Behavioral attributes influence annual mating success more than morphological traits in male collared lizards

Traits that potentially influence mating success (MS) may be"static" with low lability once they develop or "dynamic" withhighly modifiable expression. We used principal components (PCs)analyses of dynamic behavioral and morphological traits thatare static over the short term to determine their relative contributionsto the ability of territorial male collared lizards to acquireaccess to females and obtain high MS. We estimated annual MSof males as the relative frequency with which they engaged incourtship with reproductively active resident females. ThreePCs explained statistically significant phenotypic variationamong males. Morphological traits loaded significantly on 2PCs that explained 26.3% and 13.0% of the variance, respectively,whereas behavioral variables loaded significantly on a differentcomponent that explained 15.7% of the variance in male traits.The frequency with which males initiated aggressive encounterswith same-sex competitors did not load significantly on thesePC axes. Males having behavioral PC scores above the mean hadsignificantly higher MS than those with behavioral scores belowthe mean, whereas male MS was not related to component scoresfor either of the axes describing static morphological variables.Results indicate that in our population behavior patterns associatedwith advertisement, particularly to females, influence maleMS more strongly than morphological traits or the initiationof direct aggression with same-sex competitors.     

Sperm investment and alternative mating tactics in bluegill sunfish (Lepomis macrochirus)

Although alternative mating tactics are found in males of manyspecies, little is known about tactic-specific adaptations tosperm competition and the mechanism by which fertilization successis obtained. We now report on the sperm investment patternsof males that use alternative mating tactics in bluegill sunfish(Lepomis macrochirus). Bluegill males are characterized by threealternative mating tactics: parentals, sneakers, and satellites.Parentals delay maturation and adopt a courting and guardingtactic, whereas sneakers and satellites mature precociouslyand, respectively, use sneaking and female mimicry to stealfertilizations from parentals. We found differences among thetactics in testes weight, sperm longevity, and ejaculate spermdensity and competitiveness. Parentals had the largest testes,but the smallest relative to their body weight. Satellites hadintermediate-sized testes, and sneakers had the smallest testes,but the largest relative to their body weight. Within each tactic,there was a positive relationship between testes weight andbody weight, but the exact relationship differed among the tacticsand could not be attributed solely to allometry. Parentals hadlonger-lived sperm compared with that of both sneakers and satellites.Ejaculate sperm density was greatest in sneakers, satelliteswere intermediate, and parentals had the lowest. Competitionexperiments involving equal volumes of ejaculate showed thatfertilization success increased linearly with sperm number,consistent with a rafflelike process. However, independent ofsperm number, parental sperm were superior at fertilizing eggs.We interpret these sperm investment patterns in relation todifferences in sperm competition risk, number of mating opportunities,and alternative investment options.     

Sex-specific life history patterns in bluegill (Lepomis macrochirus): interacting mechanisms influence individual body size

The ultimate body size that an individual fish achieves can be a function both of direct effects of growth or indirect effects associated with the timing of sexual maturation (and associated energetic tradeoffs). These alternatives are often invoked to explain variation in body size within and among fish populations, but have rarely been considered simultaneously. We assessed how resource availability and timing of maturation interact to influence individual body size of bluegill (Lepomis macrochirus). Resource availability (high and low food) and the social structure of the population (presence or absence of large, mature males) were varied in experimental ponds. Food ration affected growth (larger fish in the high food treatments) and the social structure of the population affected timing of maturation (early maturation of males in the absence of large males). Treatment effects, however, were sex-specific; males responded to the social structure of the population and females were more responsive to resource availability. We also found individuals that became sexually mature were smaller than those that remained immature, although results were sex-specific and resource dependent. For males, individuals that matured were smaller when resources were limited; mature and immature females showed no difference in body size regardless of food ration. We show that both resource availability and the processes that control timing of maturation interact in sex-specific ways to influence body size of bluegill. These results suggest that a more robust explanation for variable body size requires consideration of sex-specific interactions between ecological (food and growth) and evolutionary (timing of maturation) mechanisms.     

Plasma levels of androgens and cortisol in relation to breeding behavior in parental male bluegill sunfish, Lepomis macrochirus

Like many teleosts, male bluegill sunfish (Lepomis macrochirus) provide sole parental care. To understand some of the proximate costs of parental care, we measured body condition and plasma levels of testosterone (T), 11-ketotestosterone (11KT) and cortisol in nesting bluegill males during pre-spawning, spawning and parental care stages. T and 11KT were at their highest mean levels during the pre-spawning period and decreased to lower levels early during the parental care period before rising again when the eggs hatched. Cortisol levels fluctuated across the breeding stages, but there was a noticeable increase from low levels on the day of spawning during the first 2 days of parental care when egg fanning is most intense. Levels of all hormones varied considerably among males, with androgen levels often correlating positively with a male's body condition. We also demonstrate, using a brood reduction experiment and repeated sampling of known individuals, that the presence of eggs affects hormone levels shortly after eggs hatch. Parental males in better body condition had higher levels of androgens during parental care. Males that were known to renest later in the season also had higher androgen levels and were in better body condition during the first nesting bout than males only known to have nested once. However, circulating levels of cortisol did not differ significantly between these groups. We discuss our findings in the context of proximate and ultimate costs of parental care and propose several reasons why elevated androgen levels may not be as incompatible with the expression of paternal care in male teleosts, as compared with avian and mammalian fathers.     

