Theory suggests females should optimize resource allocation across reproductive bouts to maximize lifetime reproduction, balancing current and future reproductive efforts according to physiological state and projected survival and reproduction. Tests of these ideas focus on long-lived vertebrates: few measure age-related reproductive output in iteroparous invertebrates, or partition reserves between those allocated to offspring versus mothers. We investigated how maternal age, and environmental and physiological factors influence reproductive investment in wild tsetse, Glossina pallidipes Austen and G. morsitans morsitans Westwood. Tsetse provide a tractable system to measure reproductive allocation. Females exhibit high maternal investment, producing single, large offspring that rely exclusively on maternal reserves. We find that mothers in better physiological condition and experiencing cooler temperatures produce larger offspring. Pupal size increases significantly but weakly with age. In both species, females with less fat invest proportionately more in offspring. Post-partum fat decreases in flies with badly frayed wings: poor flight capability may limit their feeding efficiency, or they may sacrifice more reserves as a terminal investment. Our results support evidence that offspring size increases with maternal size, investment depends on the environment, and females with lower chances of future reproduction invest more into current offspring. We discuss the implications of maternal effects for predicting vector population responses to environmental change.

1. Resource availability plays a key role in driving variation in somatic growth and body condition, and the factors determining access to resources vary considerably across life stages. Parents and carers may exert important influences in early life, when individuals are nutritionally dependent, with abiotic environmental effects having stronger influences later in development as individuals forage independently. 2. Most studies have measured specific factors influencing growth across development, or have compared relative influences of different factors within specific life stages. Such studies may not capture whether early-life factors continue have delayed effects at later stages, or if social factors change when individuals become nutritionally independent and adults become competitors for, rather than providers of, food. 3. Here, we examined variation in the influence of the abiotic, social and maternal environment on growth across life stages in a wild population of cooperatively breeding meerkats. Cooperatively breeding vertebrates are ideal for investigating environmental influences on growth. In addition to experiencing highly variable abiotic conditions, cooperative breeders are typified by heterogeneity both among breeders, with mothers varying in age and social status, and in the number of carers present. 4. Recent rainfall had a consistently marked effect on growth across life stages, yet other seasonal terms only influenced growth during stages when individuals were growing fastest. Group size and maternal dominance status had positive effects on growth during the period of nutritional dependence on carers, yet did not influence mass at emergence (at one month) or growth at independent stages (>4 months). Pups born to older mothers were lighter at one month of age, and subsequently grew faster as subadults. Males grew faster than females during the juvenile and subadult stage only. 5. Our findings demonstrate the complex ways in which the external environment influences development in a cooperative mammal. Individuals are most sensitive to social and maternal factors during the period of nutritional dependence on carers, whereas direct environmental effects are relatively more important later in development. Understanding the way in which environmental sensitivity varies across life stages is likely to be an important consideration in predicting trait responses to environmental change.

Mating between relatives often results in negative fitness consequences or inbreeding depression. However, the expression of inbreeding in populations of wild cooperative mammals and the effects of environmental, maternal and social factors on inbreeding depression in these systems are currently not well understood. This study uses pedigree-based inbreeding coefficients from a long-term study of meerkats (Suricata suricatta) in South Africa to reveal that 44% of the population have detectably non-zero (F>0) inbreeding coefficients. 15% of these inbred individuals were the result of moderate inbreeding (F≥0.125), although such inbreeding events almost solely occurred when mating individuals had no prior experience of each other. Inbreeding depression was evident for a range of traits: pup mass at emergence from the natal burrow, hind-foot length, growth until independence and juvenile survival. However, we found no evidence of significant inbreeding depression for skull and forearm length or for pup survival. This research provides a rare investigation into inbreeding in a cooperative mammal, revealing high levels of inbreeding, considerable negative consequences and complex interactions with the social environment.