Tibetan birds lay larger but fewer eggs in a clutch.

How do offspring size and number vary along elevational gradients? This is a striking but rarely addressed question in life-history evolution. Here we comparatively explore it using phylogenetically paired passerine birds of lowland China vs. the Tibetan plateau spanning an elevational range of 18-4500 m. Toward the Tibetan plateau, egg size increased, clutch size decreased and total clutch volume (= clutch size × egg size) did not change, when accounting for major confounding factors. Larger eggs and smaller clutches can be a response to harsh abiotic conditions at higher elevations and the corresponding reduction in food resources required for raising young, respectively. There was a negative correlation between egg size and clutch size after controlling for elevation effect on either trait, suggesting that the trade-off contributed to the expression of these two traits across elevations, given the lack of an elevational trend in total clutch volume. Tibetan birds had a shorter breeding season, prolonged incubation and nestling period than their lowland counterparts. While fewer clutches over the short breeding season may facilitate the increase in egg size, the concentration of annual reproductive investment did not seem to act on clutch size; despite the smaller clutches, hard environments may impose greater costs of parental care on Tibetan birds. The current research provides insight into the evolution of avian life histories across elevations.

Cell orientation characteristics of the natural combs of honey bee colonies.

The cell orientation characteristics of the natural combs of honey bees have received much research attention. Although natural combs have been shown to be composed of cells with three orientations-vertical, intermediate (oblique), and horizontal-the proportion of comb cells in these three orientations varies. Knowledge of the comb-building preferences of honey bees is essential for the installation of wax comb foundations, and clarification of the cell orientation characteristics of natural honey bee combs is important for beekeeping. The purpose of this study was to determine the cell orientation characteristics of natural combs of Eastern honey bees (Apis cerana cerana) and Western honey bees (Apis mellifera ligustica). Newly built combs were used to measure the orientation of hexagonal cells and calculate the proportion of cells in different orientations relative to the total number of cells. The number of eggs laid by queens in the cells of different orientations was also determined. The orientation of cells in the natural combs of Eastern and Western honey bees was determined based on the value of the minimum included angle between the pair of parallel cell walls and a vertical line connecting the top and bottom bars of the movable frame in the geometric plane of the comb: 0°≤θ≤10°, 10°<θ≤20°, and 20°<θ≤30° for vertical, intermediate, and horizontal orientations, respectively. Natural combs were composed of cells with at least one orientation (vertical or horizontal), two orientations (vertical + intermediate (oblique) or vertical + horizontal), or three orientations (vertical + intermediate + horizontal), and the proportions of combs with the three aforementioned configurations differed. Both Eastern honey bees and Western honey bees preferred building combs with cells in a vertical orientation. Queens showed no clear preference for laying eggs in cells of specific orientations. The results of this study provide new insight that could aid the production and cutting of wax comb foundations of Eastern and Western honey bees. Our study highlights the importance of installing wax comb foundations compatible with the comb-building preferences of bees.

From individual to population level: Temperature and snow cover modulate fledging success through breeding phenology in greylag geese (Anser anser).

Local weather conditions may be used as environmental cues by animals to optimize their breeding behaviour, and could be affected by climate change. We measured associations between climate, breeding phenology, and reproductive output in greylag geese (Anser anser) across 29 years (1990-2018). The birds are individually marked, which allows accurate long-term monitoring of life-history parameters for all pairs within the flock. We had three aims: (1) identify climate patterns at a local scale in Upper Austria, (2) measure the association between climate and greylag goose breeding phenology, and (3) measure the relationship between climate and both clutch size and fledging success. Ambient temperature increased 2 °C across the 29-years study period, and higher winter temperature was associated with earlier onset of egg-laying. Using the hatch-fledge ratio, average annual temperature was the strongest predictor for the proportion of fledged goslings per season. There is evidence for an optimum time window for egg-laying (the earliest and latest eggs laid had the lowest fledging success). These findings broaden our understanding of environmental effects and population-level shifts which could be associated with increased ambient temperature and can thus inform future research about the ecological consequences of climate changes and reproductive output in avian systems.

Temporal changes in reproductive success and optimal breeding decisions in a long-distance migratory bird.

