Glucosephosphate isomerase genotype frequencies,homozygous excess and size relationships in the sand flounder Rhombosolea plebeia

Genetic variation was studied at two polymorphic loci for 3 yr (November 1978–February 1982) in juvenile sand flounder (Rhombosolea plebeia) in Wellington Harbour, New Zealand. The Gpi-1 locus showed a heterogeneity in genotype frequencies, but this was not clearly related to season or to fish length. At the Gpi-2 locus, genotype frequencies were dependent upon length and showed a significant excess of homozygotes in juveniles but not in adults. This latter phenomenon has been reported previously in several species of marine molluscs, and so may be common to marine organisms producing large numbers of pelagic larvae. Reasons for the excess of homozygotes are discussed, and it is suggested that assortative mating is the most likely explanation and that this is an area of marine population genetic research which has been over-looked.     

Male preference for large females and assortative mating for body size in the sand lizard (Lacerta agilis)

Summary Male sand lizards (Lacerta agilis) are polygynous and guard individual females for several hours to days after copulation. Even though the copulation itself only lasts 2–4 min, the total time that a male invests per female is considerably more and may constitute a substantial investment during a mating season. In such situations, when male copulation frequency is constrained, or when variation in female fecundity is high, mate choice by males may be adaptive. Large body size in female sand lizards is correlated with higher fecundity. In choice experiments performed in the laboratory, male sand lizards preferred to court large females rather than small females. In addition, when there was little difference in size between the females in the experiment, the males visited the two females more often before they started to court the preferred female. The results from a field study during 1984 and 1987–1990 showed that females are non-aggressive, have small neighboring home ranges (c. 100 m²) and may share burrows and sites for thermoregulation. This means that females can be found close together and thus gives males the opportunity to choose a mate. Assortative mating with respect to size was observed in a natural population, as well as a limited number of direct choices of females by males. These results support the results of the choice experiment.     

Plumage characteristics, reproductive investment and assortative mating in tree swallows Tachycineta bicolor

Elaborate ornamental plumage has been associated with various measures of individual quality in many species of birds. Male plumage characteristics, which have been relatively well studied, have been shown to reflect past reproductive investment, as well as the potential for reproductive investment in the current breeding attempt. In contrast, the signalling functions of female traits remain largely unexplored. In this study, we investigated the relationship between plumage attributes of breeding adult tree swallows and past reproductive investment, current reproductive investment and social mate pairing strategy. Both males and older females possess metallic green to metallic blue iridescent plumage on their dorsal surface, making this a suitable species for this type of investigation. We did not find any effects of past reproductive investment and success on the plumage attributes of returning breeders. In contrast, female plumage hue covaried with fledging success, and female plumage brightness was positively associated with mean clutch egg mass. In addition, we found that social pairs mated assortatively with respect to plumage brightness. We argue that since plumage characteristics vary with age in both male and female tree swallows, plumage attributes in this species are indicative of breeding experience and may be honest signals of quality. Positive assortative pairing could be the result of mutual mate choice or intra-sexual competition for nest sites by both males and females.     

Age and growth of co-occurring larvae of two flounder species,Rhombosolea tapirina and Ammotretis rostratus

Daily growth increments on otoliths were used to age larvae of the pleuronectid fluonders Rhombosolea tapirina Günther and Ammotretis rostratus Günther, collected from Port Phillip Bay, Victoria, Australia, in winter 1984. Daily formation of growth increments was confirmed for R. tapirina by examining the growth of the marginal increment on otoliths of larvae collected over two 24h periods in winter 1985. The first distinctive growth increment was laid down approximately 5 d after hatching, at the onset of external feeding. Growth of flounder larvae was exponential from an early feeding stage to notochord flexion at approximately 30 d after hatching. The specific growth rate was very similar for the two species, at slightly over 4% of standard length per day. Predicted absolute growth rate of R. tapirina larvae increased from approximately 0.10 mm d^-1 in early feeding larvae to approximately 0.23 mm d^-1 in flexion-stage larvae, compared with 0.12 to 0.28 mm d^-1 for A. rostratus larvae of equivalent ages. Exponential models did not adequately describe growth of first-feeding larvae, which was slower than predicted. Growth in the field was faster than that recorded for the same species in the laboratory at higher water temperatures and prey abundances. Otolith growth accelerated markedly in relation to growth in length at the beginning of metamorphosis, causing a significant alteration in the morphology of growth increments, and eventually leading to the cessation of production of visible increments.     

