Loading constraints sexual selection and assortative mating in peracarid Crustacea

In precopula pairs of amphipod and isopod Crustacea in which males carry females, the males are larger than their mates and mating is size-assortative. Mate-guarding is a product of sexual selection. Size dimorphism and assortative mating have also been attributed to sexual selection but the supporting evidence for amphipods is equivocal. We describe a series of experiments confirming that relatively large male Gammarus pulex L. have an advantage because they can swim against stronger currents when carrying a mate. At higher current speeds, the male/female size ratio which forms is significantly greater, and in field collections size ratios of pairs are higher in streams than in lakes for a number of species. In a simulation we show that a size-assortative pattern inevitably develops if the observed size restriction is used as a rule for pairing. The results are discussed with respect to size-assortative mating, which has been attributed to male selectivity and male-male competition for access to large, fecund females.     

Size-related mating success in the shore crab Carcinus maenas (Crustacea: Brachyura)

The relationship between the size of a given mating male Carcinus maenas (L.) (Brachyura, Portunidae) and the size of the female with which it was paired was studied for 1248 pairs of crabs collected from the shore In 764 of these pairs the female was in pre-moult and so the pair were in pre-copula. In the remaining 484 pairs the female had already moulted and the pairs were in copula. There were significant correlations between the sizes of the males and females in both pre-copula and copula pairs. It was found that male Carcinus collected in mating pairs and tested in the laboratory were unable to distinguish between females in terms of their size or stage of pre-moult. The positive correlation between the sizes of males and females in mating pairs on the shore is proposed to be, in part, a function of a mechanical constraint of the size of female that a given male can hold, defend and copulate with. In addition, encounters between solitary males and males carrying females, resulting in the formation of new pairs, appear to enhance the size-related mating pattern observed.     

Factors affecting the microdistribution of Gammarus pulex (Amphipoda): an experimental study

SUMMARY. 1. The tnierodistribution of Gammarus species is size assortative: larger animals are associated with larger substrate particles. Using an artificial stream with scrubbed substrate we investigate the hypothesis that the assortative mechanism is stimulated by current avoidance rather than food searching.
2. In uniformly mixed substrate, G. pulex tend to move upstream, but in patchy substrate they are found in a predictable substrate grade. There is no discrimination between patches by different size classes of G. pulex in static water. The size assortative field pattern can only be reproduced under flow conditions in the laboratory. It occurs in the absence of food and irrespective of the sequence or location of substrate patches.
3. As density increases, G. pulex are displaced from the optimal patches but move into the open stream rather than into the cover of alternative patch types.
4. We discuss the significance of these results and suggest that some earlier models explaining mechanisms for microdistribution patterns tnay have been unnecessarily complex. Contrasts between natural and simulated stream situations and their experimental advantages are noted.     

Population dynamics and morphometries of Gammarus pulex L.: evidence of seasonal food limitation in a freshwater detritivore

SUMMARY. 1. Seasonal changes in population size structure of Gammarus pulex L. in a Cotswold stream appeared to indicate a growth check in late summer.
2. The relationships between dry weight and body length, and between the number of primary flagellar segments on the first antenna and either dry weight or body length provided further evidence of a reduction in growth in mid and late summer.
3. Body fat content was minimal (4% of dry weight) at the end of summer, when large particulate organic detritus was scarce or of poor quality, and maximal in late winter (17.9% dry weight in females; 9.4% in males), after a period of high food availability. In a field experiment, the fat content of animals in summer was raised to levels typical of winter by providing high quality food.
4. Field and experimental evidence together strongly infer that this population of G. pulex was subject to severe food limitation from early summer until leaf fall in autumn.     

State-dependent pairing behaviour in male Gammarus pulex (L.) (Crustacea, Amphipoda): effects of time left to moult and prior pairing status

Because mating can be costly in terms of time and energy, an individual's propensity to engage in courtship and mating activities might be modulated by its physiological state. However, so far, state-dependent mate choice has received little attention The present study examined the effect of both prior pairing status and time left to the moult on the ability of male Gammarus pulex (Crustacea, Amphipoda) to enter in precopula with receptive females. In the lab, males that were freshly collected in precopula pairs in the field had a higher probability of re-pairing and were quicker to enter in precopula with receptive females compared to males of similar size that were freshly collected unpaired. In addition, unpaired males found in the field were closer to their moult than paired males. Considered together, our results strongly suggest that time left to the moult and prior mating status directly influence male propensity to pair in G. pulex.     

