Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Pair and extra-pair mating success relative to male quality in red-winged blackbirds

We tested whether the reproductive success of male red-winged blackbirds (Agelaius phoeniceus) varied with male secondary sexual traits or with haematozoa prevalence, and whether these patterns were consistent with females preferring genetically superior males. We also determined whether the traits that correlated with male success on their own territories were also correlated with male success at siring young on other males territories. Our analysis included data from a 6-year study involving 617 nestlings for which paternity was determined by DNA profiling. Larger males sired more young on their territories, principally because they obtained larger harems. The success of larger males at acquiring more mates did not appear to be a consequence of larger males holding larger or better-quality territories. Older and longer-lived males sired more young by extra-pair fertilizations. Larger males sired the most offspring overall (on territory + off territory). Variation in epaulet size and color, responses to male and female models, nest defence and parasitism was not correlated with male success either on or off their territories or overall. Male success in a given year was significantly correlated with success the previous year, as expected if females were selecting genetically superior males. The male that was by far the most successful individual in this study was highly consistent from year to year. Because male body size is positively correlated with survival in this population (although not within the sample of males included in this study), female preference for larger males may have reflected a general preference for males with superior Survival ability. We propose that the direct advantage realized by older males in extra-pair matings might indicate that experience is important, such that experienced males are better at creating or exploiting the opportunities for extra-pair mating. This hypothesis is consistent with a pairwise analysis of cuckoldry that showed that cuckolders were most often older than the males they cuckolded.     

Male residence and the patterning of serum testosterone in vervet monkeys (<Emphasis Type="Italic">Cercopithecus aethiops</Emphasis>)

This study investigated hormonal and demographic processes underlying unimale and multimale mating systems in primates. Reproductive skew and challenge models of male competition provide conflicting predictions of the relationship of male residence to group composition and androgen regulation. These predictions were tested using endocrine and socioecological data from Kenyan vervet monkeys (Cercopithecus aethiops). Serum samples from 57 adult male monkeys, drawn from 19 separate groups and 4 populations, were assayed for testosterone by radioimmunoassay. Male ability to respond to conspecific challenge was assessed by their testosterone response to the capture protocol.Analyses showed that reproductive skew models were useful predictors of intergroup and interpopulation variation in male residence and T profiles. The Limited Control model of male residence was supported by positive correlations of the number of males per group with the number of females without dependent offspring, demonstrating that monopolization potential was a key determinant of male residence. Testosterone concentrations under conditions that elevated serum levels were positively correlated with infanticide risk, supporting the Concession model. Population comparisons provided evidence for increased T responsiveness where groups were predominantly unimale. Unimale populations were from sites with higher rainfall, suggesting that ecological factors contributed to population differences in male residence and T regulation.For species name, we follow the 2000 taxonomy of the IUCN/SSC Primate Specialist Groups workshop (Grubb et al. 2003)     

Reproductive plasticity in bumblebee workers (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)—reversion from fertility to sterility under queen influence

Worker sterility in the bumblebee Bombus terrestris is conditional and is linked to the social development of the colony. Workers refrain from reproducing or overtly challenging the queen until gyne production has initiated, at the so-called competition point (CP). It is not known whether this behavior is hard-wired or workers show reproductive plasticity. It also remains unclear whether worker reproductive decision is under queen and/or worker control. In this study, we tested worker reproductive plasticity in an attempt to assess whether and under which conditions worker sterility/fertility are reversible. We introduced egg-laying workers into colonies with different social structures for 1 week then monitored their reproductive status. We revealed a remarkable reproductive plasticity in the introduced workers that was social-condition-dependent. In the presence of a pre-CP queen, the introduced workers reverted to sterility, whereas in the presence of a post-CP queen, such workers remained egg-layer. Reversion to sterility does not occur when direct contact with the queen is prevented, as the introduced workers remained egg-layer in the queenright colonies with a confined queen. Egg-laying workers that were introduced into queenless colonies mostly maintained their fertility regardless of colony social phase. This shows that worker transition from cooperative to selfish behavior is reversible depending on the social context.     

Cooperative breeding increases reproductive success in the social spider <Emphasis Type="Italic">Stegodyphus dumicola</Emphasis> (Araneae,Eresidae)

Sociality in some birds, mammals, and social insects was suggested to have evolved through the lengthening and extension of parental care behaviors to nondirect descendents. In these systems, group members care for young cooperatively and, thus, increase the reproductive success of the breeders and fitness of the young. Parental care behaviors, such as regurgitation feeding and matriphagy (consumption of the mother), occur in several subsocial and social spiders. However, it is not known whether females in a colony cooperate in caring for the young of other females and whether such cooperative care improves reproductive success. To answer this question, we created experimental colonies of the social spider Stegodyphus dumicola (Araneae, Eresidae), allowing only one female in a group to produce young, simulating reproductive skew occurring in nests in nature. In this paper, we show for the first time that females of S. dumicola cooperate in providing regurgitated food for young of other females and are even eaten by those young. Young raised by a group of females were larger and had greater survival than young raised only by their mother. Thus, fitness benefits from raising broods cooperatively may have favored the evolution of sociality in spiders.     

