Models of parent-offspring conflict. IV. Suppression: Evolutionary retaliation by the parent

We have earlier analysed ESSs for the amount of parental investment (PI) that offspring are expected to solicit from their parents, given that parents acquiesce to offspring demands. The present paper considers evolutionary retaliation by the parent for species where only one parent provides PI. Two genetic loci are envisaged: one (the ‘conflictor’ locus) determines the extent of offspring solicitation; the other (the ‘suppressor’ locus) determines how parents retaliate. Solicitation is assumed to carry a cost which may affect a particular offspring uniquely if time and energy are the major costs, or may affect all offspring in a brood equally if the main cost is predation risk. Two kinds of parental retaliation are possible. Parents may supply PI in proportion to offspring demands, or may ignore solicitation altogether and give a fixed PI. Analytical models of conflict in which the parent supplies PI in proportion to solicitation yield pure ESSs with PI at a compromise level between parent and offspring interests. These are termed ‘pro rata’ ESSs. Where solicitation costs are high, an ‘offspring wins’ ESS (offspring get all they ‘want’) is possible especially for forms of conflict that affect future sibs, and a ‘parent wins’ ESS (parent supplies its optimum) is possible especially for conflict that affects contemporary sibs. When parental retaliation takes the form of ignoring offspring solicitation, this can lead to a ‘parent wins’ ESS if costs of ignoring solicitation are negligible, but where parental insensitivity carries costs, the result is an unresolvable evolutionary chase with cycling frequencies of alleles coding for parent and offspring strategies. ‘Pro rata’ ESSs cannot be invaded by ‘ignore solicitation’ mutants but ‘pro rata’ mutants can often invade at certain stages in ‘ignore solicitation’ limit cycles. We therefore conclude that the probable evolutionary end product for most species will be the ‘pro rata’ ESS in which the parent supplies more PI than would be optimal in the absence of conflict, but less PI than would be an ESS for the offspring in the absence of parental retaliation. Such ESSs will be characterized by solicitation costs; offspring will ‘ask’ for more PI than they get. In nature, under similar conditions, highest conflict will occur when both parents sustain equally the effects of conflict, or when conflict affects contemporary rather than future sibs.     

Procreative and performative kinship among residents in Victorian caravan parks

Recent senior anthropologists have heralded a new era where performative social and cultural kinship may eclipse procreational ‘biological kinship’ in contemporary western society. This article takes research on permanent residents in caravan parks as an extreme case that may act as an exemplar for future western contexts. Between 2003 and 2008, interviews with caravan‐park managers, 50 interviews with park residents and short field stays at 17 outer Melbourne and rural Victorian caravan parks, provided multi‐sited ethnographic material for analysis. I argue that contemporary contexts for relatedness give scope for creative development of social performative ties with ‘ex’ spouses and in‐laws, friends and pets. ‘Biological’ kin and family with a material dimension remain central, however, and a response to family need or rupture is often the broadening of the procreative kin links between grandparent and grandchild and between siblings. Childhood institutionalisation and divorce in particular, contributed to pathways towards caravan park housing. Other kin‐based motivations include finding safe accommodation for wives; providing housing for young adult males after injury and family divorce; and arranging for housing succession for children. Ideas of blood and folk biology demonstrate considerable resilience in contemporary kinship arrangements.     

Models of parent-offspring conflict. V. Effects of the behaviour of the two parents

In the absence of any parent-offspring conflict, the total parental investment per offspring should be less when two parents collaborate in caring for the offspring than when only one parent invests. This does not necessarily mean that offspring fare less well when both parents invest. The ‘ideal’ amount of parental investment for an offspring to take is always greater than is ‘ideal’ for the parent to allocate (Trivers 1974). The offspring's optimum is higher if the offspring's action affects the reproductive success of only one parent and lower if both parents are affected (e.g. two-parent investment, or lifelong monogamy). The difference between the parental optimum and the offspring optimum depends on the mating system and on the form of conflict (between successive broods, or within broods), and prescribes a ‘conflict range’. The extent of conflict cannot be deduced solely from a knowledge of the average relatedness between siblings. The conflict is likely to be resolved by an ESS in which intermediate (compromise) levels of investment are paid out to offspring, which nevertheless continue to make costly demands for yet more investment. The degree of conflict can be measured by the extent to which offspring subject their parents to aggressive demands for extra investment, and is likely to be greater when two parents collaborate equally over investment than when only one parent invests. When only one parent invests, conflict is higher if sibling-competition is between siblings in the same broods (intra-brood) than when it is between progeny in successive broods (inter-brood). However, the reverse will tend to be the case when both parents invest equally.     

