Behavioural plasticity in support of a benefit for aggregation pheromone use in Drosophila melanogaster

We explored behavioural plasticity in the use of aggregation pheromone in the fruit fly Drosophila melanogaster Meigen (Diptera: Drosophilidae). Based on previous field observations, we formulated two hypotheses on a benefit of using aggregation pheromone for aggregated oviposition. One hypothesis concerns a benefit to the females themselves, where reduced harassment by males can enhance oviposition rate; the other concerns a benefit to their offspring, where larvae can exploit arduous substrates more efficiently. We derive contrasting expectations on the strength of the behavioural response to pheromone for substrates that differ in nutritional quality to larvae. High quality substrates relax the strength of larval competition, which allows for stronger aggregative responses of the females, but conversely, it may yield aggregation less necessary when the benefit is related to resource exploitation by the larvae. In indoor and outdoor dual choice set-ups, we tested the behavioural responses of the adults to the aggregation pheromone with substrates of varying quality, and examined oviposition behaviour. The response of adults to the aggregation pheromone was strong and robust for low quality substrates, but significantly weaker for a high quality substrate. This supports the hypothesis on a benefit to the larvae. Females retained aggregation pheromones in the absence of oviposition substrates for at least 24 h. In the outdoor set-up, substrate with aggregation pheromone received more than three times as many eggs as control substrates, and this was directly related to the number of adults that visited each substrate. Per capita, oviposition rates were not different for differently sized aggregations, and consequently, no evidence was found in support of the hypothesis on reduced harassment. The combination of aggregation pheromone possession and a mutualistic relationship with micro-organisms for Drosophila and other insects is discussed.     

Competition with filamentous fungi and its implication for a gregarious lifestyle in insects living on ephemeral resources

Abstract. 1. Recent studies have demonstrated the existence of positive density dependence in the survival and development of Drosophila (the so‐called Allee effect); however the underlying mechanisms of such Allee effects have remained elusive. Competition with filamentous fungi have often been suggested to be involved in causing high mortality at low larval density, but it has not yet been explicitly tested if the well known spatial aggregation of insect eggs yields a fitness benefit for the developing larvae in the presence of noxious moulds. 2. Using Drosophila melanogaster, the present study tested whether larval survival is greater in aggregations when confronted with various combinations of three representative mould species (Aspergillus, Alternaria, and Penicillium) and a head start for fungal development. 3. High rates of fungal‐dependent mortality and significant positive density‐dependent larval survival (i.e. Allee effects) were observed when larvae were confronted with food resources containing established colonies of Aspergillus or Alternaria. Neither the simultaneous transfer of Aspergillus or Alternaria spores with larvae to food patches nor food infections with Penicillium affected insect larval development. 4. Significant correlations between mould growth and larval survival could be identified, although the patterns that emerged were shown to be inconsistent when the effects were compared between fungal species and fungal priority. Because mould growth only partly explained larval survival, the influence of other fungal‐borne factors, e.g. mycotoxins, needs to be elucidated in order to understand the mechanistic basis of insect–mould interactions. 5. These results are the first to argue convincingly for moulds being involved in mediating Allee effects for insects on ephemeral resources; however they also demonstrate an unexpected diversity in insect–mould interactions. Considering this diversity may be important in understanding insect spatial ecology.     

Host–parasitoid interaction as affected by interkingdom competition

Although still underrepresented in ecological research, competitive interactions between distantly related organisms (so-called “interkingdom competition”) are expected to be widespread in various ecosystems, with yet unknown consequences for, e.g. trophic interactions. In the model host–parasitoid system Drosophila melanogaster–Asobara tabida, toxic filamentous fungi have been shown to be serious competitors that critically affect the density-dependent survival of host Drosophila larvae. This study investigates the extent to which the competing mould Aspergillus niger affects key properties of the well-studied Drosophila–parasitoid system and how the host–parasitoid interaction influences the microbial competitor. In contrast to slightly positive density-dependent host mortality under mould-free conditions, competing A. niger mediated a strong Allee effect for parasitised larvae, i.e. mortality decreased with increasing larval density. It was found that the common toxic fungal metabolite kojic acid is not responsible for higher death rates in parasitised larvae. Single parasitised Drosophila larvae were less harmful to fungal reproduction than unparasitised larvae, but this effect vanished with an increase in larval density. As predicted from the negative effect of fungi on host survival and thus on parasitoid fitness at low larval densities, A. tabida females spent less time foraging in fungus-infested patches. Interestingly, even though high host larval densities increased host survival, parasitoids still reduced their search efforts in fungus-infested patches, indicating a benefit for host larvae from feeding in the presence of noxious mould. Thus, this experimental study provides evidence of the potentially important role of interkingdom competition in determining trophic interactions in saprophagous animal communities and the dynamics of both host–parasitoid and microbial populations.     

