energetics using time spent suckling, resting, moving and independent feeding. Time spent playing was unrelated to time spent suckling but negatively associated with time spent independently feeding. There were no associations with time spent moving or resting. Maternal energy via lactation was unrelated to play early in life, but energy acquired independently may constrain or enable play. Play, while a potential indicator of compromised welfare for many species when absent, can act as a highly stimulating activity for captive elephants in the absence of other forms of arousal.
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