Shiny cowbird Molothrus bonariensis egg size and chick growth vary between two hosts that differ markedly in body size

In birds, egg size affects chick growth and survival and it is an important component of reproductive success. The shiny cowbird Molothrus bonariensis is an extreme generalist brood parasite that uses hosts with a wide range of body masses. Survival of cowbird chicks decreases with host body mass, as competition for food with nestmates is more intense in large than in small hosts. We studied variation in shiny cowbird egg size and chick growth in two hosts that differ markedly in body size: the chalk‐browed mockingbird Mimus saturninus (70–75 g), and the house wren Troglodytes aedon (12–13 g). We analyzed: 1) if females parasitizing mockingbirds lay larger eggs than those parasitizing wrens, and 2) the association between egg size and chick growth. We experimentally controlled for time of parasitism and number of host chicks and evaluated growth rate of male and female parasite chicks. Shiny cowbirds parasitizing mockingbird nests laid larger eggs than those parasitizing wren nests. Chick body mass after hatching was positively associated with egg size until chicks were five days of age, but there was no association between egg size and growth rate, or asymptotic mass. There were no sexual differences in egg size or body mass at the time of hatching, but growth rate was higher in males than in females leading to sexual dimorphism in asymptotic mass. Differences in egg size between hosts and the effect of egg size on body mass after hatching support the hypothesis that different females are specialized in the use of hosts that differ in body mass.     

Spawning strategy in Atlantic bobtail squid <Emphasis Type="Italic">Sepiola atlantica</Emphasis> (Cephalopoda: Sepiolidae)

This study aimed to determine the spawning strategy in the Atlantic bobtail squid Sepiola atlantica, in order to add new information to the knowledge of its reproductive strategy. A total of 12 females that spawned in aquaria were examined. Characteristics of the reproductive traits and egg clutches were similar to those of other known Sepiolidae. Clutch size varied from 31 up to 115 eggs. Females of this species had incorporated up to 1.58 times of their body weight into laid eggs. The size of laid eggs showed a positive correlation with maternal body size, supporting the idea that female size is a determinant of egg size. Our data suggest that S. atlantica is an intermittent terminal spawner, and that its spawning strategy comprises group-synchronous ovary maturation, multiple egg laying, and deposition of egg clutches in different locations. The obtained data provide insights for future comparative studies on reproductive allocation.     

Size‐dependent reproductive success of wild zebrafish Danio rerio in the laboratory

Size‐dependent reproductive success of wild zebrafish Danio rerio was studied under controlled conditions in the laboratory to further understand the influence of spawner body size on reproductive output and egg and larval traits. Three different spawner size categories attained by size‐selective harvesting of the F₁‐offspring of wild D. rerio were established and their reproductive performance compared during a 5 day period. As to be expected, large females spawned more frequently and had significantly greater clutch sizes than small females. Contrary to expectations, small females produced larger eggs when measured as egg diameter with similar amounts of yolk compared to eggs spawned by large spawners. Eggs from small fish, however, suffered from higher egg mortality than the eggs of large individuals. Embryos from small‐sized spawners also hatched later than offspring from eggs laid by large females. Larval standard length (L_S)‐at‐hatch did not differ between the size categories, but the offspring of the large fish had significantly larger area‐at‐hatch and greater yolk‐sac volume indicating better condition. Offspring growth rates were generally similar between offspring from all size categories, but they were significantly higher for offspring spawned by small females in terms of L_S between days 60 and 90 post‐fertilization. Despite temporarily higher growth rates among the small fish offspring, the smaller energy reserves at hatching translated into lower condition later in ontogeny. It appeared that the influence of spawner body size on egg and larval traits was relatively pronounced early in development and seemed to remain in terms of condition, but not in growth, after the onset of exogenous feeding. Further studies are needed to explore the mechanisms behind the differences in offspring quality between large‐ and small‐sized spawners by disentangling size‐dependent maternal and paternal effects on reproductive variables in D. rerio.     