Why join a neighbour: fitness consequences of colony fusions in termites

The evolution of life is characterized by major evolutionary transitions during which independent units cooperated and formed a new level of selection. Relatedness is a common mechanism that reduces conflict in such cooperative associations. One of the latest transitions is the evolution of social insect colonies. As expected, they are composed of kin and mechanisms have evolved that prevent the intrusion of nonrelatives. Yet, there are exceptions an extreme case is the fusion of unrelated colonies. What are the advantages of fusions that have colonies with a high potential for conflict as a consequence? Here, we investigated fitness costs and benefits of colony fusions in a lower termite species, Cryptotermes secundus, in which more than 25% of all colonies in the field are fused. We found two benefits of colony fusion depending on colony size: very small colonies had an increased probability of survival when they fused, yet for most colony sizes mainly a few workers profit from colony fusions as their chance to become reproductives increased. This individual benefit was often costly for other colony members: colony growth was reduced and the current reproductives had an increased chance of dying when fusions were aggressive. Our study suggests that fusion of colonies often is the result of ‘selfish’ worker interests to become reproductives, and this might have been important for the termites' social evolution. Our results uniquely shows that selfish interests among related colony members can lead to the formation of groups with increased potential for conflict among less related members.     

Sexual selection modulates genetic conflicts and patterns of genomic imprinting

Recent years have seen a surge of interest in linking the theories of kin selection and sexual selection. In particular, there is a growing appreciation that kin selection, arising through demographic factors such as sex‐biased dispersal, may modulate sexual conflicts, including in the context of male–female arms races characterized by coevolutionary cycles. However, evolutionary conflicts of interest need not only occur between individuals, but may also occur within individuals, and sex‐specific demography is known to foment such intragenomic conflict in relation to social behavior. Whether and how this logic holds in the context of sexual conflict—and, in particular, in relation to coevolutionary cycles—remains obscure. We develop a kin‐selection model to investigate the interests of different genes involved in sexual and intragenomic conflict, and we show that consideration of these conflicting interests yields novel predictions concerning parent‐of‐origin specific patterns of gene expression and the detrimental effects of different classes of mutation and epimutation at loci underpinning sexually selected phenotypes.     

Support and conflict of kinsmen in Norse Earldoms,Icelandic families,and the English royalty

Data concerning the Norse Earls of the Orkney Islands, Icelandic families, and the conflicts concerning the sovereignty of England are presented. The data suggest that support vs conflict with kin is influenced by degree of biological relatedness but also by the needs for support by kinsmen, as influenced by the ability to hire retainers, and by the costs and benefits of support vs conflict.     

The problem of peers in Vietnamese interaction

In Vietnamese, address (second‐person reference) is typically accomplished by the use of a kin term regardless of whether the talk's recipient is a genealogical relative or not. All Vietnamese kin terms encode a specification of either relative age or relative generation of participants, and there are no reciprocal terms akin to English ‘brother’ or ‘sister’; rather, a speaker must select between terms such as ‘older brother’ (anh) or ‘younger sibling’ (em). Since generation is normatively associated with a difference in age, the result is a ubiquitous indexing of age and status hierarchies in all acts of address. This results in a problem for peers. How, in such a system, should they address one another (and also self‐refer)? In this article, we describe the various practices that speakers use to subvert the system and thus avoid indexing differences of age or station. Specifically, we describe four practices: (1) the use of true pronouns in address and self‐reference; (2) the use of proper names in address and self‐reference; (3) the use of kin terms in address and pronouns in self‐reference; and (4) the ironic use of kin terms in address. We conclude that the Vietnamese system well illustrates what is likely a universal tension between hierarchy and equality in acts of address and self‐reference, by showing how speakers deconstruct the vector of age and indicate that they consider one another peers. We further suggest that although the literature in this area has focused on the ways in which languages convey differences of status and rank, social order is built as much upon relations of parity and sameness – on identification of the other as neither higher nor lower than me – as it is upon relations of hierarchy.     

