Helpful Female Subordinate Cichlids Are More Likely to Reproduce

Background

In many cooperatively breeding vertebrates, subordinates assist a dominant pair to raise the dominants'' offspring. Previously, it has been suggested that subordinates may help in payment for continued residency on the territory (the ‘pay-to-stay hypothesis’), but payment might also be reciprocated or might allow subordinates access to reproductive opportunities.

Methodology/Principal Findings

We measured dominant and subordinate female alloparental brood care and reproductive success in four separate experiments and show that unrelated female dominant and subordinate cichlid fish care for each other''s broods (alloparental brood care), but that there is no evidence for reciprocal ‘altruism’ (no correlation between alloparental care received and given). Instead, subordinate females appear to pay with alloparental care for own direct reproduction.

Conclusions/Significance

Our results suggest subordinate females pay with alloparental care to ensure access to the breeding substrate and thereby increase their opportunities to lay their own clutches. Subordinates'' eggs are laid, on average, five days after the dominant female has produced her first brood. We suggest that immediate reproductive benefits need to be considered in tests of the pay-to-stay hypothesis.     

Reproductive suppression in female cooperatively breeding cichlids

Suppression by dominants of female subordinate reproduction has been found in many vertebrate social groups, but has rarely been shown experimentally. Here experimental evidence is provided for reproductive suppression in the group-living Lake Tanganyika cichlid Neolamprologus pulcher. Within groups of three unrelated females, suppression was due to medium- and small-sized females laying less frequently compared with large females, and compared with medium females in control pairs. Clutch size and average egg mass of all females depended on body size, but not on rank. In a second step, a large female was removed from the group and a very small female was added to keep the group size constant. The medium females immediately seized the dominant breeding position in the group and started to reproduce as frequently as control pairs, whereas clutch size and egg mass did not change. These results show that female subordinate cichlids are reproductively capable, but apparently suppressed with respect to egg laying. Nevertheless, some reproduction is tolerated, possibly to ensure continued alloparental care by subordinate females.     

Clutch-size adjustments and skew models: effects on reproductive partitioning and group stability

Reproductive skew theory seeks to integrate social and ecologicalfactors thought to influence the division of reproduction amonggroup-living animals. However, most reproductive skew modelsonly examine interactions between individuals of the same sex.Here, we suggest that females can influence group stabilityand conflict among males by modifying their clutch size andmay do so if they benefit from the presence of subordinate malehelpers or from reduced conflict. We develop 3 models, basedon concessions-based, restraint, and tug-of-war models, in whichfemale clutch size is variable and ask when females will increasetheir clutch size above that which would be optimal in the absenceof male–male conflict. In concessions-based and restraintmodels, females should increase clutch size above their optimaif the benefits of staying for subordinate males are relativelylow. Relatedness between males has no effect on clutch size.When females do increase clutch size, the division of reproductionbetween males is not influenced by relatedness and does notdiffer between restraint and concessions-based models. Bothof these predictions are in sharp contrast to previous models.In tug-of-war models, clutch size is strongly influenced byrelatedness between males, with the largest clutches, but thefewest surviving offspring, produced when males are unrelated.These 3 models demonstrate the importance of considering third-partyinterests in the decisions of group-living organisms.     

Fitness consequences of divorce in the oystercatcher, Haematopus ostralegus

We investigated the fitness consequences of divorce in oystercatchers. We made a distinction between two types of divorce: in desertions the disruption of the pair bond is initiated by one of the pair members, and in usurpations by a conspecific individual. Survival and reproduction prospects for oystercatchers are largely determined by their social status (nonbreeder or breeding bird at a site of a specific quality). Changes in social status in relation to divorce showed that birds taking the initiative to leave their mate increased in fitness, relative to birds that were forced to leave their partner. Status of individuals that remained in their territory after divorce was unaffected if their mate was expelled, but declined if their mate deserted. Survival after divorce was significantly lower for birds that were expelled than for those deserting. Divorce rate, and especially desertion rate, was higher among occupants of low- than high-quality territories. In general, divorce rate increased following elevated mortality. In high-quality territories usurpations increased with increasing breeder mortality, but at low-quality territories this relation was absent. Desertion rates were similarly related to mortality in both territory types. Divorce participants thus differed strongly in their fitness prospects, depending on the type of divorce, the role played in the divorce and the quality of the territory where divorce took place. Studies that do not observe the birds during divorce cannot determine the type of divorce and the role played by the individuals, and this may lead to misleading conclusions on the costs and benefits of divorce. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

