In polytocous mammals, weakling neonates, but not their stronger littermates, benefit from specialized foraging.

Adjusting foraging strategies is a common phenomenon within groups of animals competing for the same resource. In polytocous mammals, neonates concurrently compete for limited milk and alternate between two foraging (suckling) strategies: adaptable exploratory foraging with random sampling of teats, and ordered foraging with a tendency towards exploiting a particular suckling position. Some theoretical (game theory) models have shown that weaker siblings in particular benefit from foraging specialization (suckling order). Neonate piglets establish a well-defined suckling order that develops gradually and fluctuates throughout the lactation period, implying the existence of inter-individual differences in foraging strategies. We therefore analyzed suckling behavior in pigs to determine whether one foraging strategy was more beneficial to neonates in terms of their body weight and foraging environment. We found that intermediate and heavy littermates tended to adjust their suckling strategy according to the foraging environment; however, the selected foraging strategy did not affect their overall growth performance. Lighter individuals that consumed significantly less milk did not greatly alternate their foraging strategy according to the foraging environment, but their growth rate was significantly higher whenever they performed less-exploratory foraging behavior. Although suckling order appeared to be a relatively stable behavioral phenotype, it was beneficial exclusively for weaklings. These results confirm theoretical predictions and indicate that specializing in a suckling position is a beneficial strategy for weaker, light neonates. These findings suggest that physically weaker neonates might have driven the evolution of neonatal foraging specialization.

On the presence and absence of suckling order in polytocous mammals.

Mammals have developed a variety of suckling behaviours ranging from tenacious nipple attachment in some rodents and marsupials to once-a-day suckling in rabbit. However, a common feature of suckling that was found in many mammals is the suckling order, or a partial preference to suckle a particular teat (teat fidelity) or part of the udder (suckling preference). A lack of suckling order is observed only in a few mammals. In this article, the possible background of the presence or absence of suckling order in eutherian polytocous mammals is discussed either from the maternal investment or sibling competition point of view. Characteristics related to maternal investment in species in which the suckling order has already been studied at least partially, were classified using C4.5 algorithm (J48 classifier in Weka 3.8.1), and decision tree was built. In the context of sibling competition, an extensive form game (game theory) was predicted to show the optimal suckling strategy considering the basic relations among littermates in two situations (littermates of equal strength/dominance and littermates with different strength/dominance). Although no ultimate conclusion can be drawn, it appears that the suckling order is typical for species whose reproductive system requires a lower maternal investment (up to one litter/year, monogamy, biparental care, lower litter birth weight); and, it appears that the suckling order is inherent to the weaker (inferior) siblings.

Being a weakling and surviving: Keep the fittest siblings close-by when eating.

Weaklings serve as a buffer, allowing an increase in the overall prospect of offspring survival. Whereas birds are able to directly determine which offspring to invest in via selective feeding, multiparous mammals have fewer methods of excluding weaklings from milk provision. In pigs the maternal investment strategy is based on the vigorousness of the progeny, whereby weaklings can suffer hunger due to an inability to sufficiently stimulate the mammary gland. However, the vigorous massage might stimulate higher milk production in the surrounding glands and it is possible that a weakling's survivability is conditioned also by suckling at a position adjacent to heaviest littermate. Results of the present study have confirmed this prediction, showing that the surviving weaklings tended to have the heaviest littermates as nearest neighbours during suckling. It appears, therefore, that weaklings which establish a type of commensal, rather than competitive, relationship with strongest siblings might increase their survivability.