The choice of oviposition site and the effects of density and oviposition timing on survivorship in<Emphasis Type="Italic"> Rana japonica</Emphasis>

In species that lack parental care and whose larvae are restricted to a given space throughout development, choice of egg-laying or reproductive sites often greatly influences fitness. Furthermore, reproductive success of individuals may be affected not only by conditions of the breeding area but also by the time when they start to breed, because intraspecific variation in the timing of breeding may result in asymmetric competition between larvae. We addressed these issues in an experimental and field study of the Japanese brown frog, Rana japonica. We tested whether the survival success of larvae decreased with an increase in the number of deposited egg masses and with a greater delay in oviposition. We found that the number of egg masses and timing of oviposition together significantly predicted larval survival per clutch. Moreover, we observed the natural oviposition of R. japonica to examine the prediction that if the density of larvae and the timing of oviposition affect survivorship of larvae, R. japonica will avoid depositing eggs in pools in which egg masses have already been deposited. We found that for small pools only, R. japonica tended to favor oviposition at unoccupied pools (those lacking egg masses and larvae) in the spawning season. For large pools, however, adults favored those already occupied by egg masses in the spawning period. The density of egg masses and tadpoles or the timing of oviposition may have less effect on spawning or the survival of tadpoles in large pools than in small pools.     

A reproductive tradeoff in an herbivorous lady beetle: egg resorption and female survival

A reproductive tradeoff between current egg production and subsequent survival in a lady beetle, Epilachna niponica, a specialist herbivore on a thistle, Cirsium kagamontanum, was investigated at the two study sites, A and F. Survival of reproductive females decreased consistently from early May until mid-June, but apparently increased thereafter. In contrast, males showed a consistent decrease in survival throughout the reproductive season, without any sign of recovery. Dissection of ovaries of sampled females revealed that egg resorption increased late in the reproductive season, coincident with increased female survival. Reproductive females stopped oviposition immediately after a large flood in 1979 at site F. Two weeks after the habitat perturbation, females resumed oviposition in response to a flush of new leaves on damaged plants. Female survival sharply increased during the nonoviposition period, and declined when egg-laying resumed. Approximately 40% of long-lived reproductive females at site F survived up to the following reproductive season in the next year. Also, some of these long-lived females were observed ovipositing in the following reproductive season. The long-lived reproductive females which had previously invested in reproduction survived equally well as newly emerged females which had not reproduced in summer. These results suggest that there is a reproductive tradeoff between current egg production and subsequent survival. Egg resorption may be an adaptive ovipositional response to habitat perturbation such as flooding, which considerably reduces offspring fitness due to absolute shortage of food. Also, increased female survival accompanied by egg resorption enhances the likelihood of the future oviposition in the second reproductive season, thereby increasing a female's lifetime reproductive success.     

Egg-laying tactic in Phyllomorpha laciniata in the presence of parasitoids

Female insects are expected to choose oviposition sites that have the best conditions for offspring development and survival. Natural enemies, such as predators and parasitoids, may have a strong influence on the selection of oviposition substrates by phytophagous insects. The golden egg bug, Phyllomorpha laciniata (Villers) (Heteroptera: Coreidae) has an unusual reproductive strategy. Females mainly use conspecifics, both males and other females, as egg-laying substrates, but occasionally they oviposit on plants as well. Survival of the eggs is higher when eggs are carried by conspecifics than when they are laid on plants, due to predation and parasitism. We investigated egg-laying behavior in the forced presence of the egg parasitoid Gryon bolivari (Giard) (Hymenoptera: Scelionidae). Specifically, we studied whether females provide egg protection by avoiding oviposition under the risk of egg parasitization. We expected a lower oviposition rate under parasitoid presence, and the eggs, if any, to be placed preferably on conspecifics and not on plants, thus ensuring higher survival of the progeny. The results show that P. laciniata 's egg-laying rate was lower when they were enclosed with parasitoids than when parasitoids were absent, especially when plants were the only substrate to oviposit on. Moreover, females showed strong preference for laying eggs on conspecifics rather than on plants. Egg-laying in P. laciniata appears to be not only influenced by the availability of conspecifics, but also by the presence of egg parasitoids. This indicates that females may be able to detect G. bolivari and avoid oviposition when parasitoids are present. We discuss the possibility of conspecifics as enemy-free space.     