Phenotypic variability in nesting success among Osmia lignaria propinqua females in a glasshouse environment: (Hymenoptera: Megachilidae)

Abstract. 1. The phenotypic variability of several estimates of fitness among seventeen nesting female O.lignaria Cresson was examined in a glasshouse provided with abundant resources.
2. Females exhibited wide variation in: (a) number of nests and cells made and in their rate of construction, (b) the incidence of mortality of offspring, and (c) the percentage of male offspring produced (52–100%).
3. In confirmation of earlier field studies: (a) more offspring of both sexes were produced during the first half of the nesting season, (b) most female offspring were produced early in the nesting season, (c) most parental investment during the last half of the flight season was made in male offspring, (d) there was a decrease in offspring size with season, with females decreasing more than males, and (e) the ratio of female/male body weight was = 1.8. These results appear to be due to ageing rather than to a reduction in resource availability.
4. No relation between female size and fecundity was evident.
5. There was no evidence that small females tended to produce a greater proportion of the smaller sex (males) than did large females.
6. Smaller offspring of both sexes had a much greater probability of dying overwinter than did large offspring. However, where the sexes overlapped in size (large males, small females), almost all females died and almost all males survived. We hypothesize that as size of offspring produced declines with season, a greater proportion of males are produced because they have a much greater probability of surviving at small body sizes.     

Male load-lifting capacity and mating success in the swarming caddis fly Athripsodes cinereus

Abstract. Males of the caddis fly Athripsodes cinereus (Curtis) (Trichoptera: Leptoceridae) swarm above the water surface of lakes and streams. Females enter swarms and are pursued until grasped by a male. The pair couple their genitalia in the air, and then the male alone flies the pair to the shore where they settle and complete the copulation. About 8% of the pairs (total n = 384 pairs) dipped in the water soon after the coupling manoeuvre and about 25% of those then separated. Males in dipping pairs ( n = 13) were on average smaller and relatively older than the males that successfully carried their mate to the shore ( n = 54). No differences were found for flight muscle ratio (weight of flight muscles/total body weight) or relative load (total load/flight muscle weight). Males were larger than females (wing length), though typically female Trichoptera are the larger sex. Large male body size in A. cinereus may be an adaptation for flight during pairing; i.e. larger males are more likely to be able to carry larger loads.     

Growth and survivorship as proximate causes of sexual size dimorphism in peninsula dragon lizards Ctenophorus fionni

Males and females differ in body size in many animals, but the direction and extent of this sexual size dimorphism (SSD) varies widely. Males are larger than females in most lizards of the iguanian clade, which includes dragon lizards (Agamidae). I tested whether the male larger pattern of SSD in the peninsula dragon lizard, Ctenophorus fionni, is a result of sexual selection for large male size or relatively higher mortality among females. Data on growth and survivorship were collected from wild lizards during 1991–1994. The likelihood of differential predation between males and females was assessed by exposing pairs of male and female lizards to a predator in captivity, and by comparing the frequency of tail damage in wild‐caught males and females. Male and female C. fionni grew at the same rate, but males grew for longer than females and reached a larger asymptotic size (87 mm vs. 78 mm). Large males were under‐represented in the population because they suffered higher mortality than females. Predation may account for some of this male‐biased mortality. The male‐biased SSD in C. fionni resulted from differences in growth pattern between the sexes. The male‐biased SSD was not the result of proximate factors reducing female body size. Indeed SSD in this species remained male‐biased despite high mortality among large males. SSD in C. fionni is consistent with the ultimate explanation of sexual selection for large body size in males.     

Pairing success and sperm reserve of male Gammarus pulex infected by Cyathocephalus truncatus (Cestoda: Spathebothriidea)

Manipulative parasites with complex life cycles are known to induce behavioural and physiological changes in their intermediate hosts. Cyathocephalus truncatus is a manipulative parasite which infects Gammarus pulex as intermediate host. G. pulex males display pre-copulatory mate guarding as a response to male-male competition for access to receptive females. In this paper, we tested the influence that C. truncatus-infection might have on male G. pulex sperm number and pairing success. We considered 3 classes of G. pulex males in our experiments: (i) uninfected males found paired in the field, (ii) uninfected males found unpaired in the field, or (iii) infected males found unpaired in the field. Both infected males and uninfected unpaired males paired less with a new female than uninfected paired males did. Furthermore, infected males appear to be at a strong disadvantage when directly competing for females with a healthy rival male, and had fewer sperm in their testes. We discuss the potential effect of male and female mating strategies on such male host mating alteration.     

