Fitness consequences of variable maternal provisioning in quacking frogs (Crinia georgiana)

Variable maternal provisioning may evolve when there is variation in the quality of offspring environments. The frog Crinia georgiana has high variability in egg size both within and between clutches, independent of female phenotype. It breeds in ponds with high spatial and temporal variation in habitat quality. Egg size strongly affected offspring fitness in good and poor quality offspring environments, whether the egg size difference was from between or within clutches. Since there is a trade-off in egg size and number, these fitness consequences translate to strong effects on maternal fitness. In the variable and unpredictable offspring environment of C. georgiana, the maintenance of variable maternal provisioning both within and between clutches is likely to be an evolved response to the offspring environment.     

Fitness consequences of egg-size variation in the lesser snow goose

We investigated the relationship between eggsize variation and (a) egg hatching success, (b) chick survival to fledging and recruitment, and (c) adult female survival, over 12 years in the lesser snow goose (Anser caerulescens caerulescens). By comparing the means and variances of egg size for successful and unsuccessful eggs, our aim was to assess the relative fitness of eggs of different sizes and to determine the type of selection operating on egg size in this species. As both egg size and reproductive success vary with age in the lesser snow goose we controlled for the effects of female age. Egg-size variation is very marked in this population, varying by up to 52% for eggs hatching successfully. However, there was no relationship between egg size and post-hatching survival of goslings to fledging or recruitment, either within or between broods, pooling across years. Egg size varied significantly between successful and unsuccessful clutches in only 2 of 33 individual year comparisons. First-laid eggs surviving to onset of incubation, and eggs hatching successfully, were on average larger than unsuccessful eggs, but this was probably due to the confounding effects of female age-specific and sequence-specific egg survival. Variance of egg size differed significantly between successful and unsuccessful eggs in only 3 of 24, and 0 of 21, individual year comparisons for pre- and post-hatching survival respectively. We therefore found little evidence for a relationship between egg-size variation and offspring fitness, or for strong directional, normalising or diversifying selection operating on egg size, in the lesser snow goose. In addition, there was only weak support for the hypothesis that egg-size variation is maintained by temporal variation in selection pressure (sensu Ankney and Bisset 1973). It is likely that egg-size variation represents the pleiotropic expression of alleles affecting more general physiological or metabolic processes. While this does not rule out the existence of alleles with more direct effects on egg size we suggest that their contribution to heritable egg size is small.     

Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

Temperature effects on egg size and their fitness consequences in the yellow dung fly Scathophaga stercoraria

Organisms and parts of an organism like eggs or individual cells developing in colder environments tend to grow bigger. A unifying explanation for this Bergmann's rule extended to ectotherms has not been found, and whether this is an adaptive response or a physiological constraint is debated. The dependence of egg and clutch size on the mother's temperature environment were investigated in the yellow dung fly Scathophaga stercoraria. Smaller eggs were laid at warmer temperatures in the field and the laboratory, where possible confounding variables were controlled for. As clutch size at the same time was unaffected by temperature, this effect was not due to a trade-off between egg size and number. Temperature-dependent egg sizes even persisted within individuals: when females were transferred to a cooler (warmer) environment, they laid third-clutch eggs that were larger (smaller) than their first-clutch eggs. The fitness consequences of these temperature-mediated egg sizes were further investigated in two laboratory experiments. Neither egg and pre-adult survivorship nor larval growth rate were maximized, nor was development time minimized, at the ambient temperature corresponding to the mother's temperature environment. This does not support the beneficial acclimation hypothesis. Instead, this study yielded some, but by no means conclusive indications of best performance by offspring from eggs laid at intermediate temperatures, weakly supporting the optimal temperature hypothesis. In one experiment the smaller eggs laid at 24 °C had reduced survivorship at all ambient temperatures tested. Smaller eggs thus generally performed poorly. The most parsimonious interpretation of these results is that temperature-mediated variation in egg size is a maternal physiological response (perhaps even a constraint) of unclear adaptive value. This revised version was published online in November 2006 with corrections to the Cover Date.     