Females prefer mating males in the carmine triplefin,Axoclinus carminalis,a paternal brood-guarder

Synopsis This study investigates the role of male mating status in female choice patterns in the carmine triplefin, Axoclinus carminalis, a tripterygiid fish that exhibits paternal care. The distribution of daily reproductive activity is clumped, with many males receiving no mates and some receiving three or more. Females in this species do not prefer larger males, and characteristics of the oviposition site appear to have minimal effects on male mating success. When a female is removed from a male early in the daily spawning period, that male attracts fewer additional females for the remainder of the spawning period than does a control male. These changes in mating success are temporary, and do not affect mating success on subsequent days. A preference for mating males or males that are guarding eggs could provide asymmetric benefits for males to defend oviposition sites. This preference for males with eggs could be acting alone or with other factors such as high variance in oviposition site quality to favor the evolution of paternal care in fishes.     

BREEDING COMPETITION IN A PACIFIC SALMON (COHO: ONCORHYNCHUS KISUTCH): MEASURES OF NATURAL AND SEXUAL SELECTION

In the breeding system of Pacific salmon, females compete for oviposition territories, and males compete to fertilize eggs. The natural selection in females and sexual selection in males likely has been responsible for their elaborate breeding morphologies and the dimorphism between the sexes. We quantified direct-selection intensities during breeding on mature coho salmon (Oncorhynchus kisutch), measured for seven phenotypic characters, including three secondary sexual characters. Wild and sea-ranched hatchery coho were used to enhance the range of phenotypes over which selection could be examined. The fish were allowed to breed in experimental arenas where we could quantify components of breeding success as well as estimate overall breeding success. We found that without competition, natural selection acts only on female body size for increased egg production; there is no detectable selection on males for the phenotypic distribution we used. Under competition, the opportunity for selection increased sixfold among females. Natural selection favored female body size and caudal-peduncle (tail) depth. Increased body size meant increased egg production and access to nesting territories. The caudal peduncle, used in burst swimming and nest digging, influenced both successful egg deposition and nest survival. Increasing density increased competition among females, though it did not significantly intensify natural selection on their characters. In males, competition increased the opportunity for selection 52-fold, which was nine times greater than for females. Sexual selection favored male body size and hooked snout length, both characters directly influencing male access to spawning opportunities. Selection on male body size was also affected significantly by breeding density. The ability of large males to control access to spawning females decreased at higher densities reflecting an increase in the operational sex ratio. Further, the relative success of small males, which could sneak access to spawning females, appeared to increase as that of intermediate-sized males decreased. Such disruptive selection may be responsible for the evolution of alternative reproductive tactics in salmon.     

Biological and Behavioral Effects of Heavy Metals in Drosophila melanogaster Adults and Larvae

Heavy metals are essential components of biological systems but are extremely toxic at high doses. As a result, we hypothesized that perception of heavy metals through gustation may exist in Drosophila melanogaster. In this study, we investigated the behavioral effects of iron, copper, zinc, and cadmium on D. melanogaster gustation, oviposition, and pupation-site selection. In addition, we examined the biological effects of heavy metals on the fruit fly survival and reproductive success. Our results illustrate that D. melanogaster responds behaviorally to the presence of high concentrations of heavy metals in food. All metals acted as repellents to the fruit flies at high doses, with the egg-laying and feeding of the female flies significantly decreasing. Furthermore, supplementation of heavy metals in the culture medium reduced survival to the adult stage and shortened the life span of adult flies. From these observations, we speculate that D. melanogaster avoidance behavior towards high concentrations of heavy metals may have a positive effect on their survival and reproductive success in nature, particularly in the presence of metal-contaminated food sources.     

The growth bars on tail feathers in the male Styan's Grasshopper Warbler may indicate quality

We propose that growth bar size may indicate the quality of individuals in Styan's Grasshopper Warblers Locustella pleskei . Positive correlations of an individual's standardized growth bar width between years showed that some individuals of both sexes were usually in better condition than others. The survival rate of males with wider growth bars was higher than that of other males. These survivors might have survived better because of a better body condition. We also examined the relationships between growth bar width and reproductive parameters (arrival date, pairing date and reproductive success). Males with wider growth bars arrived earlier, acquired a mate earlier and achieved higher total reproductive success by breeding twice. In females, growth bar width was correlated with reproductive parameters (arrival date, pairing date and total reproductive success), but not with survival. These results suggest that males with wider growth bars were of higher quality than those with narrower bars. The contrasting results for the two sexes may be explained by differences in their breeding tactics.     

