Solitary nesting as an alternative breeding tactic in colonial nesting bluegill sunfish (Lepomis macrochirus)

Colonial breeding can evolve in response to benefits afforded by clumped individuals, such as reduced predation and increased ease of assessing potential mates. However, colonial breeding can also impose costs such as increased disease transmission or increased cuckoldry. Here, we investigate solitary nesting as a potential alternative breeding tactic in colonial breeding bluegill sunfish (Lepomis macrochirus). Most male bluegill, termed parentals, compete for nesting sites in colonies and then court and spawn with females and provide sole care of the eggs. Although nesting in a colony results in reduced predation and fungal infection of broods, it comes at a cost of increased parasitism by specialized cuckolder males that do not nest. We found that 4.5% of parentals forgo spawning in a colony and instead construct nests solitarily. Solitary males were of similar size and age to colonial males, but were in significantly better condition. Solitary males obtained as many eggs as males nesting in the center of colonies, and significantly more than males nesting on the periphery of colonies. Thus, females do not appear to discriminate against solitary males. Solitary males had smaller ear tabs, a presumed sexually selected character used by parental males in intrasexual competition, than colonial males. Tracking data revealed consistency in nesting tactic (but not nest position within the colony) between spawning attempts. We suggest that solitary nesting represents either a facultative decision made by parental males in top condition at the onset of breeding, or a life history decision to forgo spawning in colonies.Communicated by K. Lindström     

Reproductive success in male savanna baboons

Summary Different techniques have been utilized to ascertain male savanna baboon reproductive success based upon behavioral data. A 19 month field study of the reproductive behavior of savanna baboons in Kenya revealed a high degree of concordance among five different measures of male baboon reproductive success. The number of ejaculations showed the highest correlation with time spent in consort. Male reproductive success was not correlated with the number of females mated with because most males mated with most females. Female baboons regularly undergo multiple cycles prior to conception and the penultimate cycle showed no behavioral or physiological differences from the conception cycle. In nearly one-third of conception cycles a single consort partner was responsible for almost two-thirds of ejaculations during the four day optimal conception period. One may be able to reasonably infer paternity in these cases, but the available data are insufficient to support the idea that the variance in male baboon reproductive success is greater than the variance in female baboon reproductive success. The variance in male savanna baboon reproductive success will remain uncertain until genetic paternity studies are undertaken. It is suggested that mate selectivity, longevity, and stochastic factors are important components influencing male baboon reproductive success.     

Application of various mathematical models to data from the uptake of methyl mercury in bluegill sunfish (Lepomis macrochirus)

Evaluated and compared are the results of various mathematical models to data from the uptake of methyl mercury in bluegill sunfish (Lepomis macrochirus). Data were obtained from the direct uptake of methyl mercury from water in bluegill at 9°C and at a CH₃HgCl concentration of 0.2 ppb. The models considered are: (1) linear uptake relationship; (2) first-order kinetics, one-compartment or single-exponential model; (3) combination of first-order kinetics and the linear uptake relationship, designated as slope-exponential model; (4) two-compartment or double-exponential model; (5) three-compartment model, including a storage compartment for the build up of contaminant, designated as storage model; (6) three-compartment model, allowing two routes of elimination, designated the “Y” model; (7) an empirical uptake relationship having the form of a rectangular hyperbola.A comparison of the nonlinear models showed that the empirical model had fewer convergence problems in obtaining parameter estimates and its parameter values had less variation. The best fit was made by the “Y” model, the empirical model gave one of the smaller values of the residual mean squares, and other nonlinear models had computational convergence difficulties. The lack of computational problems and the comparitively small residual mean squares resulted in the selection of and the preference for the empirical model for use in describing the individual and interactive effects of temperature and methyl mercury concentration on the direct uptake of methyl mercury from water by bluegill.     

Sperm characteristics associated with different male reproductive tactics in bluegills (Lepomis macrochirus)

We examined the availability and motility of sperm from parental and sneaker male bluegills (Lepomis macrochirus), a colonially nesting sunfish (Family Centrarchidae) with male parental care and a high incidence of cuckoldry by both sneaker and satellite males. We found no differences between sneakers and parentals in length and swimming speed of sperm, or percent and duration of sperm activity. In sneaker milt, however, sperm was almost 50% more concentrated than in parental milt (16.5×10⁶ vs 11.5×10⁶ sperm/μl of milt, respectively). Despite this difference in sperm concentration, stripped ejaculates from sneakers contained almost 400 million fewer sperm (only 32% as many sperm) than those from parentals due to their much smaller stripped ejaculate volumes (only about 19% that of parentals). Thus unless sneakers can compensate by releasing more sperm or gaining closer proximity to eggs at the time of spawning, they may be at a disadvantage with respect to sperm competition. We discuss these results in relation to models for the evolution of alternative reproductive behaviours in this species and suggest that the cuckolders may be making the best of a bad situation. Received: 18 February 2000 / Revised: 23 March 2000 / Accepted: 14 September 2000     

