Environmental variation mediates the deleterious effects of Coleosporium ipomoeae on Ipomoea purpurea

Variation in the environment is common within and between natural populations and may influence selection on plant resistance by altering the level of damage or the fitness consequences of damage from plant enemies. While much is known about how environmental variation influences the amount of damage a plant experiences, few studies have attempted to determine how variation in the environment may alter the fitness consequences of damage, particularly in plant-pathogen interactions. In this work we manipulated a rust pathogen, Coleosporium ipomoeae, in field experiments and showed that this pathogen reduced several components of fitness in its natural host plant, Ipomoea purpurea. Furthermore, we showed that the deleterious effects of C. ipomoeae were variable. We identified variation in the quality of a plant's microenvironment, the abundance of secondary enemy damage, and the length of a growing season as variable components of the environment that may influence the magnitude of damage and tolerance, causing the interaction between C. ipomoeae and I. purpurea to vary from parasitism to commensalism. Considering how environmental variation impacts the magnitude and negative fitness effects of pathogen damage is important to understanding spatially variable selection and coevolution in this and other plant-pathogen interactions.     

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.     

Costing reproduction: effects of mating opportunity on mating success in male Bicyclus anynana butterflies

It is frequently assumed that males have an almost unlimited reproductive capacity, while access to receptive females is typically limiting. Consequently, sexual selection is expected to favor vigorous courtship behavior in males. If such behavior is associated with non-trivial costs, ample current mating opportunities should be accompanied by a reduction in future mating vigor. To test this hypothesis, three treatments differing in sex ratio were established using the tropical butterfly Bicyclus anynana: 50 males each were housed either with 0, 25, or 50 females. Mating trials involving the competition among males from each treatment for a single virgin female were carried out on days 3, 9, and 15 after allocation to treatments. While there was no difference on day 3, prior mating opportunity clearly reduced mating success on days 9 and 15, being lowest if identical numbers of males and females were housed together. This finding suggests accumulating costs associated with high courtship and/or mating activity. Further, older males were more active and initiated copulation earlier than the younger ones, consistent with the residual reproductive value hypothesis. We found no evidence for a survival or fertility cost of mating.     

Optimality of parental investment: The influence of nursing on reproductive success of mother and female young house mice

Summary The amount of parental milk investment determines not only the number of young the parent can produce, but also affects the offspring's fitness. The antagonism between quantity and quality of offspring was investigated in laboratory mice.In nursing first litters containing 2,4, ..., 14 pups, mothers invested increasing amounts of milk. This mainly extended the intervals between their first and second litters and slightly increased the size of their second litters, whilst third litters were not affected. In the female young, the decreasing amount each individual received as a result of increasing litter size led to delayed birth and reduced the size of their first litters. The intervals between their first and second litters and the size of the second litters were also affected, although to a lesser extent.Taking these results into consideration on a standard lifetime pattern of reproduction, the effects were calculated on net reproductive rates as an expression of fitness. Death rates were assumed that referred to population equilibrium or population growth. In both situations, the fitness curves of mothers and young showed that the actual level of milk investment and the level yielding optimal reproduction were in agreement. There was no significant deviation towards increased investment, as might result from parent-offspring conflict.     

Influence of behavior and mating success on brood-specific contribution to fish recruitment in ponds

One source of uncertainty in predicting the response of populations to exploitation is individual differences within a population in both vulnerability to capture and contribution to population renewal. For species with parental care, individuals engaged in nesting behavior are often targeted for exploitation, but predicting outcomes of this nonrandom vulnerability will depend in part on an understanding of how parental traits are related to potential for brood contribution to the population. Variation in brood-specific contribution to recruitment of largemouth bass (Micropterus salmoides), a fish species with extended parental care, was quantified to determine if differences in mating success, parental care behaviors, and timing of reproduction influenced offspring recruitment. Dependence of these relationships on brood predation was tested in communities that differed in the presence of bluegill, Lepomis macrochirus, an important nest predator. Daily snorkel surveys were conducted in experimental ponds during spring to monitor male spawning and parental care behaviors in populations of largemouth bass. Tissue samples collected from larvae in nests were used to develop brood-specific DNA fingerprints for determining nest origins of fall recruits. Largemouth bass spawning period in bluegill ponds was longer and more variable in duration, with lower, more variable mating success, than in ponds without bluegill. In all populations, only one or two broods provided the majority of recruits, and these were broods produced during the earliest days of spawning by the oldest, largest males. In bluegill ponds, brood contribution from earliest nests also increased with brood size. Earliest nesters were the oldest males, and recruits from these nests were often above average in body size. Offspring needed to be guarded to at least swim-up larval stage to contribute any recruits. Termination of parental protection before offspring were free swimming mainly occurred with broods guarded by smaller males in ponds with brood predators. These age- and size-specific differences in timing of spawning and duration of parental care are consistent with influences of residual reproductive value and energetic constraints on reproductive behavior. Furthermore, these patterns of individual contribution to recruitment imply that fisheries that selectively target either nesting individuals or larger, older males could potentially decrease recruitment at the population scale.     