Is aggregated oviposition by the blow flies Lucilia sericata and Phormia regina (Diptera: Calliphoridae) really pheromone‐mediated?

When female blow flies Lucilia sericata and Phormia regina (Diptera: Calliphoridae) oviposit in aggregations on carrion, even‐aged larval offspring reportedly develop faster, and fewer are parasitized or preyed upon. The benefits of aggregated oviposition equally affect con‐ and heterospecific larvae sharing a resource. The benefits imply that female blow flies engage in coordinated, pheromone‐mediated oviposition behavior. Yet, repeated attempts to identify oviposition pheromones have failed invoking doubt that they exist. Simply by regurgitating and feeding on carrion, flies may produce attractive semiochemicals. If flies were to aggregate in response to feeding flies rather than ovipositing flies, then the semiochemical cue(s) may be associated with the salivary gland. Working with L. sericata and P. regina and using liver as a surrogate oviposition medium, we test the hypotheses, and present data in their support, that (i) gravid or nongravid females ovipositing and/or feeding on liver enhance its attractiveness to gravid and nongravid females; (ii) females respond to semiochemicals from feeding heterospecific females; (iii) females respond equally well to semiochemicals from feeding con‐ and heterospecific females; (iv) macerated head tissues of females applied to liver enhance its attractiveness; and (v) females in direct contact with and feeding on liver, but not when next to yet physically separated from liver, enhance attraction of flies. We conclude that oviposition site‐seeking females do not respond to an oviposition pheromone. Instead, they appear to coopt semiochemicals associated with feeding flies as resource indicators, taking chances that resources are suitable for oviposition, and that ovipositing flies are present.     

Genetic Variation and the Role of Insect Life History Traits in the Ability of Drosophila Larvae to Develop in the Presence of a Competing Filamentous Fungus

Competitive interactions between organisms from distantly related phylogenetical branches have been suggested as being one of the most pervasive forms of interspecific competition. However, so-called inter-kingdom competition has rarely been the focus of ecological and evolutionary studies. Thus, a relatively novel hypothesis has been proposed on the basis that saprophagous insects might intensively compete with filamentous fungi for ephemeral resources (e.g. decaying plant tissue). Consideration that life history traits (e.g. developmental time) are adaptive in determining developmental success in the presence of con- or hetero-specifics competitors implies that these traits have been progressively established by natural selection. Because a similar scenario may apply to antagonistic interactions between saprophagous insects and filamentous fungi, one can expect the existence of heritable variation in developmental success when insect larvae are forced to grow in the presence of noxious mould. Therefore, this study aimed at discovering whether a local population of Drosophila melanogaster indeed harbours genetic variation in developmental success in the presence of the mould Aspergillus niger. By using the isofemale line technique, single larvae forced to feed on fungal infected or uninfected substrate were analysed for variation in survival probability to the adult stage, developmental time and body size of emerged adults. I found genetic variation in survival probability in fungal infected substrates but not in uninfected larval food sources. Mean developmental time and body size varied significantly among isofemale lines in both types of larval environment. Survival was negatively correlated with developmental time on fungal infected substrate, but variation in developmental time on fungal-free substrates was not correlated with survival on fungal infected food patches. Within-trait correlation between fungal infected and uninfected substrates was surprisingly weak, and developmental time was not correlated with body size. The results of this study demonstrate (a) the existence of genetic variation for larval developmental success in the presence of A. niger in a Drosophila population, and (b) heritability of important insect life history traits differed as a function of the larval environment (fungal infected or uninfected feeding substrate). I discuss models that might explain heritability differences and the evolutionary consequences of these results.     