Many avian migrants have not adjusted breeding phenology to climate warming resulting in negative consequences for their offspring. We studied seasonal changes in reproductive success of the greater snow goose (Anser caerulescens atlantica), a long-distance migrant. As the climate warms and plant phenology advances, the mismatch between the timing of gosling hatch and peak nutritive quality of plants will increase. We predicted that optimal laying date yielding highest reproductive success occurred earlier over time and that the seasonal decline in reproductive success increased. Over 25 years, reproductive success of early breeders increased by 42%, producing a steeper seasonal decline in reproductive success. The difference between the laying date producing highest reproductive success and the median laying date of the population increased, which suggests an increase in the selection pressure for that trait. Observed clutch size was lower than clutch size yielding the highest reproductive success for most laying dates. However, at the individual level, clutch size could still be optimal if the additional time required to acquire nutrients to lay extra eggs is compensated by a reduction in reproductive success due to a delayed laying date. Nonetheless, breeding phenology may not respond sufficiently to meet future environmental changes induced by warming temperatures.

Condition dependent strategies of egg size variation in the Common Eider Somateria mollissima.

We analysed intraclutch egg-size variation over the laying sequence in relation to clutch size, and the relation between clutch size and female body condition, in the Common Eider Somateria mollissima during an 8-year period. The aim was to assess if eiders adjusted egg size within the laying sequence depending on clutch sizes in response to body condition, as such an adjustment could have adaptive implications on reproductive success through a size advantage for the hatchlings. The analyses were performed on a population level; and then at the individual level using data from recaptured females that changed clutch size between years. Based on 1,099 clutches from 812 individual females, population clutch size averaged 4.13 eggs (range: 1-6), with 4- and 5-egg clutchesconstituting c.70% of all clutches, taking turns in being the most represented clutch size. Clutch size was positively related to female pre-laying body condition at both the population and individual levels. Egg size varied significantly within and between clutch sizes and changes were significantly related to the laying sequence. First eggs were significantly larger in 4-egg clutches and second eggs marginally smaller than in 5-egg clutches, a pattern also found among individual females changing clutch size between years. The relationship between female pre-laying body condition and clutch size, and the intraclutch egg-size pattern indicate that both clutch size and egg size are actively adapted to the pre-breeding body condition of the female. As egg size potentially optimise reproductive success through a size advantage in hatchlings, the observed pattern of intraclutch egg-size variation suggests that female eiders possesses a finely tuned conditional dependent mechanism that may optimize reproductive output in years were females are in suboptimal body condition for breeding.

Geographic variation in maternal investment and trade-offs between egg size and clutch size in an endemic toad of the Qinghai-Tibet Plateau.

Life history theory predicts that animals often produce fewer offspring of larger size and indicate a stronger trade-off between the number and size of offspring to cope with increasing environmental stress. In order to evaluate this prediction, we tested the life history characteristics of Bufo minshanicus at eight different altitudes on the eastern Tibetan Plateau, China. Our results revealed a positive correlation between female SVL and clutch size or egg size, revealing that larger females produce more and larger eggs. However, high-altitude toads seem to favor more offspring and smaller egg sizes when removing the effect of female SVL, which is counter to theoretical predictions. In addition, there was an overall significantly negative relationship between egg size and clutch size, indicative of a trade-off between egg size and fecundity. Therefore, we suggest that higher fecundity, rather than larger egg size, is a more effective reproductive strategy for this species of anuran living at high-altitude environments.

Egg load is a cue for offspring sex ratio adjustment in a fig-pollinating wasp with male-eggs-first sex allocation.

Fig-pollinating wasps (Agaonidae) only reproduce within fig tree inflorescences (figs). Agaonid offspring sex ratios are usually female-biased and often concur with local mate competition theory (LMC). LMC predicts less female-bias when several foundresses reproduce in a fig due to reduced relatedness among intra-sexually competing male offspring. Clutch size, the offspring produced by each foundress, is a strong predictor of agaonid sex ratios and correlates negatively with foundress number. However, clutch size variation can result from several processes including egg load (eggs within a foundress), competition among foundresses and oviposition site limitation, each of which can be used as a sex allocation cue. We introduced into individual Ficus racemosa figs single Ceratosolen fusciceps foundresses and allowed each to oviposit from zero to five hours thus variably reducing their eggs-loads and then introduced each wasp individually into a second fig. Offspring sex ratio (proportion males) in second figs correlated negatively with clutch size, with males produced even in very small clutches. Ceratosolen fusciceps lay mainly male eggs first and then female eggs. Our results demonstrate that foundresses do not generally lay or attempt to lay a 'fixed' number of males, but do 'reset to zero' their sex allocation strategy on entering a second fig. With decreasing clutch size, gall failure increased, probably due to reduced pollen. We conclude that C. fusciceps foundresses can use their own egg loads as a cue to facultatively adjust their offspring sex ratios and that foundresses may also produce more 'insurance' males when they can predict increasing rates of offspring mortality.