Reproductive stage and phonotactic preferences of female midwife toads (<Emphasis Type="Italic">Alytes cisternasii</Emphasis>)

We studied the phonotactic response of Alytes cisternasii females at different reproductive stages (before and after oviposition) toward two male call characteristics (call rate and frequency) differing in variability, as well as in the strength of female preference. We measured female phonotactic response over several repetitions by means of the frequency of selection of the population-preferred alternative, consistency of choice, frequency of reciprocal calling activity, frequency of null trials, and average approach time. Our results showed that female mating preferences in A. cisternasii are highly influenced by the variability of male call characteristics, while only female motivation is related to reproductive stage.Communicated by T. Czeschlik     

The reproductive dilemmas of queenless red dwarf honeybee (Apis florea) workers

Honeybee (Apis) workers cannot mate, but retain functional ovaries. When colonies have lost their queen, many young workers begin to activate their ovaries and lay eggs. Some of these eggs are reared, but most are not and are presumably eaten by other workers (worker policing). Here we explore some of the factors affecting the reproductive success of queenless workers of the red dwarf honeybee Apis florea. Over a 2-year period we collected 40 wild colonies and removed their queens. Only two colonies remained at their translocated site long enough to rear males to pupation while all the others absconded. Absconding usually occurred after worker policing had ceased, as evidenced by the appearance of larvae. Dissections of workers from eight colonies showed that in A. florea, 6% of workers have activated ovaries after 4 days of queenlessness, and that 33% of workers have activated ovaries after 3 weeks. Worker-laid eggs may appear in nests within 4 days and larvae soon after, but this is highly variable. As with Apis mellifera, we found evidence of unequal reproductive success among queenless workers of A. florea. In the two colonies that reared males to pupation and which we studied with microsatellites, some subfamilies had much higher proportions of workers with activated ovaries than others. The significance of absconding and internest reproductive parasitism to the alternative reproductive strategies of queenless A. florea workers is discussed.     

Sex-specific differences in site fidelity and the cost of dispersal in yellow-headed blackbirds

Male migratory birds tend to be more faithful than females to previous breeding sites, suggesting sex differences in costs or benefits of dispersal. In Illinois, greater site fidelity by male yellow-headed blackbirds (Xanthocephalus xanthocephalus) was associated with reduced reproductive success the following year for dispersers relative to non-dispersers. Dispersing females suffered no reduction in reproductive success the following year. Males that attracted few social mates, and thus had low reproductive success, were more likely to disperse, whereas females dispersed in response to low-patch reproductive success, regardless of their individual performance. Males that dispersed appeared to be successful acquiring territories because none was observed as a floater. The rate of dispersal by males in this low-density population was greater than in more dense populations where dispersing males may be less successful at acquiring territories. Despite success at obtaining territories, males that dispersed acquired territories on the periphery of wetlands where fewer females nested, resulting in lower reproductive success. In the second year after dispersing, however, males moved onto more central territories where they acquired larger harems. Thus, dispersal by males may be a long-term strategy requiring at least 2 years for benefits to be realized. Long-term success was enhanced because dispersing males moved to wetlands on which reproductive success was higher than on the wetlands they left. In addition to demonstrating that both individual and patch reproductive success affect dispersal decisions, these data indicate that when evaluating costs and benefits of dispersal, researchers should use a time frame beyond 1 year.     

Dufour’s gland pheromone as a reliable fertility signal among honeybee (Apis mellifera) workers

When a colony becomes queenless and without the possibility of requeening, honeybee workers initiate reproduction and lay male eggs about a week later. Assays in which two bees were confined in a small arena revealed that they establish a reproductive dominance hierarchy, i.e., one worker demonstrates greater ovarian development than her paired bee. Reproductive dominance is independent of relatedness, and can be established between full sisters, cousins, or random nestmates. A social environment, however, is obligatory, as singly housed bees fail to develop ovaries on the same time scale. Allowing varying degrees of social interactions between the paired bees revealed that olfaction of volatile bee compounds, as well as tactile communication, seem to provide the necessary social environment. Ovarian development was accompanied by the production of queen-like Dufours gland secretion in these workers. Especially notable was the increase in the queen-like esters. This increase was tightly linked to ovarian development and not necessarily to the dominance status of the bees in the pair. Thus, the occurrence of queen-like esters can serve as a reliable fertility signal. Advertising ovarian status may recruit helper workers with less developed ovaries (and which are less likely to successfully reproduce before colony breakdown) to assist their nestmates and thereby gain inclusive fitness. Revealing the role of Dufours gland secretion as a fertility signal adds another dimension to our understanding of how queen pheromones operate. The mandibular-gland secretion is a good predictor of dominance hierarchy, being correlated with false-queen characteristics but not fertility, whereas Dufours gland secretion is a good predictor of fertility but not dominance hierarchy.Communicated by R.F.A. Moritz