Saprophagous insect larvae,Drosophila melanogaster,profit from increased species richness in beneficial microbes

Female fruit flies, Drosophila melanogaster, lay their eggs on decaying plant material. Foraging fly larvae strongly depend on the availability of dietary microbes, such as yeasts, to reach the adult stage. In contrast, strong interference competition with filamentous fungi can cause high mortality among Drosophila larvae. Given that many insects are known for employing beneficial microbes to combat antagonistic ones, we hypothesized that fly larvae engaged in competition with the noxious mould Aspergillus nidulans benefit from the presence of dietary yeast species, especially when they are associated with increasingly species rich yeast communities (ranging from one to six yeast species per community). On a nutrient‐limited fruit substrate infested with A. nidulans, both larval survival and development time were positively affected by more diverse yeast communities. On a mould‐free fruit substrate, merely larval development but not survival was found to be affected by increasing species richness of dietary yeasts. Not only yeast diversity had an effect on D. melanogaster life‐history traits, but also the identity of the yeast combinations. These findings demonstrate the importance of the structure and diversity of microbial communities in mutualistic animal–microbe interactions.     

Aggregative Behavior is Not Explained by an Allee Effect in the Walnut Infesting Fly, Rhagoletis juglandis

Component Allee effects are considered to be a driving force in the origin and maintenance of aggregative behavior. In this study, we examine whether a pattern of active host reuse by the walnut fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), involves an Allee effect. We examined how the density of clutches deposited within a fruit, the temporal pattern in which successive clutches are deposited and the spatial distribution of clutches over a fruit’s surface influences survival to pupation and pupal size. Within the density range used in this experiment (1 to 7 clutches), increases in larval density strongly reduced pupal weight but not larval survival to pupation. The temporal staggering of clutches into a host strongly reduced offspring survival and, probably owing to competitive release, increased the pupal weight of survivors. Offspring survival and pupal weight were affected relatively little by whether two clutches were deposited within the same oviposition punctures or were evenly spaced. In contrast, in three-clutch treatments offspring survival was higher when clutches were placed within the same oviposition cavity. However, pupal weights did not significantly increase when clutches were placed together and this relatively higher survival rate was not greater than that associated with hosts that contained fewer clutches. The results of the study failed to provide evidence of an Allee effect. We put forward a scenario under which females appear to reuse larval hosts to maximize their own reproductive success, albeit at the expense of the per capita fitness of their offspring.     

Effects of aggregation pheromone on individual behaviour and food web interactions: a field study on Drosophila

Abstract.  1. The effects of an aggregation pheromone on individual behaviour and food web interactions were investigated in two ecological communities, using Drosophila melanogaster and D. simulans as focal species.
2. Fruit substrates with aggregation pheromone were significantly more attractive to adult D. melanogaster and D. simulans than control fruit substrates, and the response was positively dose dependent. Competing species and natural enemies were also significantly attracted to substrates with the aggregation pheromone of D. melanogaster and D. simulans .
3. Significantly more eggs were deposited on pheromone-treated fruits than on control fruits, and the microdistribution of eggs within fruits was correlated to the microdistribution of the pheromone. The aggregation pheromone induced more females to share the breeding site.
4. The extremely high densities of fruit flies in the large aggregations appeared to reduce the oviposition rate of females. Physical interactions with conspecific and heterospecifics were frequently observed in the aggregations, and often led to patch leaving of the fruit flies.
5. Competition for food among larvae occurred at high densities and parasitism was density dependent. Aggregation pheromones can be directly responsible for these patterns through their effects on the con- and heterospecific behaviour.
6. The combined results show that aggregation pheromones affect a multitude of aspects in the ecology of interacting animals. The importance of incorporating the communication signals in ecological theory of aggregations is discussed.     