Divergence in reproductive ecology of the three-spined stickleback,Gasterosteus aculeatus

An anadromous population (trachurus form) and three freshwater populations (leiurus form) of the three-spined stickleback,Gasterosteus aculeatus, in central Japan were compared with one another in the reproductive traits: body length at maturity, egg size, clutch size, the swell of abdomen and the number of eggs per nest. The anadromous fish which become larger in body size at maturity spawn eggs smaller in size and greater in number than the freshwater fish. The abdomen swell of gravid females expressed by the proportion of abdomen width to body length was greater in the freshwater fish. The anadromous male fish collected a mean of 2,638 eggs with a range of 1,119 to 4,052 eggs from about 6–7 females. In the three freshwater (the Yamayoke, the Tsuya and the Jizo) populations, males must have mated with about 9–22, 7–18 and 4–7 females respectively. It seems that theleiurus form increases its reproductive success by its much more mating opportunities and the parental efforts of nesting males as well as by spawning large eggs. Furthermore, among the freshwater populations, the Jizo one inhabiting the upper stream was clearly larger in body size, in egg size and in clutch size than the Yamayoke and the Tsuya ones which inhabit stable waterbodies with springs. It is possible that the Jizo population adopted the strategy of spawning a few large eggs as an adaptation to its habitat. The causal and functional explanations in reproductive characteristics among the four populations are discussed in regard to differences in the environmental conditions.     

Maternal investment increases with altitude in a frog on the Tibetan Plateau

Reproducing females can allocate energy between the production of eggs or offspring of different size or number, both of which can strongly influence fitness. The physical capacity to store developing offspring imposes constraints on maximum clutch volume, but individual females and populations can trade off whether more or fewer eggs or offspring are produced, and their relative sizes. Harsh environments are likely to select for larger egg or offspring size, and many vertebrate populations compensate for this reproductive investment through an increase in female body size. We report a different trade‐off in a frog endemic to the Tibetan Plateau, Rana kukunoris. Females living at higher altitudes (n = 11 populations, 2000–3500 m) produce larger eggs, but without a concomitant increase in female body size or clutch size. The reduced diel and seasonal activity at high altitudes may impose constraints on the maximum body size of adult frogs, by limiting the opportunity for energy accumulation. Simultaneously, producing larger eggs likely helps to increase the rate of embryonic development, causing tadpoles to hatch earlier. The gelatinous matrix surrounding eggs, more of which is produced by large females, may help buffer developing embryos from temperature fluctuations or offer protection from ultraviolet radiation. High‐altitude frogs on the Tibetan Plateau employ a reproductive strategy that favours large egg size independent of body size, which is unusual in amphibians. The harsh and unpredictable environmental conditions at high altitudes can thus impose strong and opposing selection pressures on adult and embryonic life stages, both of which can simultaneously influence fitness.     

Size Assortative Mating and Reproductive Success of the Funnel-Web Spider, Agelena limbata (Araneae; Agelenidae)

Field observations and laboratory experiments were conducted to assess the relation between male size and reproductive success in the funnel-web spider, Agelena limbata Thorell (Agelenidae), in 2 years. In this species, the body size of males is similar to that of females. In the field, size assortative mating occurred in both years. In 1 year, partial correlation coefficient analysis indicates that male cephalothorax width is a beter predictor of the copulated female cephalothorax width than of the date of pairing. In laboratory experiments, females tended to reject courting males that were smaller in relative body size, and males that were larger in relative body size had greater copulation success. Consequently female rejection of smaller courting males has some contribution to size assortative mating. Since larger females deposited more numerous eggs in the field, larger males are expected to have a higher reproductive success.     

Effects of suspended clay on Daphnia body growth and fitness

1. Suspended sediments often reduce the abundance, fitness, and feeding rates of planktonic cladocerans. This paper examines the effect of suspended clay particles on the body growth and life-history parameters of Daphnia ambigua. Cohorts were exposed to zero or 50mgl^?1 suspended clay (particle size <2μm). Individuals exposed to clay had significantly lower body lengths at a given age than individuals in the control cohort. Parameters for the von Bertalanffy body growth equation were calculated for each individual; the asymptotic maximum body length (l_max) was reduced, but the growth rate constant (k) was unaffected, by the presence of suspended clay. Body length at reproductive maturity was lower in the presence of suspended clay; this may be an adaptive response and is similar to the response of cladocerans to limiting food concentrations. 2. Age-specific survivorship and fecundity were both reduced, and the ages of maturity and first reproduction were both increased, in the presence of suspended clay. The overall effect of these demographic changes was a 70% decline in the net reproductive rate (R_o), a measure of fitness. 3. Suspended clay affected the relationship between body length and fecundity. Animals of a given length produced fewer eggs in the presence of suspended clay. In addition, there was a significant interaction in the effects of body length and clay on brood size: the slope of the regression line of brood size as a function of body length was lower in the presence of clay. This interaction has also been observed in limiting food concentrations, and may be caused by reductions in body length at maturity in low food or high clay environments. 4. The similarity between the effects of limiting food concentrations and suspended clay on Daphnia body growth, survivorship, fecundity, and brood size makes sense, given previous observations showing that suspended clay reduces the feeding rate of D. ambigua by up to 70% due to mechanical interference with feeding behaviours.     