Prolactin concentrations predict parental investment and nest survival in a free-living shorebird.

The hormonal milieu that exists during reproduction is one of the key factors influencing the trade-off between reproductive investment and self-maintenance. Much previous work in birds has focused on prolactin as a physiological mediator since prolactin is involved in the onset and maintenance of parental care. However, how prolactin relates to reproductive success in terms of altering parental behavior in wild bird populations is not fully understood. Here, we report prolactin concentrations in breeding Kentish plovers (Charadrius alexandrinus), a small shorebird with variable mating systems and parental care, as an ecological model of mating system evolution. Throughout the breeding season, we estimated the circulating prolactin concentrations in male and female plovers during incubation. In addition, we monitored parental behavior and determined the fate of nests. We found that prolactin concentrations decreased during incubation but increased with clutch completion date. In addition, males and females with high prolactin concentrations spent more time on incubation than those with low prolactin concentrations. Importantly, higher prolactin concentrations in either males or females predict higher nest survival. Our results suggest that prolactin is an indicator of parental behavior in a wild shorebird population, although additional studies including experimental manipulation of prolactin concentrations are necessary to verify this relationship.

Long-term phenology of two North American secondary cavity-nesters in response to changing climate conditions.

Wildlife populations can respond to changes in climate conditions by either adapting or moving to areas with preferred climate regimes. We studied nesting responses of two bird species, western bluebird (Sialia mexicana) and ash-throated flycatcher (Myiarchus cinerascens), to changing climate conditions (i.e., rising temperatures and increased drought stress) over 21 years in northern New Mexico. We used data from 1649 nests to assess whether the two species responded to changing climate conditions through phenological shifts in breeding time or shifts in nesting elevation. We also examined changes in reproductive output (i.e., clutch size). Our data show that western bluebirds significantly increased nesting elevation over a 19-year period by approximately 5 m per year. Mean spring temperature was the best predictor of western bluebird nesting elevation. Higher nesting elevations were not correlated with hatch dates or clutch sizes in western bluebirds, suggesting that nesting at higher elevations does not affect breeding time or reproductive output. We did not observe significant changes in nesting elevation or breeding dates in ash-throated flycatchers. Nesting higher in elevation may allow western bluebirds to cope with the increased temperatures and droughts. However, this climate niche conservatism may pose a risk for the conservation of the species if climate change and habitat loss continue to occur. The lack of significant changes detected in nesting elevation, breeding dates, and reproductive output in ash-throated flycatchers suggests a higher tolerance for changing environmental conditions in this species. This is consistent with the population increases reported for flycatchers in areas experiencing dramatic climate changes.

The traits that predict the magnitude and spatial scale of forest bird responses to urbanization intensity.

As humans continue moving to urban areas, there is a growing need to understand the effects of urban intensification on native wildlife populations. Forest species in remnant habitat are particularly vulnerable to urban intensification, but the mechanisms behind these effects are poorly understood. An understanding of how species traits, as proxies for mechanisms, mediate the effects of urban intensification on forest species can help fill this knowledge gap. Using a large point count dataset from the Second Pennsylvania Breeding Bird Atlas, we tested for the effects of species traits on the magnitude and spatial scale of the responses of 58 forest bird species to urbanization intensity in landscapes surrounding count locations. Average urbanization intensity effect size across species was -0.36 ± 0.49 (SE) and average scale of effect of urbanization intensity was 4.87 ± 5.95 km. Resident forest bird species that are granivorous or frugivorous, cavity-nesting, and have larger clutch sizes and more fledglings per clutch had more positive associations with increasing urbanization intensity in landscapes. In addition, the effect of urbanization intensity on forest birds manifested most strongly at larger spatial scales for granivorous, frugivorous, or omnivorous species that are cavity-nesting, have larger clutch sizes and longer wingspans, and flock in larger numbers. To our knowledge, the present study represents the first direct tests of the effects of species traits on both the magnitude and spatial scale of the effect of urbanization on forest birds, as well as the first evidence that migratory status, clutch size, wingspan, and fledglings per clutch are important determinants of the responses of forest birds to urbanization. We discuss the possible mechanisms underlying our results and their implications for forest bird conservation in urbanizing landscapes.

