Phylogenetic reconstruction of parental-care systems in the ancestors of birds. (9/195)

Due to the controversy surrounding incipient avian parental care, ancestral parental care systems were reconstructed in a phylogeny including major extant amniote lineages. Using two different resolutions for the basal avian branches, transitions between the states no care, female care, biparental care and male care were inferred for the most basal branches of the tree. Uniparental female care was inferred for the lineage to birds and crocodiles. Using a phylogeny where ratites and tinamous branch off early and an ordered character-state assumption, a transition to biparental care was inferred for the ancestor of birds. This ancestor could be any organism along the lineage leading from the crocodile-bird split up to modern birds, not necessarily the original bird. We discuss the support for alternative avian phylogenies and the homology in parental care between crocodiles and birds. We suggest that the phylogenetic pattern should be used as a starting point for a more detailed analysis of parental care systems in birds and their relatives.  (+info)

Evolutionary transitions in parental care and live bearing in vertebrates. (10/195)

We provide the first review of phylogenetic transitions in parental care and live bearing for a wide variety of vertebrates. This includes new analyses of both numbers of transitions and transition probabilities. These reveal numerous transitions by shorebirds and anurans toward uniparental care by either sex. Whereas most or all of the shorebird transitions were from biparental care, nearly all of the anuran transitions have been from no care, reflecting the prevalence of each form of care in basal lineages in each group. Teleost (bony) fishes are similar to anurans in displaying numerous transitions toward uniparental contributions by each sex. Whereas cichlid fishes have often evolved from biparental care to female care, other teleosts have usually switched from no care to male care. Taxa that have evolved exclusive male care without courtship-role reversal are characterized by male territoriality and low costs of care per brood. Males may therefore benefit from care through female preference of parental ability in these species. Primates show a high frequency of transitions from female care to biparental care, reflecting the prevalence of female care in basal lineages. In the numerous taxa that display live bearing by females, including teleosts, elasmobranchs, squamate reptiles and invertebrates, we find that live bearing has always evolved from a lack of care. Although the transition counts and probabilities will undoubtedly be refined as phylogenetic information and methodologies improve, the overall biases in these taxa should help to place adaptive hypotheses for the evolution of care into a stronger setting for understanding directions of change.  (+info)

Male-only care and classical polyandry in birds: phylogeny, ecology and sex differences in remating opportunities. (11/195)

It has been argued recently that the combination of male-only parental care and classical polyandry in birds is the most interesting and yet the least understood of all avian breeding systems. Despite a huge number of hypotheses, careful comparative analyses have repeatedly failed to identify consistent ecological differences between species showing male-only care and closely related species showing other patterns of care. This has led to the suggestion that such analyses fail because the crucial differences are between ancient lineages rather than between closely related species. Here, therefore, I use comparisons between families to test three well-known hypotheses: that male-only care is associated with: (i) a low rate of fecundity; (ii) large egg size relative to female size; or (iii) female-biased opportunities for remating. Families showing male-only care do not differ from families showing female-only care with respect to rate of fecundity or relative egg size. There is, however, a significant difference between these two groups of families with respect to an index of remating opportunities, nesting density. Families showing female-only care nest at high density, while those showing male-only care nest at very low density. This is one of the first times a consistent ecological correlate has been identified for male-only care in birds. It suggests that female-only care arises (or persists) in families where remating opportunities are abundant for both sexes, whereas male-only care arises (or persists) in families where remating opportunities are rare for both sexes and particularly scarce for males. This in turn suggests that sex differences in remating opportunities are the key ecological factor in determining male-only care and classical polyandry in birds.  (+info)

Intrafamilial conflict and parental investment: a synthesis. (12/195)

We outline and develop current theory on how inherent genetic conflicts of interest between the various family members can affect the flow of parental investment from parents to offspring, and discuss the problems for empirical testing that this generates. The parental investment pattern realized in nature reflects the simultaneous resolution of all the conflicts between the family players. This depends on the genetic mechanism, the mating system and reproductive constraints, on whether extra demand by progeny affects current or future sibs, and particularly on the behavioural mechanisms underlying demand (begging or solicitation) and supply (provision of parental investment by parents). The direction of deviation from the optimal parental investment for the parent(s) depends on the slope of what we term the 'effect of supply on demand', the mechanism that determines how changes in food supply affect begging levels. If increasing food increases begging (positive slope), less parental investment is supplied than the parental optimum and if increasing food decreases begging (negative slope), more parental investment is supplied. The magnitude of deviation depends on both the 'effect of supply on demand' and on the 'effect of demand on supply' (the mechanism determining how changes in begging affect food supply, which always has a positive slope). We conclude that it will often be impossible to deduce the extent of underlying conflict by establishing the amount of parental investment given relative to the ideal optimum for the parent. Some possible directions for future research are discussed.  (+info)