Paternal body size affects reproductive success in laboratory-held zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

Across many fish species, large females tend to exhibit higher individual reproductive success due to elevated fecundity and the provisioning of better conditioned eggs and offspring compared to small females. By contrast, effects of paternal body size on reproductive success are less well understood. We disentangled the maternal- and paternal-size dependent effects on reproductive output and early life history in zebrafish (Danio rerio). In the laboratory, females and males from four size categories (small, medium-sized, large and very large) were allowed to spawn freely in a full factorial design with 10 replicates per size combination. As expected, larger females produced more eggs and better conditioned offspring compared to smaller females. Male body size further contributed to zebrafish reproductive success: offspring sired by large males exhibited higher hatching probability and these offspring also hatched earlier and larger than offspring fertilized by small males. However, the largest males experienced lower mating success and received fewer eggs than males of the smaller size classes. While male body size substantially affected reproductive success in zebrafish, it remained unclear whether and to what degree direct paternal effects (e.g., related to sperm quality) or indirect paternal effects stemming from differential allocation patterns by females were the mechanism behind our findings. Answering this question constitutes an important future research topic.     

Factors affecting variation in the reproductive investment of female treefrogs,Hyla intermedia

In anurans, fecundity (clutch size) is the most important determinant of female reproductive success. We investigated three possible causes responsible for fecundity variation in female Italian treefrogs, Hyla intermedia, during four breeding seasons: (i) variation in morphological (body size and condition) and life-history (age) traits; (ii) variation in the tradeoff between the number and the size of eggs; (iii) seasonal effects and within-season differences in the timing of deposition. At the population level, we found no evidence for a tradeoff between the number and the size of eggs, because they both correlated positively with females’ body size. Conversely, neither age nor post-spawning body condition showed any effect on female reproductive investment. Independent of body size, we found no evidence for variation in reproductive effort among different breeding seasons, but strong evidence for a decrease of clutch size and an increase of egg size with the advancing of a breeding season. To test for the functional significance of the observed temporal variation in allocation strategy, we carried out a rearing experiment in semi-natural conditions on a random sample of ten clutches. The experiment showed a negative effect of clutch size and a positive effect of egg size on both tadpole growth and developmental rates, suggesting that reproductive investment, although constrained by body size, can be adjusted by females to the time of deposition to increase the chances of offspring survival.     

Interpopulation variation in egg size in the winter cherry bug relative to oviposition site selection

To investigate whether or not oviposition on substrates other than host plants (e.g., non‐host plants, abiotic materials) would affect the evolution of egg size in phytophagous insects, we studied the winter cherry bug, Acanthocoris sordidus (Thunberg) (Heteroptera: Coreidae), as a model organism for its interpopulation variation in oviposition preference. The rate of oviposition off host plants is much higher in the Amami Island population than in either the Kyoto or Kochi populations. We compared egg size and number among the three local populations from Kyoto, Kochi, and Amami Island. In addition, to evaluate the adaptive significance of larger eggs for offspring in terms of searching for host plants, we examined the relationship between egg size and first‐instar body size. We also searched for a relationship between egg size and starvation tolerance in the second instars because first instars can develop to second instars without food intake, and thus the substantial host‐searching stage is the second instar, when females lay their eggs off host plants. Females from the Amami population produced fewer larger eggs than females from either the Kyoto or Kochi population. Regardless of the local population, the body size of first instars that emerged from larger eggs was larger, and the second instars originating from larger eggs had a higher starvation tolerance. The larger body size and higher starvation tolerance should enable nymphs to disperse further, which may enhance the probability of successfully reaching host plants. These results suggest that egg size in A. sordidus may be determined in relation to its oviposition habits to maximize reproductive success, resulting in interpopulation variation in egg size.     