Oviposition Marking Behavior of <Emphasis Type="Italic">Diachasma alloeum</Emphasis>, (Hymenoptera: Braconidae), Parasitizing <Emphasis Type="Italic">Rhagoletis pomonella</Emphasis>, (Diptera: Tephritidae)

Diachasma alloeum (Muesebeck) (Hymenoptera: Braconidae) is a solitary larval endoparasitoid attacking Rhagoletis (Diptera: Tephritidae) species. Rhagoletis pomonella (Walsh) mark the surface of fruit after oviposition with an oviposition marking pheromone (OMP) which deters conspecific female flies. Herein we demonstrate that female D. alloeum wasps reared from either apple or hawthorn race R. pomenella larvae also deposit an OMP that reduces oviposition by conspecific female wasps. Significantly fewer wasps accepted fruit that had received prior wasp oviposition and OMP or OMP alone without oviposition compared with control fruit for a minimum of 7 days on both fruit types. Rinsing fruit with a 50% ethanol solution appeared to remove the OMP rendering fruit more acceptable for oviposition than marked fruit that was not rinsed. Wasps of each host race were able to detect and avoid the OMP of the sister race and fruit substrate type did not affect wasp response to the pheromone. The possibility of an internal marker deposited during oviposition is also discussed.     

Maternal effects increase survival probability in Drosophila subobscura larvae

Drosophilid flies breeding on ephemeral resource patches (e.g., decaying fruits) are assumed to transfer yeasts to their oviposition sites, presumably in order to positively affect offspring development. We tested this hypothesis with Drosophila subobscura Collin (Diptera: Drosophilidae) by manipulating their nutritional (yeast‐fed vs. non‐yeast‐fed) and reproductive status (mated vs. non‐mated). Flies were then released into vials containing decaying fruits (either sloes, crab apples, or Syrian plums). After a constant residence time in the vials, the flies were removed, 16 first‐instar larvae were transferred to the fruits and their survival probability to the adult stage was recorded. Whereas previous exposure of the larval substrate to yeast‐fed males and virgin females (yeast‐fed and non‐yeast‐fed) had no effect on survivorship, exposure to yeast‐fed and mated females that deposited eggs on the fruits (subsequently removed) led to a significant increase in the survival probability of the transferred larvae to the adult stage. Although the exact mechanism of yeast transmission remains to be determined, we suggest an active inoculation of the breeding substrates with yeast by ovipositing females. In agreement with previous studies, we found a negative effect of mould growth on larval survival, which, however, depended on the fruit type. We discuss various scenarios of yeast involvement in benefits to the insect larvae and suggest that insect–mould interactions should be examined in detail in order to better understand the behavioural and life‐history traits of insects that depend on ephemeral resources.     

The effects of the timing of exposure to cadmium on the oviposition behavior of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis>

Animals can greatly increase their fitness by choosing oviposition sites free from damaging substances such as the heavy metal cadmium (Cd). Previous research has shown that Drosophila melanogaster reared on uncontaminated media lay fewer eggs on substrates containing Cd. In this study, we examined the effects of prior exposure to Cd on oviposition site preference. We tested flies that had been exposed to Cd continuously from egg to adult, only during pre-adult stages, or for the 4 days of adulthood prior to testing. We found that flies avoid laying eggs on Cd-contaminated medium and also that flies exposed to Cd as adults laid significantly fewer eggs than controls never exposed to Cd. We did not observe consistent changes in oviposition as a result of pre-adult exposure alone. Our results suggest that the presence of cadmium in the environment, even at low doses and for short periods of time during adulthood, can harm the fitness of Drosophila melanogaster.     

Substrate selection for pit making and oviposition in an antlion, Myrmeleon bore Tjeder, in terms of sand particle size

The larvae of the pit‐making antlion Myrmeleon bore Tjeder live in open sand in riverbeds with a substratum consisting of various particle sizes. We analyzed the spatial distribution of their pits in a sandy floodplain to determine their larval and adult responses to the heterogeneous substrate. The spatial distribution pattern of their pits had an aggregated distribution, and there was a significant positive correlation between pit density and the ratio of medium‐size sand particles to total weight of sand. We examined the size of sand particles selected in the larval pit‐building behavior and the oviposition behavior of the adult. Both larvae and adults selected medium‐size sand particles. The larvae of M. bore are relatively sedentary predators and rarely move great distances. Thus, the present results suggest that habitat selection by adult females is a major factor causing the aggregative distribution of the pits.     

