Begging and hatch order can outweigh kinship
In smooth-billed anis, a communally breeding bird where several non-kin females lay in one shared nest, adults did not preferentially feed their own offspring and did not increase feeding as their genetic share of the brood rose [4]. Instead, nestlings that begged earlier and harder, and those that hatched first, were fed more. The researchers conclude that these birds attend to signals and cues of nestling condition and need rather than to kinship [4].
There was one nuance: adults were more likely to feed their own offspring in nests where they had a higher proportion of the chicks, which the authors suggest may reflect more productive individuals indirectly benefiting their offspring by chance rather than active kin recognition [4]. That distinction matters. A statistical association between relatedness and feeding can appear even when birds are not using relatedness as a cue.
Offspring sex and family competition can redirect care
In banded mongooses, patterns of alloparental care after pups emerge from the den are explained by offspring sex rather than by relatedness [2]. This is a concrete case where knowing how related a helper is to a pup would lead you to the wrong prediction about how much care that pup receives.
Kin competition offers another route by which relatedness stops being a good guide to help. A model of cooperative breeding in family groups shows that when relatives compete for limited breeding openings in a saturated habitat, parents can benefit from tolerating helpers even if those helpers reduce parental survival or fecundity [5]. Helping is also favored when staying home lowers the helper's dispersal costs or raises its chance of inheriting the territory. In such settings, the division of reproduction can emerge for reasons unrelated to the effects of help itself [5]. A helper's presence, in other words, may reflect the economics of waiting for a territory rather than a measured contribution to the brood.
Sources used in this answer
Load lightening and negotiation over offspring care in cooperative breeders
A negotiation model of cooperative breeders finds that relatedness alone does not determine helping effort, and that a helper's relatedness to the parent can matter more than relatedness to the offspring.
Biased escorts: Offspring sex, not relatedness explains alloparental care patterns in a cooperative breeder.
In banded mongooses, patterns of alloparental escort care are explained by offspring sex rather than by relatedness.
Helpers compensate for age-related declines in parental care and offspring survival in a cooperatively breeding bird
In Seychelles warblers, helpers mitigate age-related declines in breeder provisioning and offspring survival mainly because older females have recruited more helpers, not because individual helpers increase their care.
Offspring provisioning is affected by begging and hatch order but not relatedness in a communally breeding bird
Smooth-billed anis did not preferentially feed their own offspring or provision based on their genetic contribution to communal broods; instead they fed nestlings that begged earlier and harder and that hatched first.
The evolution of cooperative breeding in family groups: when should parents tolerate unhelpful helpers?
Kin competition in saturated habitats can favor parents tolerating helpers even when helpers reduce parental survival or fecundity, so division of reproduction can emerge for reasons unrelated to the effects of help.