Putting self‐organization to the test: labyrinthine patterns as optimal solution for persistence

Spatial patterns formed through the process of self‐organization are found in nature across a variety of ecosystems. Pattern formation may reduce the costs of competition while maximizing the benefits of group living, and thus promote ecosystem persistence. This leads to the prediction that self‐organizing to obtain locally intermediate densities will be the optimal solution to balance costs and benefits. However, despite much evidence documenting pattern formation in natural ecosystems, there is limited empirical evidence of how these patterns both influence and are influenced by tradeoffs between costs and benefits. Using mussels as a model system, we coupled field observations in mussel‐culture plots with manipulative laboratory experiments to address the following hypotheses: 1) labyrinthine spatial patterns, characteristically found at intermediate to high patch densities, are the most persistent over time; this is because labyrinthine patterns 2) result in adequately heavy patches that can maximize resistance to dislodgement while 3) increasing water turbulence with spacing, which will maximize food delivery processes. In the field, we observed that labyrinthine ‘stripes’ patterns are indeed the most persistent over time, confirming our first hypothesis. Furthermore, with laboratory experiments, we found the ‘stripes’ pattern to be highly resistant to dislodgement, confirming the second hypothesis. Finally, with regards to the third hypothesis, we found positive effects of this pattern on local turbulence. These results suggest that the mechanisms of intraspecific facilitation not only depend on initial organism densities, but may also be influenced by spatial patterning. We hence recommend taking into account spatial patterns to maximize productivity and persistence in shellfish‐cultivation practices and to increase the restoration success of ecosystems with self‐organizing properties.     

Reproductive competition between females in the matrilineal Mosuo of southwestern China

The matrilineal Mosuo of southwestern China live in communal households where brothers and sisters of three generations live together (duolocal residence), and men visit their wives, who reside elsewhere, only at night in ‘visiting’ marriages. Here we show that these communally breeding sisters are in reproductive conflict, in the sense that they share the resources needed to reproduce. We analyse determinants of reproductive success in females and males, and show that co-resident female kin are in competition; the more female kin reside in the household, the more reproductive success is reduced. Male reproductive success, however, is not determined by the kin in his natal household; duolocal males are not in reproductive conflict with their siblings. Competition with female cousins can be worse than that between sisters. We also find that female work on the farm (which is the main communal resource) is not equal. We use a ‘tug-of-war’ model of reproductive skew generated by incomplete control, to model the patterns of effort put into competition between sisters and cousins. The model predicts that more dominant (older) sisters will put less effort into reproductive conflict than will less dominant (younger) sisters; but younger sisters will also have lower reproductive success because they are less efficient at gaining access to the shared resource. Both predictions are consistent with our data. Younger sisters work less in the fields than do older sisters, which may represent a form of conflict or may be because their average relatedness to the household is lower than that of their more fertile older sisters.     

Dominance, Status Signals and Coloration in Male Mandrills (Mandrillus sphinx)

Where individuals contest access to a resource, escalated physical fighting presents a risk to all involved. The requirement for mechanisms of conflict management has led to the evolution of a variety of decision rules and signals that act to reduce the frequency of aggression during competitive encounters. We examined strategies of conflict management in male mandrills (Mandrillus sphinx) living in two semi‐free‐ranging groups in Gabon. Adult male mandrills are large (31 kg), with long canines, making the costs of conflict potentially very high. We found that males formed dominance hierarchies, but that male–male relationships were characterized by avoidance, appeasement and ignoring. Fights were rare, but could result in death. Examination of the relationship between dominance and signaling showed that males use facial and gestural signals to communicate dominance and subordinance, avoiding escalated conflict. Male mandrills also possess rank‐dependent red coloration on the face, rump and genitalia, and we examined the hypothesis that this coloration acts as a ‘badge of status’, communicating male fighting ability to other males. If this is the case, then similarity in color should lead to higher dyadic rates of aggression, while males that differ markedly should resolve encounters quickly, with the paler individual retreating. Indeed, appeasement (the ‘grin’ display), threats, fights and tense ‘stand‐off’ encounters were significantly more frequent between similarly colored males, while clear submission was more frequent where color differences were large. We conclude that male mandrills employ both formal behavioral indicators of dominance and of subordination, and may also use relative brightness of red coloration to facilitate the assessment of individual differences in fighting ability, thereby regulating the degree of costly, escalated conflict between well‐armed males.     