Avoidance of occupied hosts by the Hessian fly: oviposition behaviour and consequences for larval survival

Abstract 1. Unoccupied wheat plants and wheat plants occupied by conspecific eggs or larvae were presented to ovipositing female Hessian flies in choice tests.
2. The presence of conspecific eggs on the leaf surfaces of wheat plants did not appear to have any effect on the responses of ovipositing Hessian fly females.
3. The presence of conspecific larvae at the base and nodes of wheat plants for 1, 6, or 11 days had significant effects on Hessian fly oviposition. Eggs oviposited on plants were inversely proportional to larval densities and days of larval occupation.
4. Feeding by Hessian fly larvae is associated with several changes in wheat plants. One of these changes, the growth arrestment of the plant, was measured by recording the heights of plants used in oviposition tests. Plant heights were inversely proportional to both larval densities and days of occupation. Plant heights were directly proportional to eggs oviposited on plants.
5. The consequences of adult female avoidance of plants occupied by conspecific larvae were investigated by allowing females to oviposit on unoccupied plants and 1-day, 6-day, and 11-day larval occupied plants, then scoring at the end of the first larval instar the survival of the offspring that resulted from this oviposition.
6. Survival during the first larval instar was 88% for the offspring of females that oviposited on unoccupied plants, decreasing to 82, 31, and 4% on the 1-day, 6-day, and 11-day occupied plant treatments. On these four plant treatments, a positive correlation was found between larval performance (i.e. survival) and the preferences of ovipositing females.
7. On the four plant treatments, relationships between first-instar larval density and first-instar larval survival varied significantly. On unoccupied plants, survival was inversely proportional to density. On plants oviposited on at 6 days of larval occupation, survival was directly proportional to density.     

Erythritol ingestion impairs adult reproduction and causes larval mortality in Drosophila melanogaster fruit flies (Diptera: Drosophilidae)

Previous feeding studies showed the polyalcohol erythritol was toxic when ingested by adult laboratory fruit flies (Drosophila melanogaster). We asked whether erythritol could additionally affect fly population growth either through larval toxicity or through effects on adult reproduction. Females did not avoid laying on food substrates with 1M erythritol; laying rate on 1M erythritol food was similar to control food when females were given free‐choice access. Eggs laid or placed on 0.5 M to 2.5 M erythritol foods hatched at normal rates, suggesting erythritol was not toxic to eggs upon contact. Drosophila melanogaster larvae readily consumed food containing 1 M erythritol, but none of these larvae reached pupation. Longevity of larvae feeding on in 1 M erythritol food was significantly reduced relative to controls, and mean ± SE larval lifespan on erythritol was 1.54 ± 0.10 days (max. = 3 days). Exposing cohorts of second‐instar larvae to food with varying concentrations of erythritol showed the LD50 (at 24 hr) concentration was approximately 0.6 M. Taken together, these results suggest erythritol could be employed in effective larval‐sink baits. Adults flies fed with erythritol produced significantly fewer eggs on days when they fed on 1 M erythritol, and egg production was significantly reduced for one additional day after the adults were moved to control food. These findings suggest erythritol is rapid and effective at temporarily suppressing D. melanogaster reproduction, increasing its potential for use in effective insect population control.     

The relative fitness of Drosophila melanogaster (Diptera,Drosophilidae) that have successfully defended themselves against the parasitoid Asobara tabida (Hymenoptera,Braconidae)

Drosophila melanogaster larvae defend themselves against parasitoid attack via the process of encapsulation. However, flies that successfully defend themselves have reduced fitness as adults. Adults which carry an encapsulated parasitoid egg are smaller and females produce significantly fewer eggs than controls. Capsule-bearing males allowed repeated copulations with females do not show a reduction in their number of offspring, but those allowed to copulate only once did. No differences were found in time to first oviposition in females, or in time to first copulation in males. We interpret the results as arising from a trade-off between investing resources in factors promoting fecundity and mating success, and in defence against parasitism. The outcome of this investment decision influences the strength of selection for defence against parasitism.     