A silver spoon for a golden future: long-term effects of natal origin on fitness prospects of oystercatchers (Haematopus ostralegus)

1. Long-term effects of conditions during early development on fitness are important for life history evolution and population ecology. Using multistrata mark-recapture models on 20 years of data, we quantified the relation between rearing conditions and lifetime fitness in a long-lived shorebird, the oystercatcher (Haematopus ostralegus). We addressed specifically the relative contribution of short- and long-term effects of rearing conditions to overall fitness consequences. 2. Rearing conditions were defined by differences in natal habitat quality, in which there is a clear dichotomy in our study population. In the first year of life, fledglings from high-quality natal origin had a 1.3 times higher juvenile survival. Later in life (age 3-11), individuals of high-quality natal origin had a 1.6 times higher adult prebreeder survival. The most striking effect of natal habitat quality was that birds that were reared on high-quality territories had a higher probability of settling in high-quality habitat (44% vs. 6%). Lifetime reproductive success of individuals born in high-quality habitat was 2.2 times higher than that of individuals born in low-quality habitat. This difference increased further when fitness was calculated over several generations, due to a correlation between the quality of rearing conditions of parents and their offspring. 3. Long-term effects of early conditions contributed more to overall fitness differences as short-term consequences, contrary to common conceptions on this issue. 4. This study illustrates that investigating only short-term effects of early conditions can lead to the large underestimation of fitness consequences. We discuss how long-term consequences of early conditions may affect settlement decisions and source-sink population interactions.     

Genetic analysis of the population structure of socially organized oystercatchers (Haematopus ostralegus) using microsatellites

On the island of Schiermonnikoog (The Netherlands), the breeding population of oystercatchers can be divided into two groups: 'residents' and 'leapfrogs', based on their distinct social characteristics and limited probabilities of status change between breeding seasons. In order to investigate whether this social organization has caused local genetic differentiation, leapfrogs and residents were compared at eight polymorphic microsatellite loci. No significant genetic subdivision between residents and leapfrogs was observed (theta = 0.0000; 95% confidence interval (CI), -0.0027-0.0033), indicating that the oystercatcher population on the island of Schiermonnikoog has to be considered as one panmictic unit. Investigation of three additional locations in the northern part of The Netherlands did not reveal significant genetic population subdivision either (theta = -0.0005; 95% CI, -0.0045-0.0037), despite the fact that adult osytercatchers show extreme fidelity to their breeding localities. These results indicate panmixis and considerable levels of gene flow within the northern part of The Netherlands. Thus, the results from genetical analyses do not seem to be in agreement with observational data on the dispersal behaviour of breeding individuals. It is argued that the lack of population structure, locally on Schiermonnikoog as well as across larger geographical distances, is to be attributed to high levels of gene flow through dispersal of juvenile birds.     

Helpers in a cooperatively breeding cichlid stay and pay or disperse and breed, depending on ecological constraints

The theory of family-group dynamics predicts that group structure, helping behaviour and social interactions among group members should vary with the opportunities of subordinates to breed independently. We investigated experimentally whether unrelated mature helpers in the cooperatively breeding cichlid Neolamprologus pulcher reduce costly social and cooperative behaviour and choose to disperse and breed independently when offered vacant breeding sites. As predicted by the ecological constraints hypothesis,when breeding substrate was available, (i) helpers spent more time in dispersal areas and it was mainly large helpers that left the group to breed independently; (ii) all helpers invested less in costly submissive behaviours towards other group members and large helpers reduced help, supporting the 'pay-to-stay' hypothesis; and (iii) large helpers, particularly those that dispersed and bred, increased more in body mass in the treatment than those without breeding options, suggesting status-dependent strategic growth of helpers.We conclude that helpers of N. pulcher decide whether to stay and pay or disperse and breed in response to constraints on independent breeding.