Analysis of the causal components of assortative mating in water striders

Field studies demonstrate that natural populations of a group of water striders (Heteroptera: Gerridae) that share a common mating system are characterized by weak assortative mating by size and by large sizes of mating males and females relative to single individuals. This study presents an experimental assessment of the components of mating that may contribute to these mating patterns. The effects of male and female body size on each of three components of mating were studied in three water strider species in the laboratory. Large females of all three species mated more frequently, copulated for longer and were guarded longer than small females. Large males mated more frequently than small males in all three species, and also guarded females for longer in the two species where the average of mate guarding was long. However, we found an antagonistic effect of male size on copulation duration: small males copulated for longer than large males in all three species. We show that the combined effects of these size biases mimic the mating patterns found in the wild, e.g. weak and variable assortative mating, and stronger and less variable size ratios of mating versus non-mating females relative to males. We suggest that the antagonistic effects of male size on copulation and guarding duration may be a key source of interpopulational variation in assortative mating and sexual selection on male size. Further, neither spatial or temporal covariation in size, nor mechanical constraints, caused the assortative mating observed here in this group of water striders. Some combination of male and female choice (either active or passive forms) of large mates and male-male exploitation competition for mates play potentially important roles in producing population level assortative mating in water striders. Received: 17 March 1995/Accepted after revision: 28 October 1995     

Phenotypic assortative mating and genetic population structure in the crab Trapezia digitalis

Colonies of Pocillopora spp. containing the crab Trapezia digitalis were collected from Hawaii in summer of 1979 and of 1980. Individuals of T. digitalis in Hawaii vary extensively in coloration of the carapace: crabs were separated into phenotypic classes by their degree of carapace reticulation. When presented with two females of different phenotype, males preferentially paired with the female which was similar to them in reticulation. The genetic population structure of T. digitalis was unusual by comparison with other species of Trapezia. Previously reported homozygote excesses at esterase-4, hexokinase-1, phosphoglucoseisomerase, and general protein-2 cannot be explained by the single-locus Wahlund effect, since there was not significant genetic differentiation among phenotypic classes, as measured by Wright's F _ST statistic. There was significant variation among classes in allele frequencies at a fifth locus, hexokinase-2. Two pairs of loci were in significant gametic disequilibrium. Gametic disequilibrium was not detected in seven other species of Trapezia. It is suggested that multi-locus genetic analysis and an increased geographic sampling range are needed to determine the genetic population structure of T. digitalis in relation to observed variation in coloration.     

Male choice,fighting ability,assortative mating and the intensity of sexual selection in the milkweed longhorn beetle,Tetraopes tetraophthalmus (Coleoptera,Cerambycidae)

Summary Over five hundred adult longhorn milkweed beetles, Tetraopes tetraophthalmus, were individually marked and their copulatory success followed for one month in a pasture of Asclepias syriaca in northern Indiana, USA. Migration of beetles from the field site was greatest from areas of low population density. Dispersal was significantly greater for males experiencing low copulatory success; a similar but nonsignificant trend was observed for females. Large males, which displayed greater site tenacity than small males, copulated more frequently than small males because of their ability to displace small males from females. Both large and small males demonstrated a preference for large females in laboratory tests. Male preference in combination with aggressive displacement of small males results in size-assortative mating which was much stronger under conditions of high population density. It contributes to variance in male reproductive success since female size is known to be correlated with fecundity and offspring viability. Variance in copulatory success is similar for males and females, suggesting that both sexes experience similar intensities of sexual selection with respect to this component of reproductive success. Futhermore, comparison of this with other studies suggests that the intensity of sexual selection among males is positively correlated with the variance in body size which appears to be under both stabilizing and directional sexual selection in males but not in females.     

Female spawning patterns and male mating success in the sand goby Pomatoschistus minutus

In June 1989 in a study conducted near Träminne Zoological Station, Finland (60° N 23° E) I investigated whether or not male mating success could be explained by female choice for male size in sand gobies (Pomatoschistus minutus). Male mating success was constrained by nest size and increased markedly with increasing nest size. I also found a negative correlation between the length of spawning females and the fullness of the nest. As large females lay more eggs, they also need to find a nest with more space available for the eggs. The size of males without eggs was the same across nest size, whereas the size of males with eggs significantly increased with increasing nest size. This is interpreted as female discrimination against males as mates in nests that are often contested. There was no correlation between a male's size and his mating success when males with no eggs in their nests were excluded. A male removal experiment, however, showed that, in a specific nest, when male size increases so does mating success, whereas, if male size decreases, mating success also decreases. It is concluded that in the sand goby females prefer to mate with larger males, especially when the male possesses a high-quality nest that he most probably will have to defend against other males.     