Nice to kin and nasty to non-kin: revisiting Hamilton's early insights on eusociality

When helping behaviour is costly, Hamiltonian logic implies that animals need to direct helpful acts towards kin, so that indirect fitness benefits justify the costs. We revisit inferences about nepotism and aggression in Hamilton''s 1964 paper to argue that he overestimated the general significance of nepotism, but that other issues that he raised continue to suggest novel research agendas today. We now know that nepotism in eusocial insects is rare, because variation in genetic recognition cues is insufficient. A lower proportion of individuals breeding and larger clutch sizes selecting for a more uniform colony odour may explain this. Irreversible worker sterility can induce both the fiercest possible aggression and the highest likelihood of helping random distant kin, but these Hamiltonian contentions still await large-scale testing in social animals.     

Conception,De-Conception and Social Structure in Bush Mekeo Culture

According to a contradiction of endogamy vs. exogamy (or of ‘blood’ vs. ‘non-blood’ relationship) in Bush Mekeo culture, non-relatives may marry and conceive by ‘mixing’ of distinct procreative clan bloods. By means of marriage and mortuary reciprocities, on the other hand, relatives or kin are ‘de-conceived’ through the ‘unmixing’ of non-procreative clan bloods. Thus, Bush Mekeo social structure is preserved through time: While the boundaries distinguishing unilineal descent groups are maintained constant, non-relatives are translated into affines who generate bilateral kin, and bilateral kin are translated into nonrelatives and, thereby, potential affines.     

Revisiting non-offspring nursing: allonursing evolves when the costs are low

Allonursing, the nursing of another female''s offspring, is commonly assumed to have evolved through the benefits of kin selection or reciprocity. The evolution of allonursing may also be influenced by variation in the possible costs to allonurses. The relative influence of costs and benefits on the incidence of allonursing in mammals remains unexplored. We show, using comparative analyses, that where females group with kin, the presence or the absence of allonursing is not associated with further variation in relatedness. Allonursing is most common where females produce litters; here the relative investment per offspring is low, and the costs of nursing additional young are likely to be reduced. Our results suggest that variation in the potential benefits is not associated with the distribution of allonursing, but that allonursing can quickly evolve when the costs to allonurses of nursing additional offspring are low.     

Sex biases in kin shoaling and dispersal in a cichlid fish

Animal dispersal is associated with diverse costs and benefits that vary among individuals based on phenotype and ecological conditions. For example, females may disperse when males benefit more from defending territories in familiar environments. Similarly, size differences in dispersal propensity may occur when dispersal costs are size-dependent. When individuals do disperse, they may adopt behavioral strategies that minimize dispersal costs. Dispersing fish, for example, may travel within shoals to reduce predation risks. Further, kin shoaling may augment inclusive fitness by reducing predation of relatives. However, studies are lacking on the role of kin shoaling in dispersal. We explored how sex and size influence dispersal and kin shoaling in the cichlid Neolamprologus caudopunctatus. We microsatellite genotyped over 900 individuals from two populations separated by a potential dispersal barrier, and documented patterns of population structure, migration and within-shoal relatedness. Genetic differentiation across the barrier was greater for smaller than larger fish, suggesting larger fish had dispersed longer distances. Females exhibited weaker genetic differentiation and 11 times higher migration rates than males, indicating longer-distance female-biased dispersal. Small females frequently shoaled with siblings, possibly offsetting dispersal costs associated with higher predation risks. In contrast, small males appeared to avoid kin shoaling, possibly to avoid local resource competition. In summary, long-distance dispersal in N. caudopunctatus appears to be female-biased, and kin-based shoaling by small females may represent a behavioral adaptation that reduces dispersal costs. Our study appears to be the first to provide evidence that sex differences in dispersal influence sex differences in kin shoaling.     