Components of male fitness in relation to body size in Epirrita autumnata (Lepidoptera, Geometridae)

Abstract. 1. The effect of body size on different components of male fitness was studied for Epirrita autumnata , a geometrid known for its eruptive population dynamics. Body size is the main determinant of female fecundity in this species.
2. Longevity of males was found to have a weak negative correlation with body size at low temperatures. No significant correlation was found at higher temperatures.
3. We found no correlation between male size and female fecundity or egg size which is consistent with the small size of spermatophores in this species.
4. Small and large males were equally successful when allowed to compete for females in laboratory conditions.
5. In one or two field collections, males found mating were larger than males found singly. Large males also had an advantage in finding of virgin females, offered experimentally. No size-assortative mating was recorded.
6. We conclude that size-dependent mate location ability is the factor accounting for most of the variance in male fitness in E.autumnata. The dependence of fitness on body size may well be equally strong in males and females.     

Sexual reproduction in Daphnia pulex (Crustacea: Cladocera): observations on male mating behaviour and avoidance of inbreeding

1.  Mating behaviour in Daphnia appears to rely on random contact between males and sexual females rather than diffusible pheromones. Males may be able to discriminate sexually receptive females from females in other developmental stages and increase their mating efficiency. Males may also use chemical signals to avoid mating with females from the same clone and avoid the severe inbreeding depression that has been documented for intraclonal mating. The present study used experiments to test for the avoidance of intraclonal mating and assess male mating efficiency in D. pulex .
2.  Three clones were examined for the avoidance of intraclonal mating by providing males with an opportunity to mate with females of the same or two different clones. The proportion of intraclonal matings did not differ from the proportion of interclonal matings, suggesting that D. pulex males do not use kin discrimination to avoid mating with females from the same clone.
3.  The proportion of mated females decreased with increasing numbers and density of sexual females when exposed to a single male. This observation suggests that a male spends more time pursuing and copulating with sexually receptive females than non-receptive females and there is insufficient time to mate with all sexual females. The decrease in proportion of females mated could also be the result of sperm depletion in the male. Sperm depletion is unlikely to occur in nature because sexually receptive females are much rarer than in the experimental conditions.     

Size variation and reproductive success of female Aedes punctor (Diptera: Culicidae)

Abstract. 1. Variation in size (wing length) of females and males within a population of Aedes punctor (Kirby) at emergence was recorded in northern Britain during 1984 and 1985.
2. Wing length correlated well with body dry weight, confirming its usefulness as a measure of body size. In 1984, but not in 1985, mean size decreased significantly as emergence progressed.
3. The size of host-seeking females caught at human bait varied seasonally. This was partly accounted for by size variation during emergence.
4. Of females which fed to repletion, the largest spent least time in contact with the host.
5. In the laboratory, wing length correlated well with potential fecundity (number of ovarioles) and less well with actual fecundity (number of matured follicles) after feeding on human blood.
6. In the field, larger females were more successful at locating hosts, developed more egg clutches in a lifetime and generally lived longer than did smaller ones. Larger females appear to enjoy greater reproductive success than do smaller females.     

Effects of Environmental and Social Conditions on Homosexual Pairing in the Japanese Beetle (Popillia japonica Newman)

Homosexual pairing between males occurs under natural conditions in a wide variety of taxa, including many insect species, but few studies have investigated how environmental and social conditions affect same-sex pairing in insects. We investigated factors affecting homosexual pairing in male Japanese beetles (Popillia japonica Newman) in the field and in the laboratory. Specifically, we investigated how time of day, sex ratio, beetle density, and temperature affected the likelihood of homosexual pairing. In the field, male–male pairs constituted 1–6% of the pairs we collected. Homosexual pairs were more common in the afternoon than in the morning and the evening. Sex ratio, density, and temperature were all related to the likelihood of finding a homosexual pair, but the relationships were not linear. In the laboratory, higher male densities and relatively male-biased sex ratios were associated with an increase in the frequency of homosexual pairs. Homosexual pairs were more frequent at relatively low and relatively high temperatures. Males that mounted other males tended to be smaller than the males that they mounted. In addition, compared to males that were not homosexually paired, there was some indication that the mounting males were smaller, and the mounted males larger, than the unpaired males. Our data suggest that homosexual pairs are a result of males mistaking other males for females, and we hypothesize that the environmental and social factors cause changes in homosexual pairing through their effects on the frequency of pair formation and pair duration.     