Potential Costs of Selecting Good Sites For Offspring: Increased Risk of Drowning And Negative Effects on Egg Production

Many studies have shown the benefits of selecting suitable sites for offspring survival and growth. However, costs of selecting suitable sites have been little covered. If the female's costs associated with selecting suitable sites exceed the benefit from improvement of offspring performance, selecting such sites can have a negative effect on the ongoing reproductive success for ovipositing females. We investigate the potential costs of selecting suitable sites in the water strider Aquarius paludum insularis. Where there exists a risk of egg parasitism, the female A. paludum will submerge and select a deep site, more suitable for offspring survival, for oviposition. By forcing A. paludum underwater once a day for 10 d, we investigated the potential costs associated with oviposition at deep sites, firstly relating to the performance of submergence (latency to asphyxiation or proportion of buoyancy loss) and, secondly, in the number of eggs laid. Buoyancy became weaker and the latency to asphyxiation became shorter with the number of submergences. The number of eggs laid in the period of forced submergence was smaller than both before and after the period. Selecting deep sites can therefore increase the mortality risk of ovipositing females and also decrease the number of eggs laid in their lives. Selecting suitable sites in all the oviposition bouts can decrease ovipositing females' reproductive success. Costs, not only benefits, should be taken into consideration for understanding oviposition site selection during the lifespan of an ovipositing female.     

Modelling evolutionarily stable strategies in oviposition site selection, with varying risks of predation and intraspecific competition

Many ovipositing mosquitoes, as well as other species, can detect biotic factors that affect fitness. However, a female mosquito seeking a high quality oviposition site (e.g. one with low risk of predation and competition to her progeny) must often balance the competing risk of increasing probability of mortality to herself while she continues to search, against increased probability of finding a high quality site. Such oviposition site selection may affect adult population size. We examined a female mosquito’s expected strategy of oviposition site selection under conditions of varying predator prevalence and adult mortality risk, by combining a detailed structured population model with a Markov chain implementation of the adult behavioural process. We used parameter values from the specific mosquito-predator system, Culiseta longiareolata-Notonecta maculata, although the overall results can be generalised to many mosquito species. Our model finds the evolutionarily stable strategy of oviposition site selection for different parameter combinations. Our model predicts that oviposition strategy does not vary smoothly with varying environmental risk of adult mortality, but that certain oviposition strategies become unstable at some parameter values. Mosquitoes will distribute their reproductive effort between breeding sites of varying predation risk only when adult mortality is low or larval competition high. Our model predicts that females will continue searching for predator-free pools, rather than oviposit in the first site encountered, regardless of the risk of mortality to the adult. The ecological basis for a reproductive strategy with alternative behaviours is important for understanding the effect of biotic factors on the population dynamics of mosquitoes, and for the development of biological control strategies, such as the dissemination of predator-cue chemicals.     

Temperature influences host instar selection in an aphid parasitoid: support for the relative fitness rule

The relative fitness rule states that parasitoid females should adopt risk‐prone reproductive behaviours when expecting low reproductive success. Temperature influences the reproductive success of insects by affecting their basal metabolic rate during development, their egg load at emergence, and their life expectancy as adults. Using an aphid–parasitoid model system, we investigated the influence of developmental and adult temperature on the risk‐sensitive decision‐making of females. We considered the use of a low‐quality host nymphal instar by the aphid parasitoid Aphidius ervi to be a risk‐prone behaviour. Immature females were reared at 12, 20 or 28 °C and had access to the four nymphal instars of the potato aphid Macrosiphum euphorbiae for oviposition at one of these temperatures. Host selection behaviour was continuously recorded during exploitation of an aphid patch. We observed that warm‐developed females and parasitoids foraging at high temperature attacked low‐quality hosts more frequently than females from other treatments. These results support the hypothesis that a decrease in expected parasitoid reproductive success resulted in risk‐prone behaviours. To our knowledge, this is the first experimental evidence suggesting that temperature influences host stage selection and risk‐sensitive making decision in parasitoids, and the present study is the first to support the relative fitness rule. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.     

Deposition and longevity of oviposition-deterring pheromone in the cabbage seed weevil

Abstract The cabbage seed weevil ( Ceutorhynchus assimilis Payk.) lays eggs singly into pods of oilseed rape ( Brassica napus L.) through punctures bored with the mouthparts, preferring pods not recently used for oviposition. A simple new choice test has been used to test individual components of egg-laying behaviour for their effect on oviposition site selection. It is confirmed that an oviposition-deterring pheromone (ODP) is deposited during abdomen brushing of the pod which follows egg-laying. Neither pin punctures, weevil feeding punctures, oviposition punctures nor eggs had any deterrent effect. Pods walked on by female weevils were not avoided by those laying eggs. Observations suggest that the ODP is sensed by contact chemoreceptors on the antennae. The deterrent effect lasted only 1–2 h. The implications of these findings on the adaptive significance of the pheromone and its possible use in pest control are discussed.     