Reproductive aspects of the semi-aquatic snake Erythrolamprus miliaris (Dipsadidae: Xenodontini) in the state of Rio de Janeiro, southeastern Brazil.

We analyzed some aspects of reproduction and sexual dimorphism of the semi-aquatic dipsadid snake Erythrolamprus miliaris in the state of Rio de Janeiro, southeastern Brazil. We detected sexual dimorphism in body size (snout-vent length), with females averaging larger than males, but no sexual dimorphism in the relative length of the tail. Oviductal eggs and secondary follicles were found in all seasons, suggesting that female reproductive cycles are continuous, in spite of the tropical seasonal climate in the region. Reproductive males were present throughout the year, suggesting a continuous cycle for males as well. Clutch size averaged 10.3 ± 4.8 (range 4-21) and there was a positive and significant relationship between clutch size and female size. Compared to conspecific populations previously studied in other Atlantic Rainforest areas, populations of E. miliaris from the state of Rio de Janeiro appear more similar overall in their reproductive traits to a more northern population from the state of Bahia than to populations from further south in the states of São Paulo and Paraná.

Diet, sexual dimorphism and reproduction of sympatric racers Philodryas aestiva and Philodryas patagoniensis from the coastal Brazilian Pampa.

Herein we investigated diet, sexual dimorphism and reproductive biology of two sympatric congeneric species at the coastal Brazilian Pampa, Philodryas aestiva and P. patagoniensis. Analysis of the stomach content of the scansorial Philodryas aestiva revealed that it feeds mainly on Anura and small mammals, while it occasionally consumes Insecta, Squamata and Aves. The more terrestrial Philodryas patagoniensis presented a broader diet, composed mainly of Anura, Squamata and small mammals. Insecta and Actinopterygii were found in lower frequencies. In females of both species, snout-vent length was significantly larger than in males, while tail length/snout-vent length ratio was significantly higher in males. Males reach sexual maturity at smaller sizes than females in both species. Fecundity was higher in P. patagoniensis (3 to 24 eggs; 15 ± 8.15) than in P. aestiva (10 to 20 eggs; X = 14.50 ± 3.53). Females of P. aestiva showed secondary follicles in all seasons while eggs occurred from middle-winter to early-summer. Secondary follicles occurred throughout the year in P. patagoniensis, but concentrated in spring. Eggs occurred in late-summer, middle-autumn and spring. Both species presented positive correlations between SVL and clutch size. Differences in analyzed traits seem to reflect divergences in species morphology and use of habitat/microhabitat.

Age-related reproduction of female Mongolian racerunners (Eremias argus; Lacertidae): Evidence of reproductive senescence.

The reproductive maturation hypothesis, the terminal investment hypothesis, and the senescence hypothesis are the most extensively evaluated hypotheses proposed to explain age-related patterns of reproduction in iteroparous organisms. Here, we evaluated these hypotheses for the Mongolian racerunner (Eremias argus), a short-lived lacertid lizard, by comparing reproductive traits between females that completed reproductive cycles under the same laboratory conditions in two consecutive years (2008 and 2009). Reproductive females gained linear size (snout-vent length) not only as they got 1 year older but also during the breeding season. Larger females generally laid eggs earlier and invested more in reproduction than did smaller ones. Females switched from laying smaller eggs in the first clutch to larger eggs in the subsequent clutches but kept clutch size and postpartum body mass constant between successive clutches in a breeding season and between years. Females that laid more clutches or eggs in 2008 did not lay fewer clutches or eggs in 2009. Of the traits examined, only clutch frequency, annual fecundity, and annual reproductive output were susceptible to ageing. Specifically, the clutch frequency was reduced by 1.1 clutches, annual fecundity by 3.1 eggs and annual reproductive output by 1.0 g in 2009 compared with 2008. Our results suggest that the reproductive maturation hypothesis better explains patterns of reproduction in young or prime-aged females of E. argus, whereas the senescence hypothesis better explains reproductive patterns in old females. The terminal investment hypothesis does not apply to any trait examined because no trait value was maximized in old females.

