Dominance rank,female reproductive synchrony,and male reproductive skew in wild Assamese macaques

In groups with multiple males, direct mate competition may select for the evolution of dominance hierarchies that sort males into a queue for access to fertile females. The priority-of-access (PoA) model proposed by Altmann in 1962 makes explicit predictions about the resulting paternity distribution based on an interaction between male dominance rank and the overlap of female receptive phases. Here, we investigated whether the logic of the PoA model predicted the distribution of paternity across ranks in a seasonal breeder with high reproductive synchrony over six consecutive mating seasons. We studied 18 males that resided in a group of wild Assamese macaques (Macaca assamensis) in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand, between 2006 and 2011 with 5 to 13 conceptions per season. We assessed whether mate guarding increased paternity success, described “short-term” deviations from predicted paternity distribution, and examined how these are related to the number of competitors and fertile females. We determined genetic paternity of 43 (93 %) offspring born into the study group and found reproductive skew to be relatively low with 29 % alpha male paternity in accordance with the high degree of female reproductive synchrony observed. Short-term deviations from expected paternity distribution over ranks were not explained by the number of resident males or the number of conceiving females or their interaction. Within the limits of this study, these results suggest that even if males cannot discern female fertile phases, if reproduction is seasonal, and if reproductive synchrony is high, males may also compete directly over access to females.     

Sex-specific patterns of lifetime reproductive success in single and repeat breeding steelhead trout (Oncorhynchus mykiss)

All other things being equal, the lifetime reproductive success (LRS) of iteroparous and semelparous individuals should scale with the number of breeding seasons. Deviations from this relationship may occur for many reasons, including age- or size-related fecundity or life history trade-offs, which may differ between sexes. We used 19 brood years of DNA parentage analysis in a small (N = 4–143 year⁻¹) wild, unexploited population of steelhead trout (Oncorhynchus mykiss) to compare the LRS of individuals that spawned only once [“one time spawners” (OTS), N = 355 male, 371 female] to those spawning twice [“repeat spawners” (RPS), N = 13 male, 49 female]. Female RPS had nearly twice the LRS of female OTS (1.17 offspring per female vs 0.91 offspring per female), whereas male RPS had nearly three times the LRS of male OTS (1.54 offspring per male vs 0.57 offspring per male). Female RPS produced slightly more adult offspring during their second breeding season than their first (0.78 vs 0.82 offspring per female); however, male RPS produced all of their adult offspring in their second breeding season (0 vs 1.54 offspring per male). The additional growth in body size of males between breeding seasons may give them an advantage in their second breeding season, but the lack of offspring produced in their first season suggests a trade-off between survival and future reproduction that was not expressed in females.     

Survival and lifetime reproductive success in dispersing and resident red squirrels

Dispersal in red squirrels (Sciurus vulgaris) is not sex-biased and strict philopatry is rare. The immigration pattern suggests that nearly all animals have to disperse away from the natal site and that dispersal in this species is the outcome of local (intrasexual) competition. If this interpretation is correct, we predict that dispersers and residents, of both sexes, should have equal survival rates and lifetime reproductive success. Body mass, longevity, reproductive success and dominance rank of 34 resident offspring (settling within 400 m of the natal range) and 70 immigrants (dispersers) were compared. Immigrants did not weigh less than residents as adults, nor did they have a higher mortality during the pre-setding period. Survival rate, lifetime reproductive success (females) and the proportion of males obtaining a high dominance rank were similar for residents and dispersers, and no sex effect was found on either of the parameters studied. These results are consistent with the hypothesis that local competition determines whether an individual disperses further away or settles close to its birthplace.     

