Ecological significance of seasonal variation in clutch-size of Arctodiaptomus bacillifer in alpine lakes of high altitudes (Val Bognanco) [Translation from: Bollettino di Zoologia 37 491-492, 1970]

This brief paper describes the significance of seasonal variation in clutch-size of the copepod Arctodiaptomus bacillifer in alpine lakes of high altitudes (Val Bognanco). Seasonal dynamics of the zooplankton of these lakes was studied during summer and autumn of 1968 and 1969and results are summarised.

The implications of adult morphology for clutch size in the flatback turtle (Natator depressa)

When the mean adult length and mean clutch volume of marine turtles are examined, a clear pattern for larger species to lay larger clutches is evident, in accord with predictions that female size constrains the available space for carrying eggs. However, when compared with this general trend, the volume of clutches laid by flatback turtles (Natator depressa) are smaller than expected. The implication is that the unusually flat morphology of flatback turtles, provides an additional constraint on their egg carrying capacity.

Clutch size for Mediterranean loggerhead turtles (Caretta caretta)

We investigated the number of eggs laid and the total clutch volume for loggerhead turtles (Caretta caretta L.) nesting on the Greek island of Cephalonia (38° 09′ N, 20° 33′ E) during 1990. Both the number of eggs laid and clutch volume were positively related to female body size, and the mean explained variation for these relationships (36·6% and 37·7%, respectively) were not significantly different. Clutch volume decreased as the season progressed because individuals laid successively smaller clutches. The inter-nesting interval (range 15–19 days) did not significantly influence the number of eggs laid or clutch volume. For individuals seen laying more than once, female size in combination with the number of a clutch in relation to all the clutches observed for that individual, together explained 65·5% of the variation in clutch volume.

Reproductive investment and optimum clutch size of Loggerhead Sea Turtles (Caretta caretta)

(1) Data for loggerhead turtles (Caretta caretta L.) nesting on the Greek island of Cephalonia were used to develop a model which predicted the optimum clutch size. (2) There was a positive linear relationship between the number of clutches into which eggs could be divided and the total time spent by a nesting turtle on the beach, and hence a negative relationship between the time invested on the beach per egg and clutch size. (3) A previous study indicated that energy expenditure for nesting turtles on land is very high, so there may be a selective pressure to maximize clutch size in order to minimize the energy expended per egg laid. As there appeared to be no counterselective pressures favouring small clutches, clutch size should be constrained by a female's egg-carryingc apacity,w hich in turn could be expected to be related to her body size. Hence, a positive relationship between clutch size and body size was predicted, and was found in the population under study.

Nest location, clutch size and nest success in the Scarlet Ibis Eudocimus ruber

Breeding success and nest-site characteristics were studied during the 1996-1997 breeding season in a colony of Scarlet Ibises Eudocimus ruber in south-eastern Brazil to test the hypothesis that nest-site characteristics and clutch size affect nest success. Two nesting pulses produced young, the earlier being more successful. Predation accounted for most failures during the first pulse, wind destruction during the second. A third pulse with few nests produced no young. Adult Ibises abandoned nests when they lost sight of other incubating birds. Logistic regression analysis indicated that nest success during the first pulse was positively related to clutch size, number of nests in the nest tree and in the nearest tree, and negatively to the distance to the nearest neighbour. During the second pulse there were significant negative associations between success, nest height and distance to the fourth nearest nest, and a positive association between success and nest cover. The results agree with the 'selfish herd' hypothesis, indicating that nest aggregation may increase breeding success, but the nest-site characteristics affecting success can differ over the course of one breeding season.