Social parasitism as an alternative reproductive tactic in a cooperatively breeding cuckoo.

Cooperatively nesting birds are vulnerable to social parasites that lay their eggs in host nests but provide no parental care1-4. Most previous research has focused on the co-evolutionary arms race between host defences and the parasites that attempt to circumvent them5-9, but it remains unclear why females sometimes cooperate and sometimes parasitize, and how parasitic tactics arise in cooperative systems10-12. Here we show that cooperative and parasitic reproductive strategies result in approximately equal fitness pay-offs in the greater ani (Crotophaga major), a long-lived tropical cuckoo, using an 11-year dataset and comprehensive genetic data that enable comparisons of the life-histories of individual females. We found that most females in the population nested cooperatively at the beginning of the breeding season; however, of those birds that had their first nests destroyed, a minority subsequently acted as reproductive parasites. The tendency to parasitize was highly repeatable, which indicates individual specialization. Across years, the fitness pay-offs of the two strategies were approximately equal: females who never parasitized (a 'pure cooperative' strategy) laid larger clutches and fledged more young from their own nests than did birds that both nested and parasitized (a 'mixed' strategy). Our results suggest that the success of parasites is constrained by reproductive trade-offs as well as by host defences, and illustrate how cooperative and parasitic tactics can coexist stably in the same population.

Providing longer post-fledging periods increases offspring survival at the expense of future fecundity.

The cost of reproduction is a key concept in life-history trade-offs. However, our understanding of the reproductive costs is biased towards measures of reproductive effort obtained before offspring independence. During the post-fledging dependence period (PFDP), it is well known that parents feed and protect their offspring. However, the effort devoted to this reproductive activity has never been considered in the context of of the costs of reproduction. Moreover, the potential fitness benefits and costs for offspring and parents, respectively, of the duration of the PFDP are largely unknown. We estimated the duration of the PFDP over 5 years using wild common kestrels (Falco tinnunculus) and studied its association with survival probability and future parental reproductive performance. Our results show that longer PFDPs increase the survival probability of fledglings, probably due to the benefits obtained from parental care. In addition, we found that providing longer PFDPs was associated with reduced clutch sizes but not the number of fledglings in the subsequent breeding season in males. We suggest that increased parental expenditures on offspring during the PFDP may represent a potential cost of reproduction in breeding males.

Brood Reduction, Not Poecilogony, in a Vermetid Gastropod with Two Developmental Outcomes within Egg Capsules.

A small vermetid gastropod broods capsules containing nurse eggs and embryos that develop into small veligers. A few of these veligers continue development and growth while nurse eggs and developmentally arrested sibling veligers disappear. Survivors hatch as crawling pediveligers and juveniles. None of the veligers, if removed from capsules, swim in a directed way or withdraw into their shells, indicating that even the developing veligers are unsuited for extracapsular life until they can crawl. The shells of arrested veligers decalcify while their siblings grow. Few of the developmentally arrested veligers that were isolated from siblings and fed algal cells resumed detectable growth. Nurse eggs rather than cannibalism provide most of the food, but full growth of developing veligers depends on limited sharing; arrest of some siblings is a necessary adjunct of the nurse-egg feeding. Here, two developmental outcomes for larvae produced by developmental arrest of some (often termed poecilogony) serves instead as a means of brood reduction. Brood reduction is often attributed to family conflicts resulting from genetic differences. Another hypothesis is that a mother who cannot accurately sort numbers of nurse eggs and developing eggs into capsules could rely on brood reduction to adjust food for her offspring. At the extreme, an entirely random packaging would produce a binomial distribution of embryos in capsules, a very uneven distribution of food per embryo, and some capsules with no embryos. Males have yet to be found in this species, but even if reproduction is asexual, selection could still favor brood reduction.

Factors determining the occurrence of anthropogenic materials in nests of the white stork Ciconia ciconia.

Birds have been using anthropogenic materials for nest construction for the past few decades. However, there is a trade-off between the use of new nesting material, which is often linked to greater breeding success, and the higher risk of nestling mortality due to entanglement or ingestion of debris. Here, we investigate the incorporation of anthropogenic materials into nests of the white stork Ciconia ciconia, based on a long-term study of a population in Western Poland. We recorded at least one item of debris in 50 and 42% of nests at the egg and nestling stages, respectively. More debris was found in nests located in territories with higher number of anthropogenic material in the surrounding environment. We found a relationship between the age of females, the number of debris in the area surrounding a nest, and the number of debris in the nest. We found no significant effect of the total number of debris in nests on clutch size, number of fledglings, or breeding success. Studies on the influence of the age and sex of individuals in understanding this behaviour and its drivers in bird populations should be continued.