Maternal rank affects reproductive success of male Barbary macaques (Macaca sylvanus): evidence from DNA fingerprinting

Summary Paternity determination by oligonucleotide fingerprinting confirms that maternal rank affects the reproductive success of male Barbary macaques (Macaca sylvanus). High-born males began to reproduce significantly earlier and sired significantly more infants surviving to at least 1 year of age during the first 4 years of their reproductive career than low-born males. This relation was independent of the natal/non-natal status of the males, and was not affected by external conditions such as the level of intrasexual competition or the number of fertilizable females. Since high-ranking females in this population produced significantly more male offspring than low-ranking females, the data on sex ratio adjustment and comparative breeding success of sons and daughters are consistent with the predictions of the Trivers-Willard hypothesis. Offprint requests to: A. Paul     

Components of variance in male lifetime copulatory and reproductive success in a seed bug

Summary Males and females of the seed bug, Neacoryphus bicrucis Say, were individually numbered in the field in southeastern Georgia (USA) and census taken daily for 6 weeks. Variation in male mating efficiency (ME = no. copulations/no. sightings) exceeded that in females and was significantly greater than that generated by a null model. Lifetime copulatory success, estimated as the product of ME and longevity, ranged from 0 to 41, with ME accounting for over 88% of the variation. Lifetime reproductive success (LRS), estimated as the product of ME, longevity, and clutch size, ranged from 0 to 898. Among males copulating at least once, ME accounted for over 40% of the variation in LRS, while longevity alone or in combination with clutch size accounted for 21% and 46%, respectively, of the variation in LRS. Failure of some males (26%) to copulate contributed 38% of the total variation in LRS. Thus, among males surviving to adulthood, sexual selection pressure arising from variation in ME is approximately as strong a force for phenotypic change as is natural selection pressure arising from variation in longevity.     

Population trend alters the effects of maternal dominance rank on lifetime reproductive success in yellow baboons (<Emphasis Type="Italic">Papio cynocephalus</Emphasis>)

We evaluated the association between dominance rank and lifetime reproductive success of 75 free-ranging female baboons in Mikumi National Park, Tanzania. Data were evaluated over a 22-year period that included a period of troop increase (1975–1987) associated with two troop splits in 1978 and 1979, followed by a precipitous population crash (1987–1996) where the troops successively fused back together in 1989 and 1994. Lifetime reproductive success was significantly greater for high- versus low-ranking females when examined across the entire study period. High-ranking females had a longer reproductive life span (7.4 vs 3.6 years after first birth), reached menarche earlier (4.6 vs 5.2 years), lived longer (12.0 vs 8.8 years), and had more offspring of both sexes (2.25 vs 1.33 for male offspring; 3.25 vs 0.94 for female offspring), with four times the number of offspring of each sex surviving to 4 years of age compared to low ranking females. Greater offspring production was associated with shorter interbirth intervals of dominant versus subordinate females (545 vs 723 days), partly owing to lower miscarriage rates (0.05 vs 0.2) and a shorter duration of lactation (244 vs 330 days). Rank effects were then partitioned by mothers experiencing the majority of their reproductive life prior to, versus during, the population decline. The majority of rank effects on measures of lifetime reproductive success were virtually eliminated for mothers reproducing during the troop decline, indicating that the considerable impacts of social status on lifetime reproductive success can be markedly altered by intrinsically and extrinsically mediated demographic events.Ramon Rhine is deceasedCommunicated by C. Nunn     

Factors regulating annual mating success and reproductive success in spotted sandpipers (Actitis macularia)

Summary We investigated factors affecting annual mating success (MS) and reproductive success (RS) of spotted sandpipers (Actitis macularia) from 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. Analyses were restricted to ages 1–3. Mean annual female MS varied from 1.3 to 2.7 mates, and the MS pattern was indistinguishable from random. However, female MS increased with age and was affected by arrival date, territory size, and beach size. Female RS also increased with age, and number of mates and year effects were the most significant explanatory variables in each age. Older female RS was increased by priority on a territory and presence of a previous mate. Territory size and beach size varied with population density and did not predictably affect RS. The strong year affect on RS was associated with annual variation in sex ratio and predation. Males produce only one successful clutch per year, so MS greater than one is a result of nest loss and does not increase RS. Neither male MS nor RS changed with age. Male reproductive failure rate varied by year. Given that a male produced young, the degree of RS was affected by year, arrival date, priority on a territory, territory size, and beach size. In years with early-season predation, late arrivals had higher RS; territory and beach size effects varied by year. Neither the presence, nor degree, of female care was associated with male RS. Male RS was more subject to annual environmental variability than was female RS, probably because of relatively low annual potential RS among males.Offprint requests to: L.W. Oring at the current address     