SEX-RATIO SELECTION IN A BIVOLTINE THRIPS. I. CONDITIONAL SEX-RATIO MANIPULATION AND FITNESS VARIATION

Females of the bivoltine thrips Elaphrothrips tuberculatus (Hood) (Insecta: Thysanoptera) produce broods of either all males (by viviparity) or all females (by oviparity). Measurements of the sex-allocation ratio, ecological and physiological conditions affecting male and female offspring body size, and correlates of the relative fitnesses of adult males and females in relation to size indicate that female parents tend to be viviparous (produce males) if their offspring will become relatively large adults, and that males gain more in fitness from large size than do females. However, the conditions that link sex allocation with offspring fitness differ between the spring and summer generations. In spring, when breeding is synchronous, 1) oviparous and viviparous females do not differ in body size, 2) females tend to be viviparous where the fungus upon which they feed is relatively dense and where their offspring will become relatively large adults, and 3) fungus density is highly correlated with male and female offspring size. In summer, when breeding is relatively asynchronous, 1) viviparous females are much larger than oviparous females early (but not late) in the season, 2) large viviparous females begin breeding earlier than smaller ones, 3) offspring developing earlier in the season become larger adults, and 4) a higher proportion of females are viviparous earlier than later. Field experiments and field collections show that the covariation among sex allocation, conditions, and fitness is not caused by differential mortality by size or sex. Differences between the spring and summer generations in the cues used by females to adjust offspring sex ratio may be caused by seasonal variation in the factors that affect offspring size. However, in both generations, females tend to produce sons only when their offspring will become relatively large adults, whereas daughters are produced regardless of offspring size. These data suggest that females of E. tuberculatus avoid production of males (the sex with higher variance in expected fitness) when the size of their offspring is relatively uncertain.     

Sex ratios of Spalangia endius (Hymenoptera: Pteromalidae), in relation to current theory

Abstract. 1. Spulungiu endius Walker is a solitary parasitoid of house fly puparia.
2. The sex and size of S.endius was not related to host size.
3. In the laboratory the mean sex ratio of all offspring of nine groups, each comprising twenty females, was consistently female-biased (x = 83.5%, range 79–87%). The sex ratio in the field was less female-biased and showed greater fluctuation (61–75%). This may be a consequence of females laying male eggs before mating, some females remaining unmated, possible shorter adult life expectancy in the field than in the laboratory, and, perhaps, the presence of conspecific females.
4. The sex ratio of offspring of individual females varied from 66% to 100% females, and males were deposited early in the oviposition sequence.
5. Although a large number of fly puparia died before adult flies or parasitoids emerged (64.5%; n = 5874), there was no differential mortality of either sex.
6. Our results fit no general sex ratio hypothesis and we conclude that (i) the genetic nature of sex ratios in these insects needs careful examination, and (ii) the prevalence of female-biased sex ratios in solitary parasitoids needs investigation.     

Male size determines reproductive output in a paternal mouthbrooding fish

Size can have strong effects on reproductive success in both males and females, and in many species large individuals are preferred as mates. To estimate the potential benefits from mate choice for size in both sexes, I studied the effects of the size of each sex on the reproductive output of pairs of Banggai cardinalfish, Pterapogon kauderni, a sexually monomorphic obligate paternal mouthbrooder. When pairs were allowed to form freely, a size-assortative mating pattern was observed and larger pairs had a higher reproductive output as determined by total clutch weight and egg size. To separate the potential benefits from mate choice for size for each sex, I subsequently used these pairs to form reversed size-assortative pairs, that is, the largest male paired to the smallest female and vice versa. I found a positive correlation between male size and clutch size: relatively heavier clutches were found in pairs where females were given a larger male. This suggests that the size of the male influences clutch weight. For egg size, however, the size of both sexes seemed important. The study reveals the benefits of mutual mate choice on size in this species: larger females provide larger eggs and larger males can brood heavier clutches. Furthermore, these results suggest that females differentially allocate resources into the eggs according to the size of the mate.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Male-biased size dimorphism in ichneumonine wasps (Hymenoptera: Ichneumonidae) – the role of sexual selection for large male size