Reproductive success of monogamous and polygynous pied flycatchers (Ficedula hypoleuca)

One of the most interesting aspects of the mating system of pied flycatchers is the regular occurrence of polygyny. Here we present data on the reproductive success of polygynously paired pied flycatcher males compared to monogamous males based on paternity analyses through DNA fingerprinting. Males paired with two females suffered a higher loss in reproductive output per female compared to monogamous males due to (1) a greater proportion of unhatched eggs in their broods, (2) greater nestling mortality and (3) a greater probability of being cuckolded. Nevertheless, the number of fledglings was significantly greater for polygynous males. Based on the number of nestlings that returned for subsequent breeding seasons, however, the reproductive success of monogamous and polygynous males did not differ significantly. These data raise the question as to why males attempt polygyny. Received: 16 August 1999 / Received in revised form: 15 March 2000 / Accepted: 18 March 2000     

Reproductive success of solitary and communal Philoponella oweni (Araneae: Uloboridae)

Summary Philoponella oweni (Uloboridae) is a facultatively communal orb-web spider found in arid regions in the southwestern United States. Within one habitat solitary adult females and communal groups of adult females coexist.This study compared the reproductive output (egg production) and reproductive success (production of live offspring) of solitary and communal females. There was no significant difference in the mean number of egg-cases produced by solitary and communal females, but communal females produced significantly more eggs per egg-case. However, communal females suffered significantly higher rates of egg-case parasitism from the pteromalid wasp Arachnopteromalus dasys. As a result, the mean number of liver offspring produced by solitary and communal females did not differ. Thus, both strategies led to roughly equal reproductive success. The advantages of aggregation of adults may be significant in the evolution of social behavior in spiders.     

Determinants of male mating success in the red bishop (Euplectes orix)

We studied sexual selection in the red bishop, Euplectes orix, a colonial, polygynous weaverbird widely distributed over sub-Saharan Africa. Male reproductive success measured in terms of the number of nests accepted by females and the number of eggs and nestlings in all the nests on a male's territory varied considerably. The standardized variance (variance/mean²) in male reproductive success ranged from 0.505 to 1.737 in different years, indicating a high potential for sexual selection in this species. An analysis of genetic parentage for 432 nestlings by non-radioactive, multilocus DNA fingerprinting confirmed that male reproductive success (number of young sired on the territory) in this species can be reliably estimated by the measures introduced above. In all 4 study years there was a strong positive correlation between male mating success and the total number of nests that males built in their territories. The number of nests built can be partitioned into the number of weeks a male held a territory and his nest-building performance. Both factors exert a significant positive effect on male mating success and in combination explained between 53.3 and 86.3% of the variation in male reproductive success. Male morphological characters were found to be of no importance. Males that established a territory in the following season built more nests and held their territories for longer than males that did not establish a territory in the following season, suggesting that these measures might be indicators of male condition and quality. Male nest-building performance (number of nests built per week) seems to be unrelated to male condition or quality. Received: 8 January 1999 / Received in revised form: 7 June 1999 / Accepted: 13 June 1999     

Relative importance of male and territory quality in pairing success of male rock ptarmigan (Lagopus mutus)

We studied pairing success in male rock ptarmigan (Lagopus mutus) in northern Alaska to learn whether males obtaining more females possessed phenotypic traits that influenced female choice directly, whether these traits permitted males to obtain territories favored by females, or whether both processes occurred. The number of females per male varied from zero to three. Several male and territory traits were significantly correlated with number of females per male. We used multiple regression to obtain a single measure of male quality and a single measure of territory quality. These measures of male and territory quality correlated with each other and with male pairing success. We used path analysis to separate direct effects of male quality on pairing success from indirect effects due to high-quality males obtaining high-quality territories. Both direct and indirect pathways had significant effects on pairing success, and direct and indirect effects of male traits on pairing success were about equal. This study illustrates an analytical approach for estimating the relative importance of direct and indirect causal relationships in natural systems. Received: 13 January 1998 / Accepted after revision: 7 November 1998     

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.     

Individual and environmental determinants of reproductive success in male tree swallow (Tachycineta bicolor)

Evaluating the contribution of individual and environmental determinants of reproductive success is essential to improve our understanding of sexual selection. In socially monogamous bird species with high rates of extrapair paternity, traits or environmental contexts affecting the number of within-pair young (WPY) produced by males can differ from those affecting the number of extrapair young fathered (EPY). Here, we use a 4-year dataset collected in contrasted environments to assess the factors affecting male reproductive success in tree swallows (Tachycineta bicolor), a species showing high levels of extrapair paternity. Our analyses revealed that the number of WPY was higher under better environmental conditions, while the number of EPY was mainly related to male characteristics. Males nesting in more intensive agricultural areas had fewer WPY produced and a lower reproductive success. Also, males breeding earlier in the season had more WPY. The presence of parasites reduced males’ reproductive success, mainly by reducing the number of EPY. The influence of male phenotype varied according to population density: Tarsus length variation had a greater effect on reproductive success at low population density than at high density, while wing length was also positively related to the number of EPY, more so at high than at low density. Altogether, our results suggest a complex interplay between individual and environmental determinants of reproductive success and imply that sexual selection dynamics varies depending on environmental contexts.     