Status,reproductive success and fitness in Cape mountain zebra (Equus zebra zebra)

Summary Demographic data relating to herd size and stability are given for a population of Cape mountain zebra (Equus zebra zebra) under longterm observation. Temporal dispersion patterns of male and female offspring differed and were independent of the mother's status. Dispersion in females appeared to be related to physiological state, and dispersion in both sexes was related to age rather than changes in parental behaviour. Reproductive success of dominant and subordinate mares was equal and independent of age and social and reproductive variables. Fitness of dominant mares, however, was significantly higher than that of subordinates, the latter having a higher foal mortality, part of which could be attributable to dominants' aggression. The fitness of all males born was 1.6:1 compared with all females. Dominant mares produced significantly more daughters than sons. This trend was not found for subordinates. Mother's status was positively correlated with dominant status in her female offspring but not related to the subsequent status of her sons. Daughters had a more than twice as great a chance of breeding than sons. For maximum fitness gains, therefore, dominant mares should produce more daughters, since a high proportion of these would also have high status and fitness. This tendency is reflected in the sex ratio skewed towards females found for dominant mares.     

Determinants of male reproductive success in wild long-tailed macaques (Macaca fascicularis)—male monopolisation, female mate choice or post-copulatory mechanisms?

One of the basic principles of sexual selection is that male reproductive success should be skewed towards strong males in species with anisogamous sex. Studies on primate multi-male groups, however, suggest that other factors than male fighting ability might also affect male reproductive success. The proximate mechanisms leading to paternity in multi-male primate groups still remain largely unknown since in most primate studies mating rather than reproductive success is measured. Furthermore, little research focuses on a female’s fertile phase. The aim of this study was to investigate the relative importance of male monopolisation and female direct mate choice for paternity determination. We also investigated the extent to which paternity was decided post-copulatory, i.e. within the female reproductive tract. We used a combined approach of behavioural observations with faecal hormone and genetic analysis for assessment of female cycle stage and paternity, respectively. The study was carried out on a group of wild long-tailed macaques living around the Ketambe Research Station, Gunung Leuser National Park, Indonesia. Our results suggest that both male monopolisation and post-copulatory mechanisms are the main determinants of male reproductive success, whereas female direct mate choice and alternative male reproductive strategies appear to be of little importance in this respect. Female cooperation may, however, have facilitated male monopolisation. Since paternity was restricted to alpha and beta males even when females mated with several males during the fertile phase, it seems that not only male monopolisation but also post-copulatory mechanisms may operate in favour of high-ranking males in long-tailed macaques, thus reinforcing the reproductive skew in this species.     

Female reproductive qualities affect male painted turtle (Chrysemys picta marginata) reproductive success

Male reproductive success (RS) in polygamous species with minimal social systems is often determined by the number of mates. However, because male RS is translated through females, the number of offspring sired can also be influenced by female qualities. Empirically sufficient data to document how tradeoffs between mate number and quality influence male RS are seldom available for long-lived, iteroparous species. We combined long-term life history data (1983–2006) on the E. S. George Reserve (ESGR, MI, USA) with parentage data from 155 clutches of 59 female painted turtles (Chrysemys picta marginata) of varying reproductive frequencies (2003–2006) to determine the relative contribution of female numbers and qualities on male RS. One previously documented trait of female painted turtles that can have substantial influences on male RS is repeat paternity through the use of stored sperm to fertilize over 95 % of within-year clutches. In addition, our study found that second-clutch producing female painted turtles on the ESGR have higher among-year reproductive frequencies than do first-clutch only females. Multiple paternity was detected in 14.1 % of clutches (min-max?=?6.1–30.0 % annually), and the number of mates of both sexes was low annually (males 1.0; females 1.2) and over 4 years (males 1.1; females 1.7). Among successful males, RS varied substantially (1–32 offspring) and was strongly influenced by the combination of female reproductive frequency and repeat paternity (>38 % among years), but not mate number. Low mate number for both sexes was unexpected in a species without complex mating behaviors or parental care.     