Repeatable female preferences, mating order and mating success in the poeciliid fish, Heterandria formosa

Sexual selection via female choice can afford preferred males comparably higher mating success than those males that lack preferred traits. In addition, many models of sexual selection assume that both male traits and female preferences are heritable. In this study we test whether females of the poeciliid fish, Heterandria formosa, have repeatable pre-copulatory preferences for larger males. We also test whether female pre-copulatory preferences are always reliable indicators of male mating success. When given a choice between a large and a small male, females prefer larger males, and the repeatability of this preference is high. Although there are no overall differences in male mating success between large and small males, large males have a higher mating success when they are the first to mate than when they are the second to mate. Likewise, preferred males also have higher mating success when they are the first to mate than when they mate second. Therefore, the repeatable female preferences observed in this study only predict male mating success when the preferred male mates first. These results illustrate that even significantly repeatable female preferences do not translate into male mating success, which is an assumption of many examinations of the importance of female choice in sexual selection.     

Male size,mating success,and mating strategy in the mosquitofish Gambusia affinis (Poeciliidae)

Summary In laboratory choice experiments, receptive female western mosquitofish Gambusia affinis affinis (deprived of contact with males 30 days) preferred the larger of two males. When two males differing in size were placed with a receptive female, the larger was generally able to monopolize access to her, but not when the female was not receptive. In other experiments, a single male was placed with a receptive or non-receptive female. These experiments showed size-correlated differences in male mating behavior. Smaller males were more likely to chase non-receptive females and attempt to inseminate them forcibly, while larger males continued to court females even if they were non-receptive. Even when the female was receptive, there was some evidence of size-correlated differences in levels of courtship and chasing.     

Supply and demand: causes and consequences of assortative mating in common terns <Emphasis Type="Italic">Sterna hirundo</Emphasis>

Although assortative mating is widespread among long-lived monogamous birds, the underlying mechanisms still remain unclear in many species. In this study, we analysed assortative mating by age and body mass in the common tern Sterna hirundo with special regard to mate choice decisions of newly paired birds. To assess whether assortative mating by age is an active decision involving either homotypic or directional preferences or rather a passive process due to restricted availability of potential mates, we analysed the influence of age-dependent arrival date and cohort size. Furthermore, we looked for direct benefits in terms of reproductive success. Common terns mated assortatively by age and arrival date but not by body mass at arrival. Assortative mating by age was age dependent and was detected mainly in younger birds, whereas birds older than 8 years rather re-paired with younger ones. The availability of same-aged mates was restricted by cohort size and arrival date. We found no general evidence for better reproductive performance of same-aged pairs. Instead, relative reproductive success was related to own age and the relative age of mate: It should be advantageous for any bird to acquire an old mate; hence, assortative mating by age seems to be beneficial only for old terns but not for young ones. Age-assortative mating in common terns occurred by both passive and active processes, which are not mutually exclusive. Our results do not indicate a homotypic but a directional preference and support the theory that high-quality (older/experienced) individuals tend to mate assortatively if same-aged mates are available, which leaves low-quality (young/inexperienced) individuals to mate among themselves.     

Dominance, access to colonies, and queues for mating opportunities by male boat-tailed grackles

Species characterized by female-defense polygyny have extreme variance in male mating success. Many studies have considered alternative male strategies for access to females, but few have considered age-specific strategies. Male boat-tailed grackles (Quiscalus major) compete for access to colonies of females and form linear dominance hierarchies. I observed two groups of males that competed for females at seven colonies. Dominance rank was significantly correlated with mass, but not after controlling for age. In contrast, the correlation between dominance rank and age remained significant after controlling for mass. Older males dominated younger males and dominance relationships were very stable. Thus, dominance hierarchies represent queues for mating opportunities. A male's rank in the hierarchy determined how closely he approached a colony. Furthermore, males of all ranks prevented lower-ranked individuals from approaching the colonies. Dominance rank thus determined access to nesting females. One top-ranking male's loss of mass over the course of the breeding season presumably reflected the energetic cost of defending females, but he maintained his position in the hierarchy despite the small loss of mass. One alpha male held a colony for at least 4 years, and the ages of males from two queues indicated that males wait 6 or more years before becoming an alpha male. Therefore, most males die before acquiring a colony of females. Spatial structure such as that documented here could obscure recognition of queues and explain why they have not been documented in more species. Received: 13 May 1996 / Accepted after revision: 26 April 1997     

Predation risk,food deprivation and non-random mating by size in the stream water strider,Aquarius remigis