Effect of Population Density and Female Body Size on Number and Size of Offspring in a Species with Size‐Dependent Contests over Resources

When size‐dependent contests over resources influence reproductive success, the trade‐off between number and size of offspring depends on the frequency of contests. Under these circumstances, clutch size should decrease and offspring size should increase as contests become more frequent. We tested these predictions with the burying beetle Nicrophorus pustulatus through manipulation of rearing densities. Burying beetles reproduce on small vertebrate carcasses, a rare but high quality food source for the larvae. Large beetles are more likely to win contests over carcasses and gain exclusive access to a carcass. The winner of a contest kills eggs and larvae already present on a carcass. As a result of the rarity of carcasses, burying beetles are unlikely to breed more than once. As predicted, brood size of N. pustulatus decreased with increasing rearing density. Despite a negative correlation between brood size and larval mass, larval mass did not increase with increasing rearing density. This may be due to the special biology of N. pustulatus which can use snake eggs for reproduction. Potentially larger supply of resources and generally small population densities of N. pustulatus may weaken selection on body size and thus the correlation between brood size and larval mass. As size‐dependent constraints can limit reproductive phenotypes, we examined whether female size influenced reproductive phenotype. Small females produced larger broods with smaller, but more variable, offspring than large females. Mechanical constraints of egg size seem an unlikely explanation for the differences because burying beetles can compensate for small egg size through parental care. Energetic constraints may impact small females because body mass and brood size of small females decreased with increasing density. Yet, at all density levels small females produced larger, not smaller, broods than large females. The larger and more variable broods of small females seem to be in agreement with a bet‐hedging strategy.     

Nest-site size as a short-term constraint on the reproductive success of paternal fishes

Synopsis Because female signal blennies, Emblemaria hypacanthus (Pisces: Chaenopsidae), exhibited a preference for the larger of two males in laboratory mate-choice experiments, male reproductive success was predicted to be positively correlated with male size in field populations. This prediction was met at one field site, but not a second where shelter size, rather than male size, was the primary correlate of male reproductive success. This finding appeared to be the consequence of the relatively small size of shelters (vacant gastropod shells) at the second site. At this site, significantly more males had their shelters filled with eggs and variance in male reproductive success was lower than at the first site. Limited area for deposition of eggs is potentially an important constraint on the short-term reproductive success of paternal males of shelter- or crevice-spawning fishes.     

Reproductive strategy as a major factor determining female body size and fertility of a gregarious parasitoid

The generally known “adult size‐fitness hypothesis” (ASFH) is applied to the gregarious parasitic wasp Anaphes flavipes (Foerster, 1841) (Hymenoptera: Mymaridae). ASFH is dependent on the reproductive strategy of the mother, which means the larger females have more offspring compared to smaller females. Two main factors, the mother's body size and food quantity received during larval development, can affect the body size of the offspring. For the first time, we present a study on the relative effect of both factors on fitness of the same species, wasp A. flavipes. Our data confirmed that females of A. flavipes with larger body sizes had more offspring compared to smaller ones. At the same time, mother's body size does not seem to affect the body size of the offspring. The other studied factor, quantity of food received during larval development, can be influenced by reproductive strategy (number of parasitoids developing in one host egg), host quality or the duration of development. We found only the reproductive strategy to have a statistically significant effect on body size. We demonstrated that the variable reproductive strategy (VRS) of wasp A. flavipes causes a plasticity in body size and future number of offspring. The generally known “trade‐off” scheme (more small offspring or fewer bigger offspring) does not apply to A. flavipes, because their large females have more offspring and it is their reproductive strategy that determines body size.     

Nonlinear continuum of egg size-number trade-offs in a snake: is egg-size variation fitness related?