Correlates of male mating success in two species of Anthidium bees (Hymenoptera : Megachilidae)

Summary Field observations were made on the mating behavior of two congeneric species of solitary bees, Anthidium porterae in an arid grassland and A. palliventre in a coastal sand dune habitat. Males of both species exhibited resource defence polygyny and defended hostplants to gain access to females foraging for nectar and pollen. The mating frequencies of marked and measured resident (territorial) males were monitored during periods of continuous observation, following which measurements of territory size and floral resources were obtained. Mating success of A. palliventre males was strongly influenced by territory characteristics: Males that defended small areas with a few rich hostplant patches mated more often than males that held larger territories containing many hostplant patches of low floral density. Large males generally held high-quality sites and thus had a mating advantage over smaller individuals. In A. porterae, on the other hand, male mating success was unrelated to any measure of territory quality. Copulation frequency and male size were positively correlated, however, apparently due to the increased ability of large males to seize and hold females for mating. The two species also differed in the incidence of non-territorial, sneaky males. While absent in A. palliventre, sneaky males accounted for 12% of all mating observed in A. porterae. Males of A. porterae that displayed sneaky tactics mated, on average, as often as resident males. Offprint requests to: E.M. Villalobos     

Mating success on white-bearded manakin (Manacus manacus) leks: male characteristics and relatedness

Morphological, territorial, and behavioural characteristics were measured on white-bearded manakin (Manacus manacus) males from two leks in two consecutive years. These data were combined with data on marker-inferred relatedness to study possible co-variation with mating success. In one year, male size and male condition were correlated with mating success. In both years, males holding courts nearer the lek centre gained more matings. No observed male display behaviour appeared to be an independently important factor in explaining variance in male mating success. Successful males made more aggressive displays than non-successful males and more displays were between close relatives. Number of aggressive displays increased as the distance between male courts decreased. Mating success in the white-bearded manakin is most likely mediated by a combination of morphological and behavioural characteristics, influencing both male-male competition and female choice. Females could potentially use centrality on the lek as an indicator of male characteristics. However, levels of relatedness may influence spatial arrangement of males on a lek thereby affecting male-male interactions and ultimately influencing patterns of mating success.     

Egg mimicry and allopaternal care: two mate-attracting tactics by which nesting striped darter (Etheostoma virgatum) males enhance reproductive success

In a variety of fish species with paternal care of offspring, females prefer to spawn in nests that already contain eggs. This female preference has been hypothesized to explain egg thievery in male sticklebacks, allopaternal care of eggs in minnows, and the evolution of egg-mimicking body features in male cichlids and darters. Here we employ microsatellite-based parentage analyses to evaluate the reproductive success of striped darter (Etheostoma virgatum) males that appear to utilize two of these functionally related tactics to entice females to spawn in their nests. In an isolated population (Clear Creek, Ky.), we observed that breeding males develop conspicuous white spots on their pectoral fins. If these spots are egg mimics, as we suspect, then this represents the fourth independent evolutionary origin of egg mimicry documented to date in darters, the first based on pigmentation (as opposed to physical structures), and the first in which the egg mimics vary greatly in number among males. From direct counts of microsatellite genotypes in clutches of embryos, at least 3.8 females contributed to the progeny within a typical nest, and females tended to spawn preferentially with males that were larger and displayed more egg-mimic spots. In another population (Hurricane Creek, Tenn.) without egg mimics, the multi-locus genetic data document that allopaternal care is common, especially among the smallest males who sometimes tend nests containing their own as well as an earlier sire's offspring. Thus, these foster males had adopted egg-containing nests and then successfully spawned with subsequent females. Overall, the genetic data on paternity and maternity, in conjunction with field observations, suggest that egg mimicry and allopaternal care are two mate-attracting reproductive tactics employed by striped darter males to exploit female preferences for spawning in nests with 'eggs'. Electronic Publication     

Reproductive success increases with degree of kinship in cooperative coalitions of female red howler monkeys (Alouatta seniculus)