A model of the interaction between 'good genes' and direct benefits in courtship-feeding animals: when do males of high genetic quality invest less? (13/195)

Conflict between mates over the amount of parental investment by each partner is probably the rule except in rare cases of genetic monogamy. In systems with parental care, males may frequently benefit by providing smaller investments than are optimal for individual female partners. Females are therefore expected to choose males that will provide the largest amounts of parental investment. In some species, however, the preferred males provide less care than their rivals. Focusing on species in which males invest by feeding their mates, I use a simple model to demonstrate the conditions under which males preferred by females may have optimal donations that are smaller than those of less-preferred rivals. Pre-mating female choice may sufficiently bias the perception of mate availability of preferred males relative to their rivals such that preferred males gain by conserving resources for future matings. Similarly, 'cryptic' biases in favour of high-quality ejaculates by females can compensate for smaller than average donations received from preferred males. However, post-fertilization cryptic choice should not change the optimal donations of preferred males relative to their rivals. I discuss the implications of this work for understanding sexual selection in courtship-feeding animals, and the relevance of these systems to understanding patterns of investment for animals in general.  (+info)

Why is mutual mate choice not the norm? Operational sex ratios, sex roles and the evolution of sexually dimorphic and monomorphic signalling. (14/195)

Biases in the operational sex ratio (OSR) are seen as the fundamental reason behind differential competition for mates in the two sexes, and as a strong determinant behind differences in choosiness. This view has been challenged by Kokko and Monaghan, who argue that sex-specific parental investment, mortalities, mate-encounter rates and quality variation determine the mating system in a way that is not reducible to the OSR. We develop a game-theoretic model of choosiness, signalling and parental care, to examine (i) whether the results of Kokko and Monaghan remain robust when its simplifying assumptions are relaxed, (ii) how parental care coevolves with mating strategies and the OSR and (iii) why mutual mate choice is observed relatively rarely even when both sexes vary in quality. We find qualitative agreement with the simpler approach: parental investment is the primary determinant of sex roles instead of the OSR, and factors promoting choosiness are high species-specific mate-encounter rate, high sex-specific mate-encounter rate, high cost of breeding (parental investment), low cost of mate searching and highly variable quality of the opposite sex. The coevolution of parental care and mating strategies hinders mutual mate choice if one parent can compensate for reduced care by the other, but promotes it if offspring survival depends greatly on biparental care. We argue that the relative rarity of mutual mate choice is not due to biases in the OSR. Instead, we describe processes by which sexual strategies tend to diverge. This divergence is prevented, and mutual mate choice maintained, if synergistic benefits of biparental care render parental investment both high and not too different in the two sexes.  (+info)

A theoretical analysis of the energetic costs and consequences of parental care decisions. (15/195)

Should a parent care for its young or abandon them before they reach independence? We consider parental care behaviour as an adaptive decision, involving trade-offs between current and future reproduction. The condition of the parent is expected to influence these trade-offs. Using a dynamic programming model we explore how changes in the levels of energetic reserves, and time in the season, determine changes in parental care decisions. The novel feature of our model is that we have included the possibility of remating within the current breeding season in a consistent manner by explicitly modelling the behaviour of unmated animals. We show that there may be several fluctuations in the average duration of care during the breeding season. We also show that, because of the dependence of parental care behaviour on both the condition of the parent and time during the breeding season, changing some of the costs of care may increase the duration of care during one part of the season and decrease it at another. The model also shows that the conditions prevailing for animals with dependent offspring can affect the way in which an unmated animal behaves. For example, the behaviour of unmated animals may change to compensate (partly) for increases in the costs of raising offspring, which are produced at a later date (for example, by increasing the duration of foraging between breeding attempts). Overall, the model provides a good framework for understanding how various ecological and life-history variables should influence parental care behaviour during a breeding season.  (+info)

Relating paternity to paternal care. (16/195)

Intuition suggests, to most people, that parents should be selected to care for their offspring in relation to how certain they are of being the parents of those offspring. Theoretical models of the relationship between parental investment and certainty of parentage predict the two to be related only when some other assumptions are made, few of which can be taken for granted. I briefly review the models and their assumptions, and discuss two kinds of difficulty facing an empiricist wishing to test the models. The first is the problem of unmeasured (and immeasurable) variables. The second is the problem that even the most extensive models do not capture the complexity that can be demonstrated in real systems. I illustrate some of these problems, and some qualitative tests of the models, with experimental work on a population of the collared flycatcher. My conclusion is that although there are some cases where the models have qualitative support, we are a long way from understanding whether paternal care is commonly adjusted in relation to certainty of paternity.  (+info)