Drivers of clutch-size in Fork-tailed Flycatchers (Tyrannus savana) at temperate and tropical latitudes in South America

Many species of birds exhibit a latitudinal gradient in annual reproductive investment, laying more eggs and producing more nestlings at higher latitudes. However, few studies have evaluated the mechanisms that underlie such patterns and such differences in grassland birds specifically. We monitored nests of Fork-tailed Flycatchers (Tyrannus savana) over two breeding seasons at a tropical site in Bolivia (in 2010-11 and 2011-12) and three breeding seasons at a southern temperate site in Argentina (2010-11, 2011-12 and 2012-13), testing two hypotheses explaining variation in clutch-size among populations: the food-limitation hypothesis and the nest-predation hypothesis. Mean clutch-size and mean brood-size were significantly larger at the temperate study site than at the tropical site. Availability of arthropod food per individual bird was significantly higher at the temperate site. There was no relationship, positive or negative, between rates of nest predation and either clutch- or brood-size, and thus no support for the nest-predation hypothesis. We conclude that food availability explains much of the latitudinal variation in clutch-size in this species. We discuss avenues for future research on the mechanisms underlying geographical variation in the life histories of Neotropical birds.

Clutch-size adjustments and skew models: effects on reproductive partitioning and group stability

Reproductive skew theory seeks to integrate social and ecological factors thought to influence the division of reproduction among group-living animals. However, most reproductive skew models only examine interactions between individuals of the same sex. Here, we suggest that females can influence group stability and conflict among males by modifying their clutch size and may do so if they benefit from the presence of subordinate male helpers or from reduced conflict. We develop 3 models, based on concessions-based, restraint, and tug-of-war models, in which female clutch size is variable and ask when females will increase their clutch size above that which would be optimal in the absence of male-male conflict. In concessions-based and restraint models, females should increase clutch size above their optima if the benefits of staying for subordinate males are relatively low. Relatedness between males has no effect on clutch size. When females do increase clutch size, the division of reproduction between males is not influenced by relatedness and does not differ between restraint and concessions-based models. Both of these predictions are in sharp contrast to previous models. In tug-of-war models, clutch size is strongly influenced by relatedness between males, with the largest clutches, but the fewest surviving offspring, produced when males are unrelated. These 3 models demonstrate the importance of considering third-party interests in the decisions of group-living organisms.

Does clutch size evolve in response to parasites and immunocompetence?

Parasites have been argued to influence clutch size evolution, but past work and theory has largely focused on within-species optimization solutions rather than clearly addressing among-species variation. The effects of parasites on clutch size variation among species can be complex, however, because different parasites can induce age-specific differences in mortality that can cause clutch size to evolve in different directions. We provide a conceptual argument that differences in immunocompetence among species should integrate differences in overall levels of parasite-induced mortality to which a species is exposed. We test this assumption and show that mortality caused by parasites is positively correlated with immunocompetence measured by cell-mediated measures. Under life history theory, clutch size should increase with increased adult mortality and decrease with increased juvenile mortality. Using immunocompetence as a general assay of parasite-induced mortality, we tested these predictions by using data for 25 species. We found that clutch size increased strongly with adult immunocompetence. In contrast, clutch size decreased weakly with increased juvenile immunocompetence. But, immunocompetence of juveniles may be constrained by selection on adults, and, when we controlled for adult immunocompetence, clutch size decreased with juvenile immunocompetence. Thus, immunocompetence seems to reflect evolutionary differences in parasite virulence experienced by species, and differences in age-specific parasite virulence appears to exert opposite selection on clutch size evolution.

Reproductive biology of an endemic Physalaemus of the Brazilian Atlantic forest, and the trade-off between clutch and egg size in terrestrial breeders of the P. signifer group

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Reproductive biology of an endemic Physalaemus of the Brazilian Atlantic forest, and the trade-of between clutch and egg size in terrestrial breeders of the P. signifer group