Evaluation of evolutionary mechanisms proposed to promote cooperative behavior depends on the relative influence of the behavior on the reproductive success of individuals, the reproductive success of the group in which they interact behaviorally, and the degree of gene correlation among cooperators. The genetic relationship within cooperative coalitions of female red howler monkeys was examined for three populations with different densities and growth rates. Patterns of gene correlation change within coalitions is documented using data from the mitochondrial and nuclear genomes, and long-term census monitoring. Differences in fecundity and infant survivorship within and between groups of unrelated (rˉ=0) and related (rˉ≥ 0.25) females are compared. Females that emigrate from their natal groups form coalitions with other migrant females. These coalitions attempt to establish a territory and, once successful in producing offspring, exclude other females from feeding resources. Females in these coalitions had different mtDNA haplotypes and a genetically estimated mean r of 0, supporting demographic data on emigration patterns indicating that these females rarely have the opportunity to form coalitions with kin. Patterns of recruitment and rate of matriline development within social groups supported behavioral data indicating that females actively attempt to promote their own matriline as breeders over that of other females, and that some matrilines are more successful at this than others. Mean r among females was significantly higher in coalitions established as social groups for several generations (rˉ=0.44). In these groups, females all shared the same mtDNA haplotype, and mtDNA haplotype divergence was significantly higher between than within groups. Females in coalitions with kin had significantly higher reproductive success than females in unrelated coalitions in all populations. This difference was not a function of coalition size, number of males, socionomic sex ratio, or primiparity, although anecdotal evidence suggests that allomothering may compensate for inept new mothers in related coalitions more often than in unrelated ones. Differences in territory quality could not be ruled out as a potential causal factor in the saturated populations, but were unlikely in the low-density, growing population. There were substantial differences among long-established coalitions in overall reproductive output in all three populations, and this was significantly correlated with the number of breeding females. Increase in coalition size was a function of both group age and the behavioral tolerance among females. Regardless of the underlying reasons for the patterns observed, reproductive success clearly increases with degree of gene correlation among females within cooperative coalitions, and coalitions that recruit more daughters produce more offspring. The nature of the cooperative relationship among group females directly influences both of these outcomes. This is associated with substantial genetic differentiation among social groups within populations, creating conditions in which genetic tendencies towards cooperative behavior can become tightly associated with group reproductive success. Received: 15 September 1999 / Revised: 27 April 2000 / Accepted: 27 May 2000     

High male reproductive success in a low-density Antarctic fur seal (Arctocephalus gazella) breeding colony

Understanding how population density influences mating systems may lead to important insights into the plasticity of breeding behavior, but few natural systems allow for such studies. Antarctic fur seals (Arctocephalus gazella) provide an interesting model system because they breed in colonies of varying densities. Previous studies have largely focused on a high-density site at Bird Island, South Georgia. Here, 13 highly polymorphic microsatellite loci were used to conduct a genetic analysis of a low-density breeding colony of this species at Livingston Island, approximately 1,600 km south of South Georgia. The majority of adults seen ashore (n?=?54) were sampled together with every pup born (n?=?97) over four consecutive years. Paternities were confidently assigned for 34 out of the 97 pups. Two out of 23 sampled males accounted for the paternity of 28 % of all pups sampled during the study and 82 % of the pups with an assigned father. Moreover, a full likelihood pedigree inference method assigned a further eight paternities to a single unsampled male seal that is inferred to have held a territory during the season before the study began. The most successful males in our study easily surpassed the previous record for the total number of pups sired per male seal for the species. Furthermore, we identified two triads of full siblings implying that their parents remated in three consecutive years. These findings suggest that territorial male fur seals may achieve greater success in monopolizing access to breeding females when population density is relatively low.     

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.     

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     

Reproductive success of parasitized males in a marine reef fish

Parasitism is hypothesized to reduce reproductive success in heavily parasitized males because females may preferentially mate with less parasitized males (parasite-mediated sexual selection) or parasites may compromise male competitiveness. In marine systems, this hypothesis is largely unexplored. This paper provides the first confirmed record of a copepod ectoparasite (Caligus buechlerae Hewitt 1964) on the common triplefin (Forsterygion lapillum) and evaluates the hypothesis that males parasitized with C. buechlerae experience lower reproductive success than unparasitized males (as determined by the presence and area of eggs within male nests). We found that 38 % of males we surveyed were infected with at least one C. buechlerae, with a median of two individuals per infected male. About 32 % of males were defending eggs, with 62.5 % of those males infected with at least one parasite. Males of greater total length (TL) were both more likely to be infected and more likely to be defending eggs. However, when statistically accounting for the effects of TL, parasite infection had no effect on the probability of defending eggs, or the average surface area of eggs when present. Positive covariation in fish length, the presence of eggs and parasite infection observed here potentially suggest that the importance of parasitic infection on reproductive success may depend upon the strength of selection for larger male body size. Our study is one of the few studies to investigate the effects of ectoparasites on reproductive success in reef fish and also provides a quantitative measure of infection for a widespread species within New Zealand.     

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.