The influence of kinship and dominance hierarchy on grooming partner choice in free-ranging <Emphasis Type="Italic">Macaca mulatta brevicaudus</Emphasis>

In group-living animals, individuals do not interact uniformly with their conspecifics. Among primates, such heterogeneity in partner choice can be discerned from affiliative grooming patterns. While the preference for selecting close kin as grooming partners is ubiquitous across the primate order, the selection of higher-ranking non-kin individuals as grooming partners is less common. We studied a group of provisioned rhesus macaques (Macaca mulatta brevicaudus) on Hainan Island, China, to examine rank-related benefits of grooming exchanges and the influence of kin relationships. We tested four hypotheses based on Seyfarth’s model: (1) there will be kin preference in grooming relationships; (2) grooming between non-kin individuals will be directed up the dominance rank; (3) grooming between non-kin individuals will reduce aggression from higher-ranking ones; and (4) non-kin individuals will spend more time grooming with adjacent ranked ones. We found that grooming relationships between kin individuals were stronger than those between non-kin individuals. For non-kin relationships, lower-ranking individuals received less aggression from higher-ranking ones through grooming; a benefit they could not derive through grooming exchanges with individuals related by kinship. Individuals spent more time grooming adjacent higher-ranking non-kin individuals and higher-ranking individuals also received more grooming from non-kin individuals. Our results supported Seyfarth’s model for predicting partner choice between non-kin individuals. For relationships between kin individuals, we found results that were not consistent with prediction for the exchanges of aggression and grooming, indicating the importance to control for the influence of kinship in future studies.     

The socio‐genetics of a complex society: female gelada relatedness patterns mirror association patterns in a multilevel society

Multilevel societies with fission–fusion dynamics—arguably the most complex animal societies—are defined by two or more nested levels of organization. The core of these societies are modular social units that regularly fission and fuse with one another. Despite convergent evolution in disparate taxa, we know strikingly little about how such societies form and how fitness benefits operate. Understanding the kinship structure of complex societies could inform us about the origins of the social structure as well as about the potential for individuals in these societies to accrue indirect fitness benefits. Here, we combined genetic and behavioural data on geladas (Theropithecus gelada), an Old World Monkey, to complete the most comprehensive socio‐genetic analysis of a multilevel society to date. In geladas, individuals in the core social ‘units’, associate at different frequencies to form ‘teams’, ‘bands’ and, the largest aggregations, ‘communities’. Units were composed of closely related females, and females remained with their close kin during permanent fissions of units. Interestingly, female–female relatedness also significantly predicted between‐unit, between‐team and between‐band association patterns, while male–male relatedness did not. Thus, it is likely that the socio‐genetic structure of gelada society results from females maintaining associations with their female relatives during successive unit fissions—possibly in an attempt to balance the direct and indirect fitness benefits of group living. Overall, the persistence of associations among related females across generations appears to drive the formation of higher levels of gelada society, suggesting that females seek kin for inclusive fitness benefits at multiple levels of gelada society.     

The costs and benefits of resource sharing: reciprocity requires resource heterogeneity

The evolution of resource sharing requires that the fitness benefits to the recipients be much higher than the costs to the giver, which requires heterogeneity among individuals in the fitness value of acquiring additional resources. We develop four models of the evolution of resource sharing by either direct or indirect reciprocity, with equal or unequal partners. Evolution of resource sharing by reciprocity requires differences between interacting individuals in the fitness value of the resource, and these differences must reverse although previous acts of giving are remembered and both participants survive. Moreover, inequality in the expected reproductive value of the interacting individuals makes reciprocity more difficult to evolve, but may still allow evolution of sharing by kin selection. These constraints suggest that resource sharing should evolve much more frequently by kin selection than by reciprocity, a prediction that is well supported by observations in the natural world.     