Oviposition Site Selection in the Malayan Giant Frog (Limnonectes blythii) in Singapore: Conservation Implications

Amphibians require specific habitats for breeding and loss or degradation of such habitats can negatively affect reproductive success. Oviposition site selection within a habitat is also important as site quality is linked to larval survivorship and metamorphic success. We investigated oviposition site preferences of the stream-breeding frog Limnonectes blythii in Singapore through surveys and habitat measurements of breeding and non-breeding sites(N = 30 and 32, respectively). The study species L. blythii is classified as Near Threatened(NT) in the IUCN red list and is associated with medium sized forest streams. L. blythii appeared to prefer streams with higher water p H and shallower water depths for oviposition. Our findings have implications in conservation management as it provides the baseline for habitat restoration for creation of new and for preserving existing breeding habitat of L. blythii.     

Site Selection and Foraging in the Eresid Spider Stegodyphus tentoriicola

Habitat selection has profound consequences for survival and reproductive success. We investigated web relocation behaviour in relation to plant structure and body condition as well as the plasticity of foraging behaviour of the spider Stegodyphus tentoriicola. Spiders inhabiting thorny vegetation were larger, built larger webs than spiders in thornless plants and relocated their webs less frequently. Web relocation affected reproductive success through a delay of oviposition. Spiders supplemented with extra food improved body condition and built smaller webs than control spiders implying a crucial role of food in limiting fitness of S. tentoriicola. Reduced investment in webs suggests a trade-off between the benefit of more food against the cost of web-construction. We propose that S. tentoriicola exhibit a “silk and energy saving” strategy when saturated.     

Reproductive Success Associated with Territoriality, Sneaking, and Grouping in Male Rose Bitterlings, Rhodeus ocellatus (Pisces: Cyprinidae)

The spawning success of male rose bitterlings, Rhodeus ocellatus, adopting an alternative reproductive style, was estimated through behavioural data and electrophoretic paternal analyses in field observations and experiments. Three mating patterns were observed: territoriality, sneaking, and grouping. Mating patterns depended on a male's relative size and on local male density (the number of males around a spawning spot: a mussel). Spawning patterns (pair spawning, pair spawning with sneaker, and group spawning) varied with local male density. Time-budget data of the territorial males indicated a trade-off between chasing and courtship behaviour as local male density changed. Females deposited appoximately only 1 egg per egg-laying into the mussels. As a result of isozyme analysis, a minimum of 12% (two out of 17) of the offspring in the sample were found to have been fathered by sneaker males in pair spawning with sneaker. I scored through behavioural data the mating success per spawning for each pattern, on an individual basis. The average reproductive success per spawning for each pattern was: territorial (0.61), sneaking (0.31) and grouping (0.11), and thus the successes of the patterns were not equal. Accordingly, the alternative reproductive styles of male rose bitterlings are best interpreted as alternative phenotypes in a conditional behaviour.     

Oviposition in Culicoides impunctatus under laboratory conditions

We present a laboratory-based examination of oviposition preference in the Scottish biting midge, Culicoides impunctatus (Diptera: Ceratopogonidae). A variety of oviposition substrates were screened in no-choice and choice bioassays for efficacy in eliciting an egg-laying response. Both upper-layer photosynthetic Sphagnum spp. moss and Juncus articulatus infusions were identified as producing increased oviposition under no-choice conditions. During choice trials against a control of damp cottonwool, upper-layer Sphagnum spp. moss produced a significantly greater egg-laying response. These conclusions are interpreted in terms of possible cues involved in oviposition site selection and assessed for future use in colonisation of this troublesome species.     

Seasonal change in the opportunity for sexual selection

Environmental and population parameters that influence the strength of sexual selection may vary considerably over the course of the reproductive season. However, the potential for sexual selection frequently fails to translate into variation in reproductive success among individuals. We investigated seasonal changes in variation in reproductive success, measured as the opportunity for sexual selection, using parentage analysis in 20 experimental populations of the European bitterling (Rhodeus amarus, Cyprinidae), a small freshwater fish with a promiscuous, resource-based mating system. We showed that although the largest males sired most offspring over the entire reproductive season, variation in reproductive success and hence the opportunity for sexual selection was low at the start of the season but increased significantly at its end. This seasonal difference probably arose from the superior competitive endurance of large males and from a higher temporal clustering of reproductively active females at the start of the breeding season than later in the season. The spatial distribution of oviposition sites had a negligible effect on the variation in reproductive success. We discuss the potential implications of our results for the importance and strength of sexual selection in natural populations.     