Origin and dispersal of bluegill sunfish, Lepomis macrochirus, in Japan and Korea

The bluegill sunfish, Lepomis macrochirus, is a notorious exotic species in many freshwater ecosystems, currently expanding its distribution worldwide. In 1960, a small group of bluegills captured in the Mississippi River at Guttenberg in Iowa were imported to Japan as a gift from the mayor of Chicago to the Japanese government. The offspring of these fish were released into the wild in Japan and also in Korea. Over 40 years after this first introduction, L. macrochirus now occupies all the freshwater ecosystems of both countries. We compared invading populations of L. macrochirus in Japan and Korea with native populations in the USA, using PCR-RFLP (polymerase chain reaction-restricted fragment length polymorphism) analyses of mitochondrial DNA, to estimate the origin and dispersal of L. macrochirus in Japan and Korea. Five haplotypes of mitochondrial DNA detected in Japanese and Korean populations completely coincided with the haplotypes of the Guttenberg population. Haplotype diversity of invading populations was shown to be highest in populations established in the 1960s, while genetic variability was lower in more recently established populations. Our results suggest that all L. macrochirus in Japan and Korea have originated from the 15 fish first introduced in 1960. Low haplotype diversity in newly established populations is probably due to genetic drift arising from repeated population bottlenecks, while the high similarity of haplotypes among neighbouring populations is considered to reflect the history of transplantation by humans.     

Discrimination of random dot texture patterns in bluegill sunfish,Lepomis macrochirus

In order to study the ability of fish to perceive and distinguish textures visually, bluegill sunfish (Lepomis macrochirus) were trained to discriminate between pairs of artificial texture patterns. Random dot patterns with different statistical dot distributions were presented to the fish as artificial texture patterns. The results indicate that bluegills have the ability to discriminate many pairs of patterns with different statistical features of dot distributions which have different appearance in texture. This suggests that texture could be one of the important visual features bluegill sunfish recognize and utilize.     

A field observation on the influence of aggressive behaviour on mating success in the European grayling

The biggest male grayling present on a spawning area was strongly territorial and developed more aggressive behaviours than smaller males. However, its spawning frequency was similar to that of Smaller males.     

No sex‐specific markers detected in bluegill sunfish Lepomis macrochirus by AFLP

The amplified fragment‐length polymorphism (AFLP) technique was used to identify sex‐specific markers in bluegill sunfish. A total of 12 835 loci were produced by 256 primer combinations, of which nine (0·73‰) exhibited presumed sex‐associated amplifications in the pooled samples; however, none of which revealed sex specificity upon individual evaluation.     

Determinants of male mating success in a natural population of a stream goby of the genus Rhinogobius

The mating success of individually identified males of a stream goby, Rhinogobius sp. CB (cross‐band type), was recorded over one breeding season. One to 3 year‐old males were active in mating, but 3 year‐old males, which accounted for 36% of the population, amounted to c . 70% of the total males guarding eggs in the nest. Three quarters of the breeding males had only one brooding cycle, but the others had two or three. All the latter males changed their nest sites between cycles within a riffle. In 1 and 2 year‐old males, the number of brooding cycles contributed more to the mating success than egg mass size in one brooding cycle. For mating success of 3 year‐old males, the egg mass size in one brooding cycle, which can be enhanced by spawning with a large female or multiple females, was as important as the number of brooding cycles. These male reproductive tactics could be attributed to the age‐related ability of nest construction and mate acquisition.     

Costs of mating competition limit male lifetime breeding success in polygynous mammals

Although differences in breeding lifespan are an important source of variation in male fitness, the factors affecting the breeding tenure of males have seldom been explored. Here, we use cross-species comparisons to investigate the correlates of breeding lifespan in male mammals. Our results show that male breeding lifespan depends on the extent of polygyny, which reflects the relative intensity of competition for access to females. Males have relatively short breeding tenure in species where individuals have the potential to monopolize mating with multiple females, and longer ones where individuals defend one female at a time. Male breeding tenure is also shorter in species in which females breed frequently than in those where females breed less frequently, suggesting that the costs of guarding females may contribute to limiting tenure length. As a consequence of these relationships, estimates of skew in male breeding success within seasons overestimate skew calculated across the lifetime and, in several polygynous species, variance in lifetime breeding success is not substantially higher in males than in females.     