Evolution of sex ratios in social hymenoptera: kin selection,local mate competition,polyandry and kin recognition

A model is constructed to study the effects of local mate competition and multiple mating on the optimum allocation of resources between the male and female reproductive brood in social hymenopteran colonies from the ‘points of view’ of the queen (parental manipulation theory) as well as the workers (kin selection theory). Competition between pairs of alleles specifying different sex investment ratios is investigated in a game theoretic frame work. All other things being equal, local mate competition shifts the sex allocation ratio in favour of females both under queen and worker control. While multiple mating has no effect on the queen’s optimum investment ratio, it leads to a relatively male biased investment ratio under worker control. Under queen control a true Evolutionarily Stable Strategy(ess) does not exist but the ‘best’ strategy is merely immune from extinction. A trueess exists under worker control in colonies with singly mated queens but there is an asymmetry between the dominant and recessive alleles so that for some values of sex ratio a recessive allele goes to fixation but a dominant allele with the same properties fails to do so. Under multiple mating, again, a trueess does not exist but a frequency dependent region emerges. The best strategy here is one that is guaranteed fixation against any competing allele with a lower relative frequency. Our results emphasize the need to determine levels of local mate competition and multiple mating before drawing any conclusions regarding the outcome of queen-worker conflict in social hymenoptera. Multiple mating followed by sperm mixing, both of which are known to occur in social hymenoptera, lower average genetic relatedness between workers and their reproductive sisters. This not only shifts the optimum sex ratio from the workers’ ‘point of view’ in favour of males but also poses problems for the kin selection theory. We show that kin recognition resulting in the ability to invest in full but not in half sisters reverts the sex ratio back to that in the case of single mating and thus completely overcomes the hurdles for the operation of kin selection.     

Benefits of parental care: Do juvenile lizards obtain better‐quality habitat by remaining with their parents?

Abstract Evolutionary theory suggests that parental care is favoured by natural selection when the benefits to offspring fitness outweigh the costs of parental expenditure. The nature of such benefits may differ among species, however, especially in species reflecting independent evolutionary origins of parental care. Black rock skinks (Egernia saxatilis, Scincidae) of south‐eastern Australia are viviparous rock‐dwelling lizards with prolonged parent–offspring association; adult pairs vigorously defend their home range – and, when present, their offspring – against conspecifics. We addressed the hypothesis that, by remaining within their parents' (vigorously defended) home range, juveniles thereby obtain access to better‐quality habitats. Measurements of biologically significant variables (crevice size, sun exposure, vegetation cover) revealed little difference between shelter‐rocks used by solitary (‘orphan’) juveniles and those within family groups containing adults. Indeed, the only consistent differences involved larger (and therefore, less predator‐proof) crevices for juveniles within family groups than for solitary conspecifics. Our data thus falsify the hypothesis that parental care evolves because of benefits associated with habitat quality; instead, it appears that parental protection of juveniles against infanticidal conspecifics may be the most plausible benefit to parent–offspring association in this system.     

Shared and unshared parental investment,parent-offspring conflict and brood size

Taking as our starting point Trivers' (1974) account of parent-offspring conflict, we develop models of the influence of brood size on the optimal level of parental investment (PI) in the whole brood for parent and offspring, and on the magnitude of conflict between them. A modification of Trivers' model is proposed. In general, the benefit of an act of PI to an offspring in a brood of size N is (N+1)/N times the benefit to its parent. Therefore as brood size increases, offspring benefit approaches parental benefit, and this is because an increasing proportion of the offspring's benefit is being gained through siblings, to which offspring and parent are equally related. A distinction is drawn between ‘shared’ and ‘unshared’ types of PI. When PI is shared the total benefit accruing is not directly gained by all offspring but is shared amongst them (e.g. food brought to the young). In contrast, unshared PI can simultaneously benefit some or all of the brood (e.g. types of anti-predator defence). For shared investment, PI and conflict are predicted to increase with brood size. Two models of unshared anti-predator defence are described. If the predator characteristically takes the whole brood when it strikes (e.g. altricial nestlings) PI is predicted to increase and conflict decline with brood size, although this effect is inhibited or even reversed for high risk defence tactics because of the higher cost to larger broods if the parent dies. When the predator takes a single offspring (e.g. precocial birds) the parent's optimum PI is independent of brood size, the offspring's optimum PI declines in larger broods and conflict again declines with brood size. The parent is commonly expected to win the conflict over anti-predator care. Predictions concerning PI levels gain support from existing data, largely for birds, but evaluation of those for conflict must await the collection of new data. The distinction between shared and unshared investment is applicable to altruistic behaviour in general.