Summary Non-random mating by size (NRMS) plays a central role in the study of sexual selection and the evolution of mating systems. Theory suggests that NRMS should be influenced by conflicting demands (e.g., predation risk, hunger); few experimental studies, however, have addressed these effects. We used a factorial experiment to examine the influence of predatory green sunfish and food deprivation on NRMS in male and female stream water striders, Aquarius remigis. As predicted by theory, food deprivation decreased the large-male mating advantage. The influence of predation risk, however, went against existing theory; that is, risk increased the large male mating advantage. The degree of large-male mating advantage was negatively related to a measure of the rate of male harassment of females. A behavioral mechanism that can explain these patterns emphasizes the contrasting effects of different competing demands on male harassment rates, female resistance and the role of male size in overcoming female resistance. Females usually resist male mating attempts. Successful mating occurs when males overcome female resistance. If harassment rates (of females by males) are low, larger males have a mating advantage over smaller males perhaps because females resist heavily and thus only larger males can overcome female resistance. If, however, male harassment rates are very high, female resistance might be swamped; mating should then be more random with respect to male size. Food deprivation increases gerrid activity and thus increases harassment rates which should then reduce NRMS. In contrast, risk decreases gerrid activity, thus decreasing harassment rates and increasing NRMS. Females did not show significant NRMS. Females did, however, show a pattern of change in NRMS that is consistent with male choice for larger females. Correspondence to: A. Sih     

Multiple sexual pigments,assortative social pairing,and genetic paternity in the yellow warbler (Setophaga petechia)

In socially monogamous species, extra-pair paternity may increase the reproductive success of highly ornamented males, mediating the evolution of sexual ornaments. However, ornaments may also attract social mates, and a tradeoff between extra-pair paternity (EPP) and within-pair paternity (WPP) may complicate mating strategies. Further, in many socially monogamous species, females are also ornamented, and the relationship between female ornamentation and patterns of EPP has been neglected. We investigated the patterns of genetic paternity with respect to carotenoid- and melanin-based pigmentation in yellow warblers (Setophaga petechia) of both sexes. We asked whether males face tradeoffs between EPP and WPP, how paternity patterns relate to carotenoid- versus melanin-based pigmentation, and whether less EPP occurs in broods when males and females are assortatively paired. Males faced a tradeoff between EPP and WPP. Moreover, non-additive relationships existed between paternity patterns and the two pigment types in both sexes. Males with high melanin coverage but dull carotenoid pigmentation achieved EPP but lost WPP, whereas males with high levels of both pigment types had high WPP but gained little EPP. A parallel pattern occurred in females. Warblers paired assortatively by pigmentation and EPP was less common in broods when the sexes were assortatively paired by carotenoid pigmentation. Results suggest that the most colorful birds obtain high quality social mates and advance reproductive success through WPP, show that correlations can arise between female ornamentation and patterns of EPP, and also uniquely suggest that social pairing patterns may influence extra-pair mating strategies.     

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     

Sex ratio, sexual dimorphism and mating structure in bethylid wasps

Sexual dimorphism has been linked to parasitoid mating structure by several authors. In turn mating structure has an important influence on predicted sex ratio optima. Here we test the relationship between sexual dimorphism and sex ratio using data from 19 species of bethylid wasps. Using phylogenetically based comparative methods we confirm the findings of a previous cross-species analysis that sex ratio (proportion of males) is strongly and negatively correlated with clutch size. Using cross-species comparisons we show an additional positive correlation of sex ratio and relative male size, as predicted. The relationship however is not significant when using phylogenetically based methods. The cross-species result is largely due to differences between two bethylid sub-families: the Epyrinae have relatively large males and relatively high sex ratios, whereas the Bethylinae have relatively small males and lower sex ratios. Our study illustrates the benefits and drawbacks of using cross-species versus phylogenetically based comparisons. Received: 13 May 1997 / Accepted after revision: 12 January 1998     

Polyandry, multiple mating, and female fitness in a water strider Aquarius paludum

Contrary to Bateman’s principle, polyandry appears to be a common female mating strategy. Several hypotheses have been proposed to explain the evolution of polyandry. It is assumed that females gain either material or genetic benefits from multiple matings, or that they are coerced into mating by males. In water striders, mating is generally assumed to be costly to females, and they are thought to mate for reasons of convenience, adjusting their resistance to mating according to male harassment. Here, we tested the effect of number of matings (with the same male) and number of partners on female fitness in a water strider Aquarius paludum. In the first experiment, we regulated the time females spent with a male and found that females’ egg production increased with multiple matings up to a point. The result supports the existence of an optimal female mating frequency. In the second experiment, we tested how polyandry affects the number of eggs laid and egg hatching success. We conducted three different trials: females mated four times with either a single male, two different males, or with four different males. Females that mated with four different males laid the lowest number of eggs and had the lowest egg hatching success, suggesting that polyandry reduces females’ egg production and egg hatching success in A. paludum. We conclude that A. paludum females probably gain material benefits from mating but no genetic benefits were found in this study.