Laying sequence interacts with incubation temperature to influence rate of embryonic development and hatching synchrony in a precocial bird.

Incubation starts during egg laying for many bird species and causes developmental asynchrony within clutches. Faster development of late-laid eggs can help reduce developmental differences and synchronize hatching, which is important for precocial species whose young must leave the nest soon after hatching. In this study, we examined the effect of egg laying sequence on length of the incubation period in Wood Ducks (Aix sponsa). Because incubation temperature strongly influences embryonic development rates, we tested the interactive effects of laying sequence and incubation temperature on the ability of late-laid eggs to accelerate development and synchronize hatching. We also examined the potential cost of faster development on duckling body condition. Fresh eggs were collected and incubated at three biologically relevant temperatures (Low: 34.9°C, Medium: 35.8°C, and High: 37.6°C), and egg laying sequences from 1 to 12 were used. Length of the incubation period declined linearly as laying sequence advanced, but the relationship was strongest at medium temperatures followed by low temperatures and high temperatures. There was little support for including fresh egg mass in models of incubation period. Estimated differences in length of the incubation period between eggs 1 and 12 were 2.7 d, 1.2 d, and 0.7 d at medium, low and high temperatures, respectively. Only at intermediate incubation temperatures did development rates of late-laid eggs increase sufficiently to completely compensate for natural levels of developmental asynchrony that have been reported in Wood Duck clutches at the start of full incubation. Body condition of ducklings was strongly affected by fresh egg mass and incubation temperature but declined only slightly as laying sequence progressed. Our findings show that laying sequence and incubation temperature play important roles in helping to shape embryo development and hatching synchrony in a precocial bird.

Estimating Intermittent Individual Spawning Behavior via Disaggregating Group Data.

In order to understand fish biology and reproduction, it is important to know the fecundity patterns of individual fish, as frequently established by recording the output of mixed-sex groups of fish in a laboratory setting. However, for understanding individual reproductive health and modeling purposes it is important to estimate individual fecundity from group fecundity. We created a multistage method that disaggregates group-level data into estimates for individual-level clutch size and spawning interval distributions. The first stage of the method develops estimates of the daily spawning probability of fish. Daily spawning probabilities are then used to calculate the log likelihood of candidate distributions of clutch size. Selecting the best candidate distribution for clutch size allows for a Monte Carlo resampling of annotations of the original data which state how many fish spawned on which day. We verify this disaggregation technique by combining data from fathead minnow pairs, and checking that the disaggregation method reproduced the original clutch sizes and spawning intervals. This method will allow scientists to estimate individual clutch size and spawning interval distributions from group spawning data without specialized or elaborate experimental designs.

Artificial wetlands as tools for frog conservation: stability and variability of reproduction characteristics in Sahara frog populations in Tunisian man-made lakes.

Amphibian populations are in decline principally due to climate change, environmental contaminants, and the reduction in wetlands. Even though data concerning current population trends are scarce, artificial wetlands appear to play a vital role in amphibian conservation. This study concerns the reproductive biology of the Sahara frog over a 2-year period in four Tunisian man-made lakes. Each month, gonad state (parameters: K, GSI, LCI), fecundity, and fertility of females (using 1227 clutches) were evaluated in the field under controlled conditions. Clutches were present for 110-130 days at two of the sites, but only for 60-80 days at the other two. Maximum egg laying occurred in May, corresponding to the highest point in the gonad somatic index. Clutch densities were higher in the smaller lakes. Female fecundity was in relation to body size; mean clutch fecundity attained 1416 eggs, with no differences observed according to site. Egg fertility varied over a 1-year period, with a maximum in May followed by a decrease when water temperature was at its highest. Eggs were smaller at the beginning of spawning; maximum size was in May, which might explain the higher fertility, but no maternal influence was detected. Embryonic development was strictly dependent on temperature. The population at each site appeared as a small patch within a metapopulation in overall good health, as shown by the relative temporal stability in reproduction variables. Constructed wetlands may therefore play an important role in the conservation of amphibians, especially in semi-arid zones.