Mothers front-load their investment to the egg stage when helped in a wild cooperative bird

In many cooperative societies, including our own, helpers assist with the post-natal care of breeders' young, and may thereby benefit the post-natal development of offspring. Here we present evidence of a novel mechanism by which such post-natal helping could also have hitherto unexplored beneficial effects on pre-natal development: by lightening post-natal maternal workloads, helpers may allow mothers to increase their pre-natal investment per offspring. We present the findings of a decade-long study of cooperatively breeding white-browed sparrow weaver, Plocepasser mahali, societies. Within each social group, reproduction is monopolized by a dominant breeding pair, and non-breeding helpers assist with nestling feeding. Using a within-mother reaction norm approach to formally identify maternal plasticity, we demonstrate that when mothers have more female helpers they decrease their own post-natal investment per offspring (feed their nestlings at lower rates) but increase their pre-natal investment per offspring (lay larger eggs, which yield heavier hatchlings). That these plastic maternal responses are predicted by female helper number, and not male helper number, implicates the availability of post-natal helping per se as the likely driver (rather than correlated effects of group size), because female helpers feed nestlings at substantially higher rates than males. We term this novel maternal strategy 'maternal front-loading' and hypothesize that the expected availability of post-natal help allows helped mothers to focus maternal investment on the pre-natal phase, to which helpers cannot contribute directly. Such cryptic maternally mediated helper effects on pre-natal development may markedly complicate attempts to identify and quantify the fitness consequences of helping.

What kind of maternal effects are selected for in fluctuating environments?

Adaptation to temporally fluctuating environments can be achieved through direct phenotypic evolution, by phenotypic plasticity (either developmental plasticity or trans-generational plasticity), or by randomizing offspring phenotypes (often called diversifying bet-hedging). Theory has long held that plasticity can evolve when information about the future environment is reliable while bet-hedging can evolve when mixtures of phenotypes have high average fitness (leading to low among generation variance in fitness). To date, no study has studied the evolutionary routes that lead to the evolution of randomized offspring phenotypes on the one hand or deterministic maternal effects on the other. We develop simple, yet general, models of the evolution of maternal effects and are able to directly compare selection for deterministic and randomizing maternal effects and can also incorporate the notion of differential maternal costs of producing offspring with alternative phenotypes. We find that only a small set of parameters allow bet hedging type strategies to outcompete deterministic maternal effects. Not only must there be little or no informative cues available, but also the frequency with which different environments are present must fall within a narrow range. By contrast, when we consider the joint evolution of the maternal strategy and the set of offspring phenotypes we find that deterministic maternal effects can always invade the ancestral state (lacking any form of maternal effect). The long-term ESS may, however, involve some form of offspring randomization, but only if the phenotypes evolve extreme differences in environment-specific fitness. Overall we conclude that deterministic maternal effects are much more likely to evolve than offspring randomization, and offspring randomization will only be maintained if it results in extreme differences in environment-specific fitness.

Ontogeny of swimming and diving in northern fur seal (Callorhinus ursinus) pups

Time spent in the water and diving behavior of northern fur seal (Callorhinus ursinus) pups were monitored between birth and weaning at St. Paul Island, Alaska. The median age when pups began swimming was 26 days, but prior to 40 days of age they spent virtually all their time on shore and only rarely took brief (15-20 min) swims. Pups began spending substantial time in the water at approximately 40-50 days of age, coinciding both with the early growth of insulating underfur and a seasonal peak in sea surface temperature. This suggests that pups had earlier been constrained to remain on shore by their undeveloped thermoregulatory capabilities. Time in the water increased up to approximately 100 days of age, when molted pups spent about 35% of their time in the water and swim bouts were several hours in duration. Moreover, the presence of a pup's mother on shore, photoperiod, and precipitation also influenced the amount of time pups spent in the water. Pups (mean age = 100 days) dove to very shallow depths (mean = 3 m) for short durations (mean = 11 s). Because pups did not gain mass unless suckled by their mothers, it is unlikely that they fed extensively while diving prior to migration. The pattern of development of swimming and diving in northern fur seals is intermediate between typical phocid and otariid patterns, as is the maternal strategy.