Sexual signaling in the European bitterling: females learn the truth by direct inspection of the resource

In many taxa females appear to base their mate choice on multipletraits. But the relative importance of different traits inmate choice has rarely been determined. Here we show that femalesof a freshwater fish, the European bitterling, Rhodeus sericeus,base their mate choice on multiple traits that differ in theirreliability as indicators of expected reproductive successand are used at different stages of the decision process. Theinitial decision to inspect a male is based on male behaviorand red coloration, whereas the final spawning decision isbased on the quality of the live unionid mussel, Anodonta anatina,that the male is defending as an oviposition site. Male traitsmay indicate which males are worth inspecting by reflectingmale quality, such as reproductive condition and genetic constitution.Male traits do not, however, reflect mussel quality, as bright males also court females vigorously toward mussels that yielda low probability of survival of the offspring. Females, onthe other hand, are choosier than males in their choice ofspawning site and seem to gain reliable information about thesurvival probability of the eggs by inspecting the mussel directly.     

Simulating smallmouth bass reproductive success in reservoirs

An individual-based model of nesting smallmouth bass, Micropterus dolomieu, is used to predict the effects of water level fluctuations on reproductive success. The model simulates daily nest site selection, spawning, nest guarding, and development and survival of eggs, embryos, and larvae until dispersal. The model was configured for Brownlee Reservoir, Idaho-Oregon. Each reservoir bank is represented as a rectangular grid of cells, with each cell characterized by an assigned slope, substrate, and elevation, and by a water depth that varied daily. Nest site (cell) selection is determined from a habitat suitability index (HSI) based on substrate, slope, and depth. Development of young is temperature-dependent. Mortality of young occurs via attrition, abandonment by the guarding male, and probabilistic whole-nest catastrophe. Simulations were performed that used observed data for 1991 to 1996. Model predicted nest habitat selection and egg-to-dispersal survival rates were similar to observed values. Additional model simulations showed that: (1) nest habitat selection was best predicted when depth, slope, and substrate were considered together rather than singly, (2) egg-to-dispersal survival was related to the magnitude of water level fluctuations during the peak spawning period, (3) relating mortality to HSI values resulted in lower survival, while relating growth to HSI values resulted in higher survival, and (4) spawning habitat is not limiting in Brownlee Reservoir. Suggestions for future monitoring of reservoir-based smallmouth bass populations are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

A generalist brood parasite modifies use of a host in response to reproductive success

Avian obligate brood parasites, which rely solely on hosts to raise their young, should choose the highest quality hosts to maximize reproductive output. Brown-headed cowbirds (Molothrus ater) are extreme host generalists, yet female cowbirds could use information based on past reproductive outcomes to make egg-laying decisions thus minimizing fitness costs associated with parasitizing low-quality hosts. We use a long-term (21 years) nest-box study of a single host, the prothonotary warbler (Protonotaria citrea), to show that local cowbird reproductive success, but not host reproductive success, was positively correlated with the probability of parasitism the following year. Experimental manipulations of cowbird success corroborated that female cowbirds make future decisions about which hosts to use based on information pertaining to past cowbird success, both within and between years. The within-year pattern, in particular, points to local cowbird females selecting hosts based on past reproductive outcomes. This, coupled with high site fidelity of female cowbirds between years, points to information use, rather than cowbird natal returns alone, increasing parasitism rates on highly productive sites between years.     

Host selection in insects: reproductive interference shapes behavior of ovipositing females

In nature, closely related species often utilize different host species, but it is still unclear what factors contribute to the evolution and maintenance of such diversified host selection. In this review, I describe how negative interspecific mating interactions (reproductive interference) can shape host selection by animals, focusing mainly on phytophagous and predatory insects. First, I explain an important premise of this hypothesis, which is that the adult reproductive site is the same as the feeding site for the offspring. Next, I describe several mathematical models and well-studied empirical systems to show that reproductive interference can sufficiently drive and maintain different host selection between phylogenetically related species. Then, I argue for the first time that reproductive interference can cause an oviposition preference in insects that is not optimal for the survival and development of the offspring, as a result of maternal adaptive behavior that maximizes the mother's own fitness. Furthermore, I argue that in insects, reproductive interference probably shapes oviposition behavior before the female alights on the host (e.g., habitat preference), without affecting post-alighting decision making. I would like to emphasize that these two arguments represent the novel approach to clarify the unrevealed pattern of complex insect oviposition behavior.