The relationship between offspring size and offspring number is crucial to life history evolution. To examine how these two life history variables are coupled and whether an altered balance between them will result in changes in maternal fitness, we manipulated clutch size of the Chinese cobra (Naja atra) by using the techniques of hormonal manipulation and follicle ablation. Females receiving exogenous follicle-stimulating hormone produced more but smaller eggs, and females undergoing follicle ablation produced fewer but larger eggs. Neither body size (body mass and snout-vent length) at hatching nor egg mass at oviposition had a role in determining hatchling survival and growth. Female hatchlings were more likely to die in early post-hatching days and grew more slowly than male hatchlings. Our data show that: (1) there is a nonlinear continuum of egg size-number trade-offs in N. atra within which there is a single inflexion where the rate at which egg size decreases with increasing clutch size, or clutch size increases with decreasing egg size, is maximized; (2) there is a fixed upper limit to egg size for a given-sized female, and the limit is not determined by her body volume; (3) egg size has no role in determining hatchling survival and growth; and (4) the extent to which females may enjoy reproductive benefits in a given reproductive episode depends on how well egg size and egg number are balanced.     

Subsociality and female reproductive success in a mycophagous thrips: An observational and experimental analysis

Oviparous females of the haplodiploid, facultatively viviparous thrips Elaphrothrips tuberculatus(Thysanoptera: Phlaeothripidae) guard their eggs against female conspecifics and other egg predators. The intensity of maternal defense increases with clutch size. Field and laboratory observations indicate that cannibalism by females is an important selective pressure favoring maternal care. Experimental removals of guarding females showed that egg guarding substantially increases egg survivorship and that the survivorship of undefended eggs is higher in the absence of nonguarding female conspecifics than in their presence. The fecundity of viviparous females increases with the number of eggs cannibalized. The reproductive success of oviparous females increases with body size and local food density and decreases with local density of breeding females. Social behavior may not have advanced beyond maternal care in Elaphrothrips tuberculatusbecause, relative to Hymenoptera, capabilities for helping relatives are few or nonexistent, and the causes of variation in female reproductive success are not influenced easily by cooperation among females.     

Reproductive ecology of the jacky dragon (Amphibolurus muricatus): an agamid lizard with temperature‐dependent sex determination

Abstract The jacky dragon, Amphibolurus muricatus (White, ex Shaw 1790) is a medium sized agamid lizard from the southeast of Australia. Laboratory incubation trials show that this species possesses temperature‐dependent sex determination. Both high and low incubation temperatures produced all female offspring, while varying proportions of males hatched at intermediate temperatures. Females may lay several clutches containing from three to nine eggs during the spring and summer. We report the first field nest temperature recordings for a squamate reptile with temperature‐dependent sex determination. Hatchling sex is determined by nest temperatures that are due to the combination of daily and seasonal weather conditions, together with maternal nest site selection. Over the prolonged egg‐laying season, mean nest temperatures steadily increase. This suggests that hatchling sex is best predicted by the date of egg laying, and that sex ratios from field nests will vary over the course of the breeding season. Lizards hatching from eggs laid in the spring (October) experience a longer growing season and should reach a larger body size by the beginning of their first reproductive season, compared to lizards from eggs laid in late summer (February). Adult male A. muricatus attain a greater maximum body size and have relatively larger heads than females, possibly as a consequence of sexual selection due to male‐male competition for territories and mates. If reproductive success in males increases with larger body size, then early hatching males may obtain a greater fitness benefit as adults, compared to males that hatch in late summer. We hypothesize that early season nests should produce male‐biased sex ratios, and that this provides an adaptive explanation for temperature‐dependent sex determination in A. muricatus.     

Contrasting reproductive traits in two species of mangrove‐dwelling littorinid snails in a seasonal tropical habitat

Previous studies on the reproductive biology of littorinid snails have focused on rocky shore species, investigating how these gastropods can achieve maximal reproductive success, as well as on processes of sexual selection. This study documented differences in the reproductive traits of two mangrove‐dwelling littorinids, Littoraria ardouiniana and L. melanostoma, in Hong Kong. Reproductive activity of both species was most intense during the summer months. Mating pairs of the two species generally occurred in the tree canopies. Few false mating pairs (same sex or heterospecific pairs: <10%) were recorded, and members of both species showed size‐assortative mating. Littoraria ardouiniana had a shorter reproductive season but a higher intensity of mating and higher seasonal fecundity, than did L. melanostoma. Members of both species showed bi‐lunar periodicities of egg or larval release, synchronized with spring tides. Fecundity showed a strong positive relationship with body size in L. ardouiniana, but not in L. melanostoma. Females of L. ardouiniana released entire broods of larvae in a single brief event, whereas females of L. melanostoma released fewer eggs over 1–8 d. Release of larvae in L. ardouiniana involved a series of short bursts and was much faster than the trickle release of eggs in L. melanostoma. The contrasting reproductive traits in these two species represent different strategies to optimize reproductive success in mangrove habitats.     