Comparative fecundity and reproductive strategies in seventeen species of the Cumacea (Crustacea: Peracarida)

The fecundity of 17 species of cumaceans belonging to 5 families from the eastern and western temperate regions of the Atlantic and the Canadian Arctic and Subarctic Oceans are discussed separately and then compared. The species range from 0.78 to 10.11 mm (mean carapace length) with fecundities ranging from 15.1 to 168.9 (mean number of eggs brood^-1 female^-1). Interspecific comparisons show that carapace volume versus volume of brood yields a very strong relationship (r=0.99). Therefore, it is a combination of carapace volume plus egg volume which determines the fecundity in the Cumacea. A discussion of the various life-history and reproductive strategies employed by the Cumacea is included.     

Female reproductive success and nest predation in polyterritorial wood warblers (Phylloscopus sibilatrix)

Summary Polyterritorial polygyny in passerine birds, i.e., when already-paired males attract females in secondary territories, has been explained by male deception. Secondary females are assumed to make a poor choice, because they lack information about males' pairing status. To date, the deception hypothesis has focused mainly on the hole-nesting pied flycatcher. In this study of polyterritorial wood warblers, an open-nesting species, secondary females did not differ in number of fledged young compared with females of monogamous males, and they fledged even more young than primary females. Nest predation influenced the allocation of paternal care. Almost half of the secondary females had exclusive male assistance because nests of primary females were preyed upon.     

Alliances in winter flocks of willow tits; effects of rank on survival and reproductive success in male-female associations

Summary The value of being dominant in winter flocks of willow tits may not only be a matter of personal survival for males, although their survival probability was highest among flock members. Winter flocks of willow tits contain male-female subunits, and the dominant pair may be viewed as an alliance based on mutual benefits. Dominant males provide their mates with protection in winter, and females return the help in reproductive success. Survival was directly related to rank only within sexes. Mates of dominant males survived significantly better than low-ranked males even though these females were subordinate to these males in direct interactions. Mates of dominant males were relatively spared from costs of low rank as they were subject to aggression less often than expected. Presumably dominant males prevented such attacks, as they excluded other flock members from high tree sections where their mates fed. Males should gain from improving the survival probability of their mates since it was not always possible to replace lost mates in spring populations having a male-biased sex ratio. Dominance in avian winter flocks may thus have more indirect effects than merely to improve the personal survival probability through resource priority. The benefits of being a dominant suggest that subordinate willow tits join groups because there is no space available for them to take up a territory as dominants.     

Helper contributions to reproductive success in the splendid fairy-wren (Malurus splendens)

Summary In Malurus splendens, helpers were present in 65% of 226 group-years with at least one helper female in 37% of group-years. Most females helped for only one year, while many males did so for at least two years. Most were offspring of one or both present breeders, and in 53% of helper-years, helped both parents. For 159 helpers of known age and parentage, the mean coefficient of relatedness to the offspring was 0.47. Novice females with or without helpers produced fewer fledglings per season than females with one year breeding experience and the same level of help. Helpers did not affect production of fledglings per year by females with one year of experience. Females with two or more years experience and at least two helpers produced more fledglings than equivalent birds with one or no helpers. Experience and helpers have little effect on production of fledglings per nest but they lead to more females renesting after a first brood has been raised. Fewer than 20% of novices renest after fledging one brood, while for females with at least two years experience, the percent renesting after success is 40% with no help, 56% with one helper and 69% with 2 or more helpers. Experienced females begin their first clutch earlier than novices, and helpers reduce the time to renest after success from 66 days for an experienced female with no helpers to 50 days for females with at least two years experience and two or more helpers. Breeding females with helpers survive better (76%) than those with no helpers (55%), and helpers thus gain future indirect fitness. Despite their close relatedness to breeders and offspring, in only 19% of group-years did helpers increase their indirect fitness from an increase in productivity.     