Oviposition behaviour of two tephritid fruit flies,Dacus tryoni and Dacus jarvisi,as influenced by the presence of larvae in the host fruit

Summary When offered a choice, females of the fruit flies Dacus tryoni (Frogg.) and D. jarvisi (Tryon) strongly preferred to lay in fruits without larvae rather than fruits which already contained larvae. Fruits which contained even low densities of larvae, including newly hatched ones, received many fewer eggs than control fruits. This preference was not influenced by the species of larvae present in the fruits nor by the distance to uninfested fruits. Discrimination occurred when fruits with and without larvae were close together (10 cm apart) and also when they were separated by distances of about one metre. Laboratory assays suggested that the flies detect chemical changes in the fruit associated with the decomposition which accompanies larval feeding, but they do not seem to detect the larvae Perse. This behaviour may be significant when these two species utilise the same host since the species which is able to infest fruits first will reduce the availability of hosts for the other species. In contrast to many other Tephritids (e.g. Rhagoletis, Anastrepha and Ceratitis) female Dacus don't discriminate against fruits which contain eggs nor do they deposit a pheromone to deter oviposition by females that subsequently visit the fruit. An hypothesis is proposed to explain the absence of oviposition-deterring pheromones in Dacus, and their presence in many other species of Tephritidae, on the basis of differences in life history and population structure.     

Efficacy and longevity of newly developed catnip oil microcapsules against stable fly oviposition and larval growth

The stable fly, Stomoxys calcitrans (Diptera: Muscidae), is one of the most important pests of cattle and costs U.S. cattle producers billions of dollars in losses annually. In this study, the efficacy of catnip oil encapsulated in gelatin in oviposition deterrence and larval growth inhibition in stable flies was examined under laboratory conditions. More than 98% inhibition of stable fly larval growth and female oviposition was observed in larval and oviposition media treated with encapsulated catnip oil (0.5 g). Further, dose–response tests showed that as little as 0.1 g of encapsulated catnip oil provided > 85% oviposition deterrence. The release of nepetalactones from the capsules was more rapid when the capsules were placed on a moist substrate rather than a dry substrate. Encapsulated catnip oil also exhibited antibacterial activity, supporting the hypothesis that its inhibition of larval growth may be based on its killing of the bacteria on which larvae feed. The use of encapsulated catnip oil can provide an alternative control strategy for stable fly management.     

Butterfly oviposition preference is not related to larval performance on a polyploid herb

The preference–performance hypothesis predicts that female insects maximize their fitness by utilizing host plants which are associated with high larval performance. Still, studies with several insect species have failed to find a positive correlation between oviposition preference and larval performance. In the present study, we experimentally investigated the relationship between oviposition preferences and larval performance in the butterfly Anthocharis cardamines. Preferences were assessed using both cage experiments and field data on the proportion of host plant individuals utilized in natural populations. Larval performance was experimentally investigated using larvae descending from 419 oviposition events by 21 females on plants from 51 populations of two ploidy types of the perennial herb Cardamine pratensis. Neither ploidy type nor population identity influenced egg survival or larval development, but increased plant inflorescence size resulted in a larger final larval size. There was no correlation between female oviposition preference and egg survival or larval development under controlled conditions. Moreover, variation in larval performance among populations under controlled conditions was not correlated with the proportion of host plants utilized in the field. Lastly, first instar larvae added to plants rejected for oviposition by butterfly females during the preference experiment performed equally well as larvae growing on plants chosen for oviposition. The lack of a correlation between larval performance and oviposition preference for A. cardamines under both experimental and natural settings suggests that female host choice does not maximize the fitness of the individual offspring.     

Superparasitism of larval hosts by the walnut fly, <Emphasis Type="Italic">Rhagoletis juglandis</Emphasis>, and its implications for female and offspring performance

The oviposition-preference–offspring-performance hypothesis predicts that female insects should prefer to deposit clutches on or in hosts that maximize offspring performance. An important assumption behind this prediction is that female fitness is tightly correlated with the fitness of any one offspring. In this study, we evaluate offspring performance in the walnut fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), in relation to a previously described oviposition preference for previously exploited host fruit. In particular, we examined how superparasitism of walnut hosts influences offspring survival and weight at pupation under field conditions. We found that superparasitism was common and that increases in larval densities within fruit were associated with reduced larval survival and weight at pupation. In a laboratory experiment, female size was correlated with lifetime fecundity. In this system, oviposition preference is therefore negatively, not positively, correlated with offspring performance. We argue that patterns of female preference in this system reflect direct benefits to females that are traded off against costs in terms of offspring fitness. Because female fitness is a product not only of offspring quality but also of the total number of offspring produced, female walnut flies may be optimizing their fitness by producing many less fecund offspring. Studies examining the preference-performance hypothesis should consider the reproductive conflicts between parents and offspring as potential factors that influence the congruence between parental preference and offspring performance.     

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.