Temporal variation in male traits, nesting aggregations and mating success in the peacock blenny

At the beginning of the breeding season male Salaria pavo that have eggs in their nests are larger, have more developed anal glands and less intense eye-spots and are located in breeding aggregations. These differences cease to occur from the peak of the breeding season (June—July) onwards. Two scenarios are presented that may explain these results: (1) smaller and younger males may begin to breed later devoting part of the warm season to growth; (2) females may cease to be selective as the nesting space begins to be saturated with eggs. These results raise one methodological and one conceptual question. The search for correlates of male reproductive success must cover different phases of the breeding season to capture the overall dynamics of the processes involved. The operational sex ratio for cavity-spawning fishes should take into account the availability of spawning sites rather than just counting the sexually mature members of each sex.     

Effects of fine‐scale genetic structure on male mating success in gynodioecious Beta vulgaris ssp. maritima

Plant mating systems are known to influence population genetic structure because pollen and seed dispersal are often spatially restricted. However, the reciprocal outcomes of population structure on the dynamics of polymorphic mating systems have received little attention. In gynodioecious sea beet (Beta vulgaris ssp. maritima), three sexual types co‐occur: females carrying a cytoplasmic male sterility (CMS) gene, hermaphrodites carrying a non‐CMS cytoplasm and restored hermaphrodites that carry CMS genes and nuclear restorer alleles. This study investigated the effects of fine‐scale genetic structure on male reproductive success of the two hermaphroditic forms. Our study population was strongly structured and characterized by contrasting local sex‐ratios. Pollen flow was constrained over short distances and depended on local plant density. Interestingly, restored hermaphrodites sired significantly more seedlings than non‐CMS hermaphrodites, despite the previous observation that the former produce pollen of lower quality than the latter. This result was explained by the higher frequency of females in the local vicinity of restored (CMS) hermaphrodites as compared to non‐CMS hermaphrodites. Population structure thus strongly influences individual fitness and may locally counteract the expected effects of selection, suggesting that understanding fine scale population processes is central to predicting the evolution of gender polymorphism in angiosperms.     

Hemolymph loss during nuptial feeding constrains male mating success in sagebrush crickets

Although costs of mating have been widely documented in females,intrinsic costs of copulation have been poorly documented inmales, and there is little evidence that such costs constrainmale mating success under natural conditions. Male sagebrushcrickets, Cyphoderris strepitans, offer females an unusual somaticfood gift at copulation that may constitute a significant costof copulation: females chew on the ends of the males' fleshyhind wings and ingest hemolymph seeping from the wounds theyinflict. Previous studies have shown that once a male has mated,his probability of obtaining an additional copulation is reducedrelative to that of a virgin male seeking to secure his firstmating. If the future mating prospects of nonvirgin males arediminished because of the costs of copulation, this could stemeither from the resources required to manufacture a new spermatophoreor through the energy needed to replenish hemolymph lost throughfemale wing-feeding. To distinguish between these two alternatives,we experimentally depleted virgin males of varying amounts hemolymphin a way that mimicked hemolymph loss of nonvirgin males, withoutthe attendant costs of spermatophore production. After theyhad been treated, males were released in the field and recapturedover the course of the breeding season to monitor their matingsuccess. Control males mated significantly sooner than did malesdepleted of hemolymph. We conclude, therefore, that the depletionof hemolymph that occurs through female wing feeding is sufficientby itself to diminish a nonvirgin male's ability to secure anothermating.     

Male mating success in a North American pitviper: influence of body size,testosterone, and spatial metrics

Males with enhanced traits relative to conspecifics often show increased mating and reproductive success and thus have a fitness advantage. The opportunity or potential for sexual selection is predicted to occur under these conditions. Here, we investigated proximate determinants of mating success in male copperhead snakes (Agkistrodon contortrix), a medium‐sized pitviper of North America. Specifically, we investigated the relationships of body size (snout‐vent length, body mass), body condition index, spatial metrics (total distance moved, home range size), and plasma testosterone concentration on mating success in males. The single mating season lasts from August through September. We compared a set of candidate linear mixed models and selected the best‐fitting one using the adjusted Akaike Information Criterion (AICc). The AICc‐selected model (model 2), with testosterone, body condition index, and home range size as predictor variables, showed that male mating success was positively correlated with testosterone. To our knowledge, this is the first report to show the relationship of testosterone and individual mating success in any snake species. A parallel study conducted on male fitness in A. contortrix of the same population used microsatellite markers to assign parentage of fathers (known mothers). Unlike our study, they found that snout‐vent length was positively correlated with reproductive success and that males were experiencing greater sexual selection. This relationship has been detected under natural conditions in other species of snakes. Although behavioural data are important in any mating system analysis, they should not stand alone to infer parentage, relationships or selection metrics (e.g. Bateman gradients). Long‐term sperm storage by females, female cryptic choice, and other factors contribute to the complexity of mating success of males. Accordingly, we thus conclude that estimates of reproductive success and fitness in cryptic species, such as copperheads and other snakes, require robust molecular methods to draw accurate conclusions regarding proximate and evolutionary responses. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 185–194.     