Physalaemus crombiei is a small foam-nesting frog endemic to the Atlantic forest. It is a member of the P. signifer group known only from its type locality in Santa Teresa, State of Espírito Santo, and from another locality in the State of Bahia, Brazil. Most Physalaemus species are aquatic breeders, and species in the P. signifer group are the only ones exhibiting a tendency toward terrestrial reproduction in the genus. Here we describe the reproductive period, breeding site and reproductive modes of P. crombiei from a third population in the Atlantic forest, southeastern Brazil. We also investigated reproductive effort and size-fecundity relationships in females. Reproductive traits were compared to other species in the genus Physalaemus, especially those included in the P. signifer group. Physalaemus crombiei is a prolonged breeder, reproducing throughout the year with a peak of activity during the most rainy months (October-March). Males called from the humid forest foor and eggs embedded in foam nests were deposited in the water as well as on the humid foor amidst the leaf litter, or inside fallen leaves or tree holes containing rainwater on the forest foor. As expected, P. crombiei exhibited three alternative reproductive modes, as described for other species of the P. signifer group. The number of eggs produced per female varied from 91 to 250. Female body size is positively correlated both with ovary mass and clutch size (number of eggs per clutch). Variation in the number and size of eggs observed in Physalaemus species may be explained not only by female size, but also by the terrestrial reproductive mode exhibited by the species in the P. signifer group.

Seasonal variation of reproduction rates and body size of Calanus sinicus in the Southern Yellow Sea, China

Annual variations of egg production rate (EPR) and clutch size of Calanus sinicus, as well as body size of females (prosome length and dry weight), were investigated at a series of stations in the Southern Yellow Sea by onboard incubation. Calanus sinicus was spawning in all the 11 cruises investigated, and the annual variation of EPR was bimodal. Monthly average EPR was highest from May to July, respectively, 5.97, 5.36 and 6.30 eggs female(-1) d(-1), then decreased dramatically to only 1.37 eggs female(-1) d(-1) in August and attained the lowest 1.07 eggs female(-1) d(-1) in October. In November, average EPR increased again to 4.31 eggs female(-1) d(-1). Seasonal variation of clutch size was similar to EPR, except that it decreased gradually after August rather than dramatically as did EPR. Prosome length of females was maximum in May and minimum in October, but dry weight was highest in November. Monthly average EPR correlated better with prosome length than dry weight, while clutch size was rather determined by dry weight of females. It is suggested that egg production of C. sinicus was active during two discontinuous periods when both surface and bottom temperature fell into its favorite range (i.e. 10-23degreesC), and different reproductive strategies were adopted in these two reproductive peaks: other than the highest EPR, longer prosome length was also achieved by C. sinicus from May to July, while females in November developed shorter bodies but accumulated more energy for reproduction.

Does female nuptial coloration reflect egg carotenoids and clutch quality in the Two-Spotted Goby (Gobiusculus flavescens, Gobiidae)?

Summary

1 - Carotenoid-based ornamentation has often been suggested to signal mate quality, and species with such ornaments have frequently been used in studies of sexual selection.

2 - Female Gobiusculus flavescens (Two-Spotted Goby) develop colourful orange bellies during the breeding season. Belly coloration varies among mature females, and previous work has shown that nest-holding males prefer females with more colourful bellies. Because males invest heavily in offspring during incubation, the evolution of this preference can be explained if colourful females provide males with eggs of higher quality.

3 - We tested this hypothesis by allowing males to spawn with ‘colourful’ and ‘drab’ females and comparing parameters including egg carotenoid concentration, clutch size, hatchability and larval viability between groups. We also investigated relationships between egg carotenoid concentration and clutch quality parameters.

4 - Eggs from colourful females had significantly higher concentrations of total carotenoids than drab females, and photographically quantified belly coloration was a good predictor of egg carotenoid concentration.

5 - Colourful females produced slightly larger clutches, but female belly coloration was not related to any measure of clutch quality. In addition, there were no significant relationships between egg carotenoids and clutch quality. Females with high levels of egg carotenoids spawned slightly earlier, however, possibly because they were more ready to spawn or because of male mate choice.