Experienced males have higher mating success than virgin males despite fitness costs to females

It is generally assumed that mating history has potentially important effects on the mating biology of insects, and differences in mating success of males, in relation to their mating history, have been commonly documented in Lepidoptera. Mating success of male European corn borer Ostrinia nubilalis, in relation to their mating history, and consequent fitness parameters for their female mates, were investigated. Mating experience significantly increased mating success for O. nubilalis males. Mating with experienced males significantly reduced lifetime fecundity and longevity of females. Sperm depletion, transfer of less nutritive substances or more toxic substances may explain the female response to mating with experienced males. Male persistence or some other compensatory mechanism may account for female acceptance of experienced males.     

Mating system and mating success of the desert spider Agelenopsis aperta

Field studies of the desert spider Agelenopsis aperta revealed a primarily monogamous mating system. However polygyny, polyandry and polygynandry were superimposed upon the primary system, with 9% of the marked males and 11% of the marked females in a field population mating more than once. In the laboratory males commonly mated multiply with fertile offspring resulting, while females were less likely than males to mate multiply. Monogamy under field conditions was enforced by two factors: (1) high travel costs to males, and (2) a significant decline in female receptivity after the first mating. Heavy males were more likely to be accepted by females both in the field, and in female choice experiments conducted in the laboratory. Finally, male weight determined the outcome of male-male agonistic interactions over females. One possible explanation for female choice in this system which lacks male parental investment is that females may be using male size as an indicator of future success of their offspring.     

Bower decorations are good predictors of mating success in the spotted bowerbird

Variation in the mating success of males can often be predicted by considering measures of their exaggerated sexual traits. Male spotted bowerbirds Chlamydera maculata build and decorate elaborate structures--bowers--that function in mate choice. I show that numbers of certain decorations correlate with the mating success of the bower owner. Specifically, numbers of Solanum berries used as decorations, accurately predict variation in mating success over 2 years. I show a relationship between changes in rank berry number and changes in rank mating success. These predictive decorations differ from those shown to relate to mating success in another population of spotted bowerbirds, suggesting that the exact form of sexual display may differ between populations of the same species.     

Dietary carotenoids and male mating success in the guppy: an environmental component to female choice

Summary This study examined the relationship between dietary carotenoids, female choice, and male mating success in the guppy, Poecilia reticulata. Using a split-brood design, male siblings were either raised on a diet enhanced with astaxanthin and canthaxin or fed a basal diet without carotenoids. Males were photographed, and the location, size, and brightness of their red and orange pigment spots on the body were measured. Courtship behaviors were recorded during visual and mating trials. Males fed the carotenoid-enhanced diet had red and orange spots that averaged 2.5 times brighter, spent significantly more time near the female in visual response trials, were preferred by females in visual choice tests, and had a higher mating success than their siblings raised on the carotenoid-free diet. Diet did not affect male size, location or size of the red and orange pigment spots, or the intensity of courtship behavior. The results of this study show that females respond to environmentally-induced variation in the expression of a secondary sexual trait and that this has important consequences for male mating success.     

A radiotracer technique for the determination of male mating success in natural populations

Summary Males of a natural population of Antechinus stuartii (Marsupialia) were injected at the beginning of their short, synchronous breeding period with one of twelve gamma-emitting nuclides which are individually recognizable by their unique spectial properties. This label passed to the females during ejaculation and was identified when the females were captured and counted in a whole-body counter. This technique established mating success of males and provided an estimate of reproductive success.     

Sexual selection in lekking sage grouse: phenotypic correlates of male mating success

Summary Mate choice cues in sage grouse were reinvestigated by analyzing relationships between male mating success and a range of suggested cues. Display cues were implicated by significant relationships between mating status (whether or not a male mated) and lek attendance, display rate (corrected for effects of female proximity and time of day) and an acoustic component related to temporal and frequency measure of a whistle emltted during the strut display. Although display rate and the acoustic component were intercorrelated, both exerted significant partial effects on mating success in mutivariate analyses. These display measures also differed significantly between males. In contrast, mating success was not significantly related to measures of territory characteristics, including size and proximity to the lek center, or to body size. These results resolve discrepancies between previous studies and provide a basis for experimental analysis of the role of female choice in this lek system.     

