Life‐history trade‐off theory predicts that current reproduction can negatively affect survival and future reproduction. Few studies have assessed breeding costs for males of polygynous species compared to females, despite substantial variation in breeding success among individual males (e.g. subordinate cf. dominant breeders). Specifically, differentiating between the cost of attending breeding events, and the additional cost of successfully securing and mating females is lacking. We investigated whether trade‐offs are present in the highly polygynous male southern elephant seal Mirounga leonina using 34 years of individual‐level data. We compare age‐specific survival, recruitment and future breeding success probabilities of pre‐breeders (males yet to recruit) and breeders (subordinate and dominant social ranks) using multievent models. Pre‐breeders and breeders of overlapping ages had similar survival probabilities, suggesting that there was no attendance cost for early recruits. In addition, the probability of recruiting as a dominant breeder never exceeded recruitment probability as a subordinate breeder of the same age. Therefore, older pre‐breeders that delayed attendance costs generally did not improve their breeding success (probability of being dominant) at recruitment more than younger recruits. Rather, recruitment age may be a function of individual quality, with lower quality individuals requiring more time to socially mature. When comparing subordinate and dominant breeders, we found clear evidence for survival senescence, with subordinate breeders having a higher baseline mortality. In contrast, age‐specific future breeding success (probability of being dominant at t + 1) increased with age, with dominant breeders maintaining higher subsequent breeding success than subordinate breeders. The opposite trends in survival and future breeding success for both subordinate and dominant breeders may indicate a lifetime, population‐level trade‐off. However, we found no evidence to suggest that being a dominant breeder consecutively (and having a higher accumulated breeding cost) accelerated the rate of senescence when compared to individuals that were previously subordinate. Thus, males experienced actuarial senescence regardless of social rank, with dominant (and possibly high quality) breeders showing a reduced trade‐off between survival and future breeding success. We make several novel contributions to understanding polygynous male life histories and southern elephant seal demography.
Individual heterogeneity is variation in trait expression observed among individuals of a population. Individual heterogeneity in the pre-breeder stage of development is of importance given its eventual contribution to the breeding population's overall reproductive performance. Yet most studies do not consider the role of individual heterogeneity in pre-breeders when investigating population processes. We investigated individual heterogeneity in the survival and recruitment probabilities of pre-breeding male southern elephant seals Mirounga leonina using 34 years of data collected at Marion Island. Elephant seals are highly polygynous capital breeders, with few male offspring surviving to compete in breeding events and even fewer breeding successfully. Specifically, we fitted finite-mixture models with two hidden groups that represented 'robust' and 'frail' individuals and compared the demographic rates of these groups to that of the population. Survival and recruitment as first-time subordinate breeders could be distinguished between groups. Survival of both robust and frail pre-breeders decreased from age 2, whilst differences between groups were maintained (ontogeny processes). Frail pre-breeders had a much lower survival than the population average, suggesting that frail individuals were preferentially removed from the population (selective disappearance). Differences in recruitment probabilities were apparent from age 5 (earliest recorded age) and increased until age 7, with most recruits comprising robust pre-breeders. Male pre-breeders were more likely to be robust at age 2 when born in years with few conspecifics, suggesting that individual heterogeneity was determined, in part, by density-dependent effects. A population projection model revealed that male breeders of every age class never consisted of more than 5% of frail recruits. Rather, the breeding population increasingly consisted of robust recruits that obtained social dominance with age. We demonstrated that individual heterogeneity in male pre-breeder demographic rates was present and persistent throughout much of development and determined the proportion of adult males that obtained social dominance.
The objective of the study was to record the tick species collected from three species of tortoise, each in a different province of South Africa. Ticks were collected from leopard tortoises, Stigmochyles pardalis, in the southern region of the Kruger National Park, Mpumalanga province; from hingeback tortoises, Kinixys zombensis, in the Enseleni Nature Reserve, KwaZulu-Natal province and from angulate tortoises, Chersina angulata, in the West Coast National Park, Western Cape province. Of the 63 leopard tortoises examined, 58 were infested with Amblyomma marmoreum and 49 with Amblyomma hebraeum, and all stages of development of both species were recovered. Amblyomma nuttalli was collected from 25 hingeback tortoises, and all stages of development were present. All 24 angulate tortoises examined were infested with Amblyomma sylvaticum, and large numbers of larvae, nymphs and adults were collected. Three snake species and a sand lizard were also infested with A. sylvaticum. The adults of A. marmoreum, A. nuttalli and A. sylvaticum were identified as specific parasites of the family Testudinidae, whereas all stages of development of A. hebraeum were classified as generalists.
Trophic generalists tolerate greater habitat change than specialists; however, few studies explore how generalist trophic ecology is affected. We established how the trophic ecology of an extreme generalist, Rhabdomys pumilio, changed in relation to a directionally changing woody‐encroached savannah in Eastern Cape, South Africa by investigating (a) foraging behaviour, (b) trophic niche and (c) feedback effects. (a) Giving‐up densities showed that R. pumilio preferred foraging in subcanopy microhabitat during the night as a result of lower thermoregulatory costs, but had similar preferences for sub‐ and intercanopy microhabitats during the day. (b) An isotope analysis revealed that the dietary composition and trophic niche occupied by R. pumilio differed among tree canopy cover levels (0%, 30% and 80%), which appeared to be related to changes in C4 grass material and invertebrate availability. (c) Artificial seed patches suggested that R. pumilio was a potentially important postdispersal seed predator of the woody‐encroaching species, Vachellia karroo. Thus, an increase in tree canopy cover altered the trophic niche of R. pumilio by reducing foraging costs at night and providing alternative food resources in terms of availability and source. These findings demonstrate how an extreme generalist adapted to human‐induced habitat change through changes in its trophic ecology.
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