6 - Our results call into question the generality of a causal link between egg carotenoids and offspring quality.

Size-fecundity relationships and reproductive investment in female frogs in the Pantanal, south-western Brazil

The ovarian complement of anurans exhibiting different reproductive modes is highly diverse, and intraspecific variation in reproductive output of anurans is generally related to variation in female properties and/or environmental conditions. The size-fecundity relationships, reproductive investment, and correlation between ovary mass and fat body mass were investigated for females of an anuran assemblage in the Pantanal, Brazil. Female body size was positively correlated with clutch size among seven of the eight species analyzed. However, these results seem to be influenced by seasonal variation in fecundity. Interspecific size-fecundity relationships revealed that female SVL was positively correlated with clutch size and egg size regardless of reproductive mode. Among 11 species analyzed, the reproductive investment (RI: ovary mass relative to body mass) varied from 5.5 to 18%, and there were no differences among reproductive modes and activity patterns (explosive/prolonged). RI correlated negatively with female size. Among three Leptodactylus species examined, negative correlation between ovary mass and fat body mass was verified for two species. Variations in the reproductive strategies are closely related to the reproductive activity patterns and reproductive modes exhibited by individual species, but are also influenced by environmental conditions. However, regardless of reproductive mode or activity pattern, each species seems to exhibit a reproductive strategy that allows them to respond differently to the same environmental restrictions.

Living on voles - plastic life of the Ural owl

Individuals face variable environmental conditions during their life. This may be due to migration, dispersion, environmental changes or, for example, annual variation in weather conditions. Genetic adaptation to a novel environment happens through natural selection. Phenotypic plasticity allows, however, a quick individual response to a new environment. Phenotypic plasticity may also be beneficial for individual if the environment is highly variable. For example, eggs are costly to produce. If the food conditions vary significantly between breeding seasons it is useful to be able to adjust the clutch and egg size according to the food abundance. In this thesis I use Ural owl vole system to study phenotypic plasticity and natural selection using a number of reproduction related traits. The Ural owl (Strix uralensis) is a long-lived and sedentary species. The reproduction and survival of the Ural owl, in fact their whole life, is tied to the dramatically fluctuating vole densities. Ural owls do not cause vole cycles but they have to adjust their behaviour to the rather predictable population fluctuations of these small mammals. Earlier work with this system has shown that Ural owl laying date and clutch size are plastic in relation to vole abundance. Further, individual laying date clutch size reaction norms have been shown to vary in the amount of plasticity. My work extends the knowledge of natural selection and phenotypic plasticity in traits related to reproduction. I show that egg size, timing of the onset of incubation and nest defense aggressiveness are plastic traits with fitness consequences for the Ural owl. Although egg size is in general thought to be a fixed characteristic of an individual, this highly heritable trait in the Ural owl is also remarkably plastic in relation to the changes in vole numbers, Ural owls are laying the largest eggs when their prey is most abundant. Timing of the onset of incubation is an individual-specific property and plastic in relation to clutch size. Timing of incubation is an important underlying cause for asynchronous hatching in birds. Asynchronous hatching is beneficial to offspring survival in Ural owl. Hence, timing of the onset of incubation may also be under natural selection. Ural owl females also adjust their nest defense aggressiveness according to the vole dynamics, being most aggressive in years when they produce the largest broods. Individual females show different levels of nest defense aggressiveness. Aggressiveness is positively correlated with the phenotypic plasticity of aggressiveness. As elevated nest defense aggressiveness is selected for, it may promote the plasticity of aggressive nest defense behaviour. All the studied traits are repeatable or heritable on individual level, and their expression is either directly or indirectly sensitive to changes in vole numbers. My work considers a number of important fitness-related traits showing phenotypic plasticity in all of them. Further, in two chapters I show that there is individual variation in the amount of plasticity exhibited. These findings on plasticity in reproduction related traits suggest that variable environments indeed promote plasticity.