Abstract.  1. Sexual differences in body size are expected to evolve when selection on female and male sizes favours different optima.
2. Insects have typically female-biased size dimorphism that is usually explained by the strong fecundity advantage of larger size in females. However, numerous exceptions to this general pattern have led to the search for selective pressures favouring larger size in males.
3. In this study, the benefits of large size were investigated in males of four species of ichneumonine wasps, a species-rich group of parasitoids, many representatives of which exhibit male-biased size dimorphism.
4. Mating behaviour of all ichneumonine wasps are characterised by pre-copulatory struggles, in the course of which males attempt to override female reluctance to mate. A series of laboratory trials was conducted to study the determinants of male mating success.
5. A tendency was found for larger males as well as those in better condition to be more successful in achieving copulations. Size dimorphism of the species studied, mostly male-biased in hind tibia length but female-biased in body weight, indicates that sexual selection in males favours longer bodies and appendages rather than larger weight.
6. The qualitative similarity of the mating patterns suggests that sexual selection cannot completely explain the considerable among-species differences in sexual size dimorphism.
7. The present study cautions against using various size indices as equivalents for calculating sexual size dimorphism.
8. It is suggested that female reluctance in ichneumonine wasps functions as a mechanism of female mate assessment.     

The effects of temperature and body size on the mating pattern of a gregariously nesting bee, Colletes cunicularius (Hymenoptera: Colletidae)

Abstract. 1. In a 3-year study of the solitary bee Colletes cunicularius L. in Sweden, average body size and population density fluctuated greatly between years.
2. In this protandrous population, females mated just once and the sex ratio was slightly male biased. Males were smaller than females.
3. Size assortative mating (homogamy), associated with an increase in population density during the central days of female emergence and mating, was observed in two out of three years. Homogamy was also observed in pairs with remating males.
4. Most of the mating males had emerged the day they mated, but 42% were older. We found no support for a general large-male mating advantage.
5. Weight of emerging females and mating males were negatively correlated with ground temperature, indicating thermoregulatory influence on the process of sexual selection in this species.     

Effects of size at metamorphosis on stonefly fecundity, longevity, and reproductive success

Many organisms with complex life cycles show considerable variation in size and timing at metamorphosis. Adult males of Megarcyssignata (Plecoptera: Perlodidae) are significantly smaller than females and emerge before females (protandry) from two western Colorado streams. During summer 1992 stoneflies from a trout stream emerged earlier in the season and at larger sizes than those from a colder fishless stream, and size at metamorphosis did not change over the emergence period in either stream. We performed two experiments to determine whether variation in size at metamorphosis affected the fecundity, reproductive success and longevity of individuals of this stonefly species and if total lifetime fecundity was affected by the number of matings. In the first experiment, total lifetime fecundity (eggs oviposited) was determined for adult females held in small plastic cages in the field. Males were removed after one copulation, or pairs were left together for life and allowed to multiply mate. Most copulations occurred in the first few days of the experiment. Females in treatments allowing multiple matings had significantly lower total lifetime fecundity and shorter adult longevity than females that only mated once. Multiple matings also reduced longevity of males. Fecundity increased significantly with female body mass at emergence, but only for females that mated once. While multiple matings eliminated the fecundity advantage of large female body size, number of matings did not affect the significant positive relationship between body mass at metamorphosis and longevity of males or females. In a second experiment designed to determine if body mass at emergence affected male mating success, we placed one large and one small male Megarcys in an observation arena containing one female and recorded which male obtained the first mating. The large and the small male had equal probabilities of copulating with the female. Copulations usually lasted all night, and the unmated male made frequent, but unsuccessful attempts to take over the copulating female. Our data suggest that selection pressures determining body size at metamorphosis may operate independently on males and females, resulting in evolution of sexual size dimorphism, protandry, and mating early in the adult stage. We emphasize the importance of interpreting the fitness consequences of larval growth and development on the timing of and size at metamorphosis in the context of the complete life cycle. Received: 1 July 1997 / Accepted: 12 November 1997