Space use,longevity, and reproductive success in meadow voles

Summary We addressed the question of how reproductive success (RS) was limited in the shortlived but highly fecund meadow vole, Microtus pennsylvanicus. In so doing, we asked how differential space use patterns could affect longevity and hence RS in each sex. The sample comprised all voles achieving sexual competency over the course of a 40-week breeding season in a live-trapped population in Manomet, MA USA. Matrilineal families were determined using a radionuclide labelling technique; paternity was estimated using a maximum likelihood model. Individual RS was defined as the number of offspring successfully recruited into the trappable population per adult. We found that the variance in RS among female meadow voles was greater than the variance among males. In an attempt to explain this pattern, reproductively successful individuals were compared to reproductively unsuccessful individuals with regard to survivorship, maximum body weight achieved, and spatial mobility. The only difference between fathers and reproductively unsuccessful males was that fathers were heavier. In contrast, mothers differed from unsuccessful females in every measurement. Females lived longer than males, and mothers lived longer than either fathers or reproductively unsuccessful females. The observed differences in longevity may have been largely the result of differences in levels of mobility, assuming more mobile voles were more susceptible to predation. Mothers were significantly more site tenacious than were either males or unsuccessful females. These patterns explain the distribution of RS in our population if predation differentially affects male and female meadow voles. The meadow vole is the only non-polyandrous vertebrate reported to date in which the variance in RS among females exceeds the variance in RS among males.     

Dominance rank,resource availability,and reproductive maturation in female savanna baboons

Summary Ten years of data collected from a population of savanna baboons, Papio cynocephalus anubis, residing near Gilgil, Kenya were analyzed to ascertain the extent to which social and ecological factors influence reproductive maturation in females. First sexual swelling occurred at an average age of 4.79 years and first birth occurred at an average age of 6.92 years. Age at first menses was significantly correlated with age at first sexual swelling, but age at first sexual swelling was not a good predictor of age at first birth. The amount of rainfall in the 6 months preceding first sexual swelling and resource availability were significantly correlated with age at first sexual swelling. When ecological factors were taken into account, dominant females had an earlier age at onset of puberty, but not an earlier age at first birth, than did subordinate females. We suggest that nutritional and social stress operate at the same physiological level to disrupt GnRH pulsatility and retard reproductive maturation in some females. Given that socioecological variables modify the timing of life history events related to fitness in female baboons, the task for the future is to unravel how socioecological factors influence different life history components and generate variation in lifetime reproductive success. Correspondence to: F.B. Bercovitch     

Correlates of male reproductive success in the browncheek blenny,Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

Summary Acanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.     

MHC, health, color, and reproductive success in sand lizards

Good genes are genetic elements that contribute to lifetime reproductive success, regardless of an individuals additional genotype. Their existence is debated, and most work has targeted their viability benefits to the offspring of choosy females. In the present study, we analyze a case of potential good genes effects in adult male sand lizards (Lacerta agilis). We show that males with a particular RFLP (Restriction Fragment Length Polymorphism) MHC genotype (O-males), as opposed to those that lack this genetic element (NO-males), have less ectoparasites under increasing physiological stress (indexed by baseline corticosterone level), and are not constrained by parasites at production of status coloration. Furthermore, O-males are more successful at mate acquisition and guard their partners longer. Ultimately, they have a higher genetic reproductive success as assigned by microsatellites.Communicated by W. Cooper