Reproductive success in male Japanese minnows,Pseudorasbora parva: observations under experimental conditions

To evaluate the spawning success of male Japanese minnows,Pseudorasbora parva, and female mate choice, spawning behaviour was observed under both artificial and experimental conditions. Larger males had larger territories and greater reproductive success. The body weight of territorial males decreased during the maintenance of territories, while that of non-territorial males increased significantly. When the weight of non-territorial males exceeded that of territorial males, the former began to establish new territories on the substrate, suggesting a conditional strategy by non-territorial males to trade off immediate reproductive success with growth and hence improve future reproductive success. Females chose males with larger body size, probably based on dominance rank rather than the quality (or size) of territory. It was concluded that females choose males of higher dominance rank and that males compete for large territories, both of which play an important part in male reproductive success.     

The effect of petal size manipulation on pollinator/seed-predator mediated female reproductive success of Hibiscus moscheutos

The effects of petal-size manipulations on the behavior of pollinators and pollen/seed predators, and on pollen removal and deposition, were studied in Hibiscus moscheutos (Malvaceae) populations. The ultimate effects on the female reproductive success of flowers, such as fruit set, seed predation rate, and final seed set were also measured. We applied three levels of petal removal (100%, 50%, and 0% size reduction in radius) to flowers in natural populations. Two pollinators (Bombus pennsylvanicus and Ptilothrix bombiformis) ignored flowers without petals, suggesting that pollinators use petals as a visual cue to locate flowers. Consequently, 100% petal removal reduced female reproductive success considerably, mainly through a higher rate of fruit abortion due to failure of pollen deposition on stigmas. No significant differences between the 50% petal removal treatment and uncut control were detected in any components of female success examined. The results, therefore, suggest that differences in petal size have little influence on female reproductive success of Hibiscus flowers at our study site. Final seed set varied considerably depending on the larval densities of two coleopteran seed predators (Althaeus hibisci and Conotrachelus fissunguis). A. hibisci responded to petal size, and a higher density of adults was found in flowers in which petal size had not been reduced. Because Althaeus feed on pollen as adults and no effect of petal size on seed predation was detected, the preference of Althaeus for larger flowers may represent a foraging strategy for adult beetles and may exert counteracting selection pressure on petal size through male reproductive success of flowers. Received: 30 November 1997 / Accepted:12 June 1998     

THEORY OF MAST REPRODUCTION IN PLANTS: STORAGE-SIZE DEPENDENT STRATEGY

The amount of available resource can be considered to restrict mast seeding behavior in plants, regardless of the cause of masting itself. The reproductive strategy of a plant that has stopped growing, but continues to maintain a constant size, is based on the allocation of assimilated resources between reproduction and storage. If both plant death and population growth rates are constant, the plant strategy is dependent only on its own storage size. Conditions for the evolution of mast reproduction were analyzed under both constant and varying environments, from the view point of storage-size dependent strategy, which is influenced by three parameters: the advantage coefficient of mast seeding, β; the cost coefficient of delayed seeding, γ; and the assimilation rate, P. The model indicated that: (1) mast seeding evolves only with a greater than linear increase in reproductive success with effort (β > 1) under both constant and varying environments; (2) in mast seeding, a critical storage size, S_c*, occurs, above which plants utilize all storage for reproduction; (3) in a constant environment, S_c* increases with increasing β and P and/or decreasing γ; (4) concomitant with (3), an intermasting period, τ, also increases with increasing β and decreasing 7, but is independent of P; (5) in a varying environment, S_c* and the average of τ increase with increasing variance of the assimilation rate, P; and (6) concomitant with (5), the intermasting period has a certain probability distribution. In addition, reproductive synchrony within the population and the relationship between mast seeding and species diversity are discussed.