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     

The genetic mating system of a sea spider with male-biased sexual size dimorphism: evidence for paternity skew despite random mating success

Male-biased size dimorphism is usually expected to evolve in taxa with intense male–male competition for mates, and it is hence associated with high variances in male mating success. Most species of pycnogonid sea spiders exhibit female-biased size dimorphism, and are notable among arthropods for having exclusive male parental care of embryos. Relatively little, however, is known about their natural history, breeding ecology, and mating systems. Here we first show that Ammothella biunguiculata, a small intertidal sea spider, exhibits male-biased size dimorphism. Moreover, we combine genetic parentage analysis with quantitative measures of sexual selection to show that male body size does not appear to be under directional selection. Simulations of random mating revealed that mate acquisition in this species is largely driven by chance factors, although actual paternity success is likely non-randomly distributed. Finally, the opportunity for sexual selection (I _s), an indirect metric for the potential strength of sexual selection, in A. biunguiculata males was less than half of that estimated in a sea spider with female-biased size dimorphism, suggesting the direction of size dimorphism may not be a reliable predictor of the intensity of sexual selection in this group. We highlight the suitability of pycnogonids as model systems for addressing questions relating parental investment and sexual selection, as well as the current lack of basic information on their natural history and breeding ecology.     

Correlates of male mating success in the ruff Philomachus pugnax,a lekking shorebird

Summary Male ruffs (Philomachus pugnax), a lekking shorebird, can generally be divided into two morphs according to behavior and plumage coloration. Predominantly dark-colored, independent male ruffs defend small territories on a lek, whereas lighter colored satellite males are non-territorial and less site-faithful to a particular lek. The traits associated with the mating success of independent male ruffs were studied at two leks during two seasons on the island of Öland, Sweden in May and June of 1985 and 1987. Using multivariate analyses, three characteristics were found to be significantly related to mating success: high frequency of visits by satellites to an independent male's residence, consistency of lek attendance, and low rates of aggressive behavior. In contrast, mating success was unrelated to the degree of darkness of the independent male nuptial plumage, territory location on the lek, or rate of courtship displays. The use of multivariate analyses helped to resolve conflicting conclusions from previous studies which employed simple statistical analyses, or none at all. Experimental manipulations are suggested which will help to further determine the influence of female mate choice in this lekking system.     

Fighting behaviour as a correlate of male mating success in black grouse Tetrao tetrix

Fighting is a fundamental determinant of male fitness in species where females prefer socially dominant males as mates or where dominants can prevent subordinates from mating. This in turn can lead to the evolution of honest inter- and intra-sexual cues of male dominance. Fighting as a behaviour comprises both fighting rate (number of fights per unit of time) and fighting performance (success in winning fights), but it is not always clear which of these components are important for female choice and how they link to signals of male quality. To quantify the relative importance of fighting as a cue for females, we recorded detailed behavioural data from male black grouse Tetrao tetrix at leks. We explored the relationship between phenotypic traits (body mass, eye comb size, tail (lyre) length and blue chroma colouration) and fighting performance and rates and how these were related to male mating success. In older males' pairwise fights, winners had lower blue chroma than losers, but there were no differences in other morphological traits. In yearlings, no morphological trait predicted success in pairwise contests. Both fighting rate and performance were positively related to the number of copulations acquired by a male; however, when controlled for lek centrality, fighting performance and not fighting rate was significantly related to mating success. Our results indicate that females may be using components of fighting behaviour as cues for mate choice.     

Breeding success of a brood parasite is associated with social mating status of its host

Reproductive success of brood parasites varies considerably both among and within host species, mainly due to differences in host egg-rejection rates and survival of parasitic chicks. Here, we investigated the breeding success of the cuckoo (Cuculus canorus) in one of its major hosts, the great reed warbler (Acrocephalus arundinaceus), with respect to host social mating status. In this passerine, polygynous males provide less parental care to their young per nest than monogamous males. Consequently, their less-assisted females may fledge lower numbers of nestlings than monogamous females. This may be especially true for secondary females, which often receive limited or no paternal help with young at all. Based on these findings, we expected higher cuckoo reproductive success in nests of socially monogamous than polygynous great reed warbler males. More specifically, we predicted lower fledging success of cuckoo young in nests of secondary than primary or monogamous females. In line with the prediction, we found higher cuckoo fledging success in nests of monogamous than polygynous males, monogamous nests being more than twice as successful as secondary nests. We detected, however, only a tendency to lower cuckoo success in primary compared to monogamous nests and no differences between primary and secondary nests. Moreover, neither parasitism nor host egg-rejection rates differed among the nests of different status. Our results show, for the first time, that the social mating status of a host may influence the overall reproductive success of a brood parasite and thus should be considered in further studies.