Extra Nestlings That Are Condemned to Die Increase Reproductive Success in Hoopoes.

AbstractThe adaptive value of routinely laying more eggs than can be successfully fledged has intrigued evolutionary biologists for decades. Extra eggs could, for instance, be adaptive as insurance against hatching failures. Moreover, because recent literature demonstrates that sibling cannibalism is frequent in the Eurasian hoopoe (Upupa epops), producing extra offspring that may be cannibalized by older siblings might also be adaptive in birds. Here, directed to explore this possibility in hoopoes, we performed a food supplementation experiment during the laying period and a clutch size manipulation during the hatching stage. We found that females with the food supplement laid on average one more egg than control females and that the addition of a close-to-hatch egg at the end of the hatching period increased the intensity of sibling cannibalism and enhanced fledging success in hoopoe nests. Because none of the extra nestlings from the experimental extra eggs survived until fledging, these results strongly suggest that hoopoes obtain fitness advantages by using temporarily abundant resources to produce additional nestlings that will be cannibalized. These results therefore suppose the first experimental demonstration of the nutritive adaptive function of laying extra eggs in vertebrates with parental care.

New colonisers drive the increase of the emerging loggerhead turtle nesting in Western Mediterranean.

The loggerhead sea turtle (Caretta caretta) is sensitive to climate change and is responding by colonising the Western Mediterranean. To understand the rapid nesting increase in recent years in Spain, we sampled 45 hatchlings from 8 nests between 2016 and 2019. We sequenced a mtDNA D-loop region, genotyped 2291 SNPs using 2bRAD and collected data on clutch size, hatching success, and incubation duration. We confirmed that the colonisation has a Mediterranean and Atlantic mixed origin and we detected that these nests were laid by different females, except for two nests within the same season. Our results suggest that the recent increase in nesting is due to an increase in the number of colonising individuals rather than females born in the same area returning to breed. We hypothesize that this increase in the number of colonisers results from successful conservation efforts, feminisation of the populations of origin and earlier sexual maturation. However, the percentage of offspring females produced in Spain suggests that future returning individuals will aid to the settlement of the new population. These results allow defining the current status of this colonisation although future efforts are needed to detect remigrants to confirm the establishment of a resident population.

Transcriptome analysis of ovarian tissues highlights genes controlling energy homeostasis and oxidative stress as potential drivers of heterosis for egg number and clutch size in crossbred laying hens.

Heterosis is the major benefit of crossbreeding and has been exploited in laying hens breeding for a long time. This genetic phenomenon has been linked to various modes of nonadditive gene action. However, the molecular mechanism of heterosis for egg production in laying hens has not been fully elucidated. To fill this research gap, we sequenced mRNAs and lncRNAs of the ovary stroma containing prehierarchical follicles in White Leghorn, Rhode Island Red chickens as well as their reciprocal crossbreds that demonstrated heterosis for egg number and clutch size. We further delineated the modes of mRNAs and lncRNAs expression to identify their potential functions in the observed heterosis. Results showed that dominance was the principal mode of nonadditive expression exhibited by mRNAs and lncRNAs in the prehierarchical follicles of crossbred hens. Specifically, low-parent dominance was the main mode of mRNA expression, while high-parent dominance was the predominant mode of lncRNA expression. Important pathways enriched by genes that showed higher expression in crossbreds compared to either one or both parental lines were cell adhesion molecules, tyrosine and purine metabolism. In contrast, ECM-receptor interaction, focal adhesion, PPAR signaling, and ferroptosis were enriched in genes with lower expression in the crossbred. Protein network interaction identified nonadditively expressed genes including apolipoprotein B (APOB), transferrin, acyl-CoA synthetase medium-chain family member (APOBEC) 3, APOBEC1 complementation factor, and cathepsin S as hub genes. Among these potential hub genes, APOB was the only gene with underdominance expression common to the 2 reciprocal crossbred lines, and has been linked to oxidative stress. LncRNAs with nonadditive expression in the crossbred hens targeted natriuretic peptide receptor 1, epidermal differentiation protein beta, spermatogenesis-associated gene 22, sperm-associated antigen 16, melanocortin 2 receptor, dolichol kinase, glycine amiinotransferase, and prolactin releasing hormone receptor. In conclusion, genes with nonadditive expression in the crossbred may play crucial roles in follicle growth and atresia by improving follicle competence and increasing oxidative stress, respectively. These 2 phenomena could underpin heterosis for egg production in crossbred laying hens.

Elucidation of the genetic determination of clutch traits in Chinese local chickens of the Laiwu Black breed.

BACKGROUND: Egg laying rate (LR) is associated with a clutch, which is defined as consecutive days of oviposition. The clutch trait can be used as a selection indicator to improve egg production in poultry breeding. However, little is known about the genetic basis of clutch traits. In this study, our aim was to estimate genetic parameters and identify quantitative trait single nucleotide polymorphisms for clutch traits in 399 purebred Laiwu Black chickens (a native Chinese breed) using a genome-wide association study (GWAS). METHODS: In this work, after estimating the genetic parameters of age at first egg, body weight at first egg, LR, longest clutch until 52 week of age, first week when the longest clutch starts, last week when the longest clutch ends, number of clutches, and longest number of days without egg-laying until 52 week of age, we identified single nucleotide polymorphisms (SNPs) and potential candidate genes associated with clutch traits in Laiwu Black chickens. The restricted maximum likelihood method was used to estimate genetic parameters of clutch pattern in 399 Laiwu Black hens, using the GCTA software. RESULTS: The results showed that SNP-based heritability estimates of clutch traits ranged from 0.06 to 0.59. Genotyping data were obtained from whole genome re-sequencing data. After quality control, a total of 10,810,544 SNPs remained to be analyzed. The GWAS revealed that 421 significant SNPs responsible for clutch traits were scattered on chicken chromosomes 1-14, 17-19, 21-25, 28 and Z. Among the annotated genes, NELL2, SMYD9, SPTLC2, SMYD3 and PLCL1 were the most promising candidates for clutch traits in Laiwu Black chickens. CONCLUSION: The findings of this research provide critical insight into the genetic basis of clutch traits. These results contribute to the identification of candidate genes and variants. Genes and SNPs potentially provide new avenues for further research and would help to establish a framework for new methods of genomic prediction, and increase the accuracy of estimated genetic merit for egg production and clutch traits.

Protective Geometry and Reproductive Anatomy as Candidate