Influence of previous mating experience on future mating success in maleLucilia cuprina (Diptera: Calliphoridae)

Receptive and virginL. cuprina females were placed with a virgin male and an experienced male that had mated between one and five times previously. The experienced males secured significantly more matings than virgin males. Males with previous matings also gained experience in competing with males. Males directed mating attempts at each other, seemingly in the context of intrasexual competition. Experienced males directed more mating attempts at virgin males than vice versa. As their number of previous matings increased, experienced males made the first mating attempt at females more often and directed more mating attempts at females compared with virgin males. Females did not actively discriminate against experienced males, even though the proportion of matings secured on the first attempt by experienced males declined with increasing mating experience. Alternative behavioral explanations are discussed.     

Body size,male aggression,and male mating success in the cottonwood borer,Plectrodera scalator (Coleoptera: Cerambycidae)

The natural history and mating system ofPlectrodera scalator exhibit several unusual characteristics. Larvae and adults feed on the wood and foliage, respectively, of the same plant,Populus deltoides. The population sex ratio, based on censuses of oviposition areas, is female-biased. Females are significantly larger than males, yet males are intensely aggressive. Larger males tend to win escalated battles, which involve grasping of antennae with mandibles, but smaller males can defeat larger males if they grasp their opponent's antenna first. Most escalated fights involve possession of a female, but prior possession does not play a role in determining the outcome of these fights. The size-dependent fighting advantage does not translate into a mating advantage for larger males. There is no significant difference in elytron length or body mass between mating and single males. Larger females are not preferred as mates. The mating system appears to be a mixture of female-defense and scramble-competition tactics. One advantage to males of aggression may be in its effect on sperm precedence. Males appear to be able to remove previously deposited sperm from a female's reproductive tract.     

An analysis of mating biases in trees

Assortative mating is a deviation from random mating based on phenotypic similarity. As it is much better studied in animals than in plants, we investigate for trees whether kinship of realized mating pairs deviates from what is expected from the set of potential mates and use this information to infer mating biases that may result from kin recognition and/or assortative mating. Our analysis covers 20 species of trees for which microsatellite data is available for adult populations (potential mates) as well as seed arrays. We test whether mean relatedness of observed mating pairs deviates from null expectations that only take pollen dispersal distances into account (estimated from the same data set). This allows the identification of elevated as well as reduced kinship among realized mating pairs, indicative of positive and negative assortative mating, respectively. The test is also able to distinguish elevated biparental inbreeding that occurs solely as a result of related pairs growing closer to each other from further assortativeness. Assortative mating in trees appears potentially common but not ubiquitous: nine data sets show mating bias with elevated inbreeding, nine do not deviate significantly from the null expectation, and two show mating bias with reduced inbreeding. While our data sets lack direct information on phenology, our investigation of the phenological literature for each species identifies flowering phenology as a potential driver of positive assortative mating (leading to elevated inbreeding) in trees. Since active kin recognition provides an alternative hypothesis for these patterns, we encourage further investigations on the processes and traits that influence mating patterns in trees.     

Effect of male and female age on the mating success of the obliquebanded leafrollerChoristoneura rosaceana (Lepidoptera: Tortricidae) under different ecological conditions

The present study was undertaken to test the hypothesis that male and female age, as well as temperature, can affect the mating success ofChoristoneura rosaceana, given the role that these two factors play in the pheromone biology of this species. In the laboratory, the mating success of females generally declined linearly with age, whereas in males, it increased during the first 3 days and then decreased. The decline in female mating success was more pronounced under warm than cool thermocycles, while the changes observed in males were greater under fluctuating than constant temperature regimes. The onset time of mating was unaffected by male age, however, older females always mated earlier than younger ones, with the advance being more pronounced at cool than warm temperatures. Similar results were also obtained under field conditions. When a single 3-day-old male was provided with 0-, 3-, and 5-day-old females simultaneously, older females obtained mates significantly more often than younger individuals at all temperatures, indicating that calling earlier may afford a reproductive advantage to older females. At both constant temperatures, the time spent mating was longer in older than in younger females, but not under fluctuating thermocycles. Very young and very old males generally spent more time in copula than middle-aged individuals both at constant temperatures and under the warm thermocycle. Under the cool thermocycle, the duration of mating was considerably prolonged at all ages, which could increase the risk of predation.