Egg components, egg size, and hatchling size in leatherback turtles

Relationships between egg size, egg components, and neonate size have been investigated across a wide range of oviparous taxa. Differences in egg traits among taxa reflect not only phylogenetic differences, but also interactions between biotic (i.e., maternal resource allocation) and abiotic (i.e. nest environment conditions) factors. We examined relationships between egg mass, egg composition, and hatchling size in leatherback turtles (Dermochelys coriacea) because of the unique egg and reproductive characteristics of this species and of sea turtles in general. Albumen comprised 63.0%+/-2.8% (mean+/-S.D.) of egg mass and explained most of the variation in egg mass, whereas yolk comprised only 33.0%+/-2.7%. Additionally, leatherback albumen dry mass was approximately 16% of albumen wet mass. Whereas hatchling mass increased significantly with egg mass (n = 218 clutches), hatchling mass increased by only approximately 2 g for each 10 g increase in egg mass and was approximately 10-20 g greater than yolk mass. Taken together, our results indicate that albumen might play a particularly significant role in leatherback embryonic development, and that leatherback eggs are both capable of water uptake from the nest substrate and also possess a large reservoir of water in the albumen. Relationships between egg mass and egg components, such as variation in egg mass being largely explained by variation in albumen mass and egg mass containing a relatively high proportion of albumen solids, are more similar to bird eggs than to eggs of other non-avian reptiles. However, hatchling mass correlates more with yolk mass than with albumen mass, unlike patterns observed in bird eggs of similar composition.     

Effects of temperature on reproductive output, egg provisioning, juvenile hormone and vitellogenin titres in the butterfly Bicyclus anynana

Environmentally induced phenotypic plasticity is common in nature. Hormones, affecting multiple traits and signaling to a variety of distant target tissues, provide a mechanistic link between environments, genes and trait expression, and may therefore well be involved in the regulation phenotypic plasticity. Here, we investigate whether in the tropical butterfly Bicyclus anynana temperature-mediated plasticity in egg size and number, with fewer but larger eggs produced at lower temperatures and vice versa, is under control of juvenile hormone, and whether different temperatures cause differences in egg composition. Female B. anynana butterflies showed the expected response to temperature, however, we found no evidence for an involvement of juvenile hormone. Neither haemolymph JH II and JH III titres nor vitellogenin levels differed across temperatures. The smaller eggs produced at the higher temperature contained relatively higher amounts of water, free carbohydrates and proteins, but relatively lower amounts of lipids. While these smaller eggs had a lower absolute energy content, total reproductive investment was higher at the higher temperature (due to a higher fecundity). Overall, our study indicates that temperature-mediated plasticity in reproduction in B. anynana is mechanistically related to a biophysical model, with oocyte production (differentiation) and oocyte growth (vitellogenesis) having differential temperature sensitivities.     

Social and environmental influences on egg size evolution in frogs

The reproductive strategies of frogs are highly diverse, but analysis of these strategies in a phylogenetic context has lagged behind other taxa. Here we investigate associations between aspects of parental care and egg size in a phylogenetic context. We obtained data on egg size and parental care strategies in various species of frogs from the scientific literature. We developed a phylogenetic supertree of frogs by combining the results of multiple phylogenetic analyses using matrix representation parsimony. We used phylogenetic pairwise comparisons to investigate the correlation between various forms of parental care and egg size across the order Anura. We also investigated correlations between tadpole carnivory and egg size, and phytotelm breeding and egg size. We also investigated the association of egg size with several environmental factors. Parental care, male parental care, direct development, stream breeding and montane breeding habitats were all associated with large egg size. Female care (in species with trophic egg feeding), carnivory, use of small pools (phytotelmata) and use of temporary pools were not associated with egg size.     

Copulatory behaviors and body condition predict post-mating female hormone concentrations, fertilization success, and primary sex ratios in Japanese quail

Environmental cues and social interactions are known to influence reproductive physiology and behavior in vertebrates. In female birds, male courtship displays can result in the growth of ovarian follicles, the production of reproductive hormones, and stimulation of oviduct development, all of which have the potential to influence maternal investment. Male Japanese quail follow a typical sequence of copulatory behaviors during a mating interaction and often force copulations with unreceptive females. We hypothesized that female Japanese quail could adjust maternal investment in response to male copulatory behaviors during a single mating interaction. We investigated the relationships between 1) male copulatory behaviors and post-mating concentrations of steroids in the female, 2) female steroid concentrations and fertilization success of inseminations and 3) female steroid concentrations and the offspring sex ratio. We found that male condition and copulatory behaviors predicted female steroid concentrations and maternal investment in eggs laid after a mating trial. The body condition of one or both mates was a significant predictor of the changes in female corticosterone and testosterone concentrations after mating, whereas specific male copulatory behaviors significantly predicted changes in female progesterone concentrations. Male and female body condition, male neck grabs and post-mating concentrations of female corticosterone, progesterone, and testosterone were all significant predictors of egg fertilization rates. Female body condition, male copulation efficiency, and female testosterone concentrations were significant predictors of offspring sex ratios. Our results show that phenotypic and behavioral characteristics of male Japanese quail modulate female steroid concentrations and result in changes in maternal investment.     

Body temperature and resistance to evaporative water loss in tropical Australian frogs

Although the skin of most amphibians measured to date offers no resistance to evaporative water loss (EWL), some species, primarily arboreal frogs, produce skin secretions that increase resistance to EWL. At high air temperatures, it may be advantageous for amphibians to increase EWL as a means to decrease body temperature. In Australian hylid frogs, most species do not decrease their resistance at high air temperature, but some species with moderate resistance (at moderate air temperatures) gradually decrease resistance with increasing air temperature, and some species with high resistance (at moderate air temperatures) abruptly decrease resistance at high air temperatures. Lower skin resistance at high air temperatures decreases the time to desiccation, but the lower body temperatures allow the species to avoid their critical thermal maximum (CT(Max)) body temperatures. The body temperatures of species with low to moderate resistances to EWL that do not adjust resistance at high air temperatures do not warm to their CT(Max), although for some species, this is because they have high CT(Max) values. As has been reported previously for resistance to EWL generally, the response pattern of change of EWL at high air temperatures has apparently evolved independently among Australian hylids. The mechanisms involved in causing resistance and changes in resistance are unknown.     

Intraclonal variation in propensity for mixis in several rotifers: variation among females and with maternal age

In 14 previously published experiments with clones of three species of Brachionus, Epiphanes ukera, and Rhinoglena frontalis, females cultured singly in small volumes from birth through most of their reproductive period usually produced some mictic daughters. Here, these data are analyzed to test two hypotheses: the propensity of females in a clone to produce mictic daughters will vary significantly among females, and will decrease with female age. Significant heterogeneity (χ²) among replicate females was demonstrated in two clones of B. calyciflorus from Florida, in single clones of this species from Georgia, Texas, Spain, and Australia, in a clone of B. angularis from Argentina, and in single clones of E. ukera and R. frontalis from Germany. No significant heterogeneity was found in two other clones of B. calyciflorus from Florida and in a clone of B. variabilis from Spain. Significant heterogeneity among females of a clone could be caused by chance fluctuations during development, differences in birth order, or epimutations. This heterogeneity is an important component of a bet-hedging strategy to balance production of diapausing fertilized eggs and future population growth by female parthenogenesis. The propensity of a female to produce mictic daughters did not decrease with her age. In experiments with six different clones from four strains of B. calyciflorus, and with single clones of E. ukera and R. frontalis, only one clone of B. calyciflorus showed significant variation (ANOVA) in the proportion of mictic daughters produced in three or four successive one- or two-day periods. In this clone the proportion of mictic daughters increased with maternal age. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer research     

Clutch size, intraclutch egg size variation, and breeding strategy in the GoldcrestRegulus regulus

Summary In Fennoscandia the Goldcrest regularly lays two clutches per breeding season. Near Trondheim in Central Norway (about 63° N) the mean size of 11 first clutches was 10.4 (9–12) eggs and of 10 second/repeat clutches 9.4 (8–11) eggs. Mean egg length was 13.49 mm, breadth 10.35 mm, mean weight of freshly-laid eggs 0.78 g, volume 737 mm³ and shape index 130.6. The correlation coefficient for egg length against weight was 0.47, and for egg breadth against weight 0.77. For the four clutches studied in more detail, all egg dimensions, except the shape index, increased with the laying sequence. On average, the last-laid egg was 20.1 % heavier than the first-laid egg, or 5.6 % heavier than the mean weight for the whole clutch. The 3–5 last-laid eggs in the clutch deviated by about the same magnitude from the clutch mean. Because of the pronounced asynchrony in hatching, the size-hierarchy within a Goldcrest brood may be considerable, at least 1:5. The survival chances of the smallest young in a brood apparently depend mainly on the prevailing food supply. The primary advantage of hatching from a relatively large-sized egg is probably that such a hatchling is strong enough to secure itself a favourable feeding position in the crowded nest, in which the young often form 2–3 horizontal layers. The parental selection of food is adjusted to the size of the young, even in a brood with a marked size-hierarchy. It is concluded that the Goldcrest has evolved a brood reduction strategy.

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Gelegegröße, Variation der Eigröße und Brutstrategie beim Wintergoldhähnchen (Regulus regulus)

Zusammenfassung In Fennoskandien zeitigt das Wintergoldhähnchen regelmäßig zwei Gelege pro Brutsaison. In einem Kontrollgebiet bei Trondheim in Mittelnorwegen (ca. 63° N) betrug die mittlere Gelegegröße für 11 Erstgelege 10,4 (9–12) und für 10 Zweit- bzw. Ersatzgelege 9,4 (8–11) Eier. Die mittlere Eilänge betrug 13,5 mm, die Eibreite 10,35 mm, das mittlere Frischvollgewicht 0,78 g; das Volumen maß 737 mm³, der Eiformindex (=Länge/Breite × 100) 130,6. Die Korrelation zwischen Länge und Gewicht betrug 0,47, zwischen Breite und Gewicht 0,77. Bei vier genauer untersuchten Gelegen nahmen alle Eidimensionen mit Ausnahme des Eiformindex mit der Legefolge zu. Im Mittel war das letzte Ei 20,1 % schwerer als das erste oder 5,6 % schwerer als das mittlere Gewicht des gesamten Geleges; letzteres gilt auch für die 3 bis 5 zuletzt gelegten Eier. Als Folge des bemerkenswert asynchronen Schlupfes sind die Größenunterschiede innerhalb einer Goldhähnchenbrut beachtlich, nämlich mindestens 1:5. Die Überlebenschancen des kleinsten Jungen einer Brut hängen offensichtlich vor allem vom Nahrungsangebot ab. Der primäre Vorteil eines Jungen aus einem großen Ei liegt möglicherweise darin, daß dieser Nestling stark genug ist, um sich eine günstige Position im dichtbesetzten Nest bei den Fütterungen zu sichern. Die Jungen liegen oft in zwei bis drei Schichten übereinander. Die Nahrungswahl der Altvögel richtet sich nach der Jungengröße, sogar in einer Brut mit deutlichen Größenunterschieden. Das Wintergoldhähnchen hat wahrscheinlich die Strategie einer Brutreduzierung in Anpassung an knappes Nahrungsangebot entwickelt.

     

Body temperature and resistance to evaporative water loss in tropical Australian frogs.

Although the skin of most amphibians measured to date offers no resistance to evaporative water loss (EWL), some species, primarily arboreal frogs, produce skin secretions that increase resistance to EWL. At high air temperatures, it may be advantageous for amphibians to increase EWL as a means to decrease body temperature. In Australian hylid frogs, most species do not decrease their resistance at high air temperature, but some species with moderate resistance (at moderate air temperatures) gradually decrease resistance with increasing air temperature, and some species with high resistance (at moderate air temperatures) abruptly decrease resistance at high air temperatures. Lower skin resistance at high air temperatures decreases the time to desiccation, but the lower body temperatures allow the species to avoid their critical thermal maximum (CT(Max)) body temperatures. The body temperatures of species with low to moderate resistances to EWL that do not adjust resistance at high air temperatures do not warm to their CT(Max), although for some species, this is because they have high CT(Max) values. As has been reported previously for resistance to EWL generally, the response pattern of change of EWL at high air temperatures has apparently evolved independently among Australian hylids. The mechanisms involved in causing resistance and changes in resistance are unknown.     

Patterns of nest provisioning and parental investment in the solitary digger wasp Ammophila sabulosa

Abstract.

• 1 In nest-building wasps there is a continuum from species that provision each offspring cell with one large prey item (single provisioning) to species that provision each cell with many small items (multiple provisioning). The significance of number of prey per cell was examined during two seasons in Ammophila sabulosa (L.), a species that provisions between one and five caterpillars per cell.
• 2 There was no difference between the total weight of prey in singly-provisioned and multiply-provisioned cells: individual caterpillars placed in multiply-provisioned cells were smaller.
• 3 Small caterpillars were captured and transported to the nest faster than large ones.
• 4 If the first prey provisioned was large, the cell was usually then permanently closed. If the first prey was small, additional prey were captured, and females appeared to become more selective: additional prey were rarely large.
• 5 It is suggested that the flexible provisioning strategy used by A.sabulosa allows a wide range of prey sizes to be utilized without affecting maternal control of offspring size and sex. This could be important when available prey sizes vary temporally and spatially.
• 6 There was a male-biased first generation investment ratio. Female-producing cells were provisioned with a higher total prey weight than male-producing cells, and conversion of prey weight to adult weight was more efficient for females.
• 7 Total prey weight provisioned is probably a good indirect measure of parental time investment in A.sabulosa. Other measures, particularly number of prey provisioned, will be less accurate.

     

Cuticular hydrocarbon synthesis and its maternal provisioning to embryos in the viviparous cockroach Diploptera punctata

Embryos of the viviparous cockroach Diploptera punctata accumulate large amounts of hydrocarbon (HC) of either maternal or embryonic origin. HC synthesis and its accumulation in maternal and embryonic tissues were measured over the course of gestation. Female abdominal integument was the only tissue that synthesized appreciable amounts of HC in vitro, and did so at an increasing rate from the time of mating to mid-pregnancy, when rates of synthesis declined. The embryos synthesized HC at rates <1% those of the female, showing that the majority of HC detected in and on embryos was of maternal origin. The brood sac that houses the developing embryos did not synthesize HC in vitro, indicating that HC must be transported from the female abdominal integument to the embryos. The mass of female epicuticular HC was constant at approximately 183 microg, while her internal HC increased fourfold from mating to mid-pregnancy, then declined until parturition. The decline in internal HC reflected both declining HC synthesis in the female and greater export to the embryos, as embryonic internal HC increased 250-fold prior to parturition. An external HC coating over the oothecal covering and chorion of the embryos increased to mid-pregnancy, then declined. Unlike oviparous cockroaches, D. punctata females fed throughout the reproductive cycle, reflecting the nutritional demands of continuously provisioning the developing embryos.     

Seasonal variation in sex ratio and sexual egg dimorphism favouring daughters in first clutches of the spotless starling

We investigated possible pre‐hatching mechanisms of sex‐differential investment by females that may contribute to offspring sex‐ratio adjustment enhancing the fitness return from reproductive effort in the spotless starling (Sturnus unicolor). We found a seasonal shift in sex ratio from daughters to sons as the season advances. Furthermore, the probability of breeding at 1‐year old and recruitment into the breeding population in daughters is associated with laying date but not with mass at fledging. The reverse is true for males which rarely bred at 1‐year old. We also found that eggs containing female embryos are significantly heavier than those containing males in spite of the slight sexual dimorphism in favour of males. This suggests maternal control of provisioning, favouring daughters that may balance sibling mortality and competition with their brothers. Our results on seasonal variation in sex ratio and differential egg provisioning are consistent with an adaptive tactic in which mothers increase their reproductive return by enhancing the probability that daughters survive and breed in their first year of life.     

Effects of plant density and nutrient levels on fruit polymorphism in Atriplex sagittata

Atriplex sagittata is a heterocarpic species producing three types of fruits which differ in morphology and ecological properties. This study focused on variation in biomass allocation into particular fruit types under different density and fertilization levels. The reduction in total weight under stressful conditions was accompanied by a reduction of reproductive structures in terms of both total fruit production and mean fruit weight. Allocation of biomass to particular fruit types under different density-fertilization treatments showed considerable variation. The non-dormant, bracteolate fruit type (termed C) contributed 80–90% to the total fecundity and its production was not affected by density-fertilization treatments. The production of this fruit type is higher in the upper part of the maternal plant stem. Production of the very dormant ebracteate fruit type (termed A) increased under favourable conditions and was greater on the lower part of the stem, whereas more of the so-called type B fruit (covered by bracteoles and exhibiting dispersal and dormancy characteristics intermediate between types A and C) was produced under suboptimal conditions and was concentrated in the middle part of the plant body. This pattern contradicts to some extent the theory that deeply dormant and less dispersible fruits will be produced under stressful conditions and may be considered a new model for the behaviour of plants with more than two heterocarpic fruits. The type B fruit, intermediate in both morphology and ecological behaviour, provides a continuum between extreme strategies. Received: 8 February 1998 / Accepted: 19 November 1998     

Brood size and the cost of provisioning nestlings: interpreting Lack's hypothesis

A recent model of parental provisioning (the tradeoff model) suggests that the maximum delivery rate of food to nestlings represents a tradeoff between parental residual reproductive value and nestling survival. In contrast, Lack's hypothesis suggests that maximum provisioning rate determines brood size and therefore delivery rates are limited by shortages of food or foraging time, not by tradeoffs of parental investment. Several authors have examined the shape of the per-nestling feeding curves to test the tradeoff model against Lack's hypothesis. We show that Lack's hypothesis can produce per-nestling feeding curves consistent with the tradeoff model. Therefore, the shape of the per-nestling feeding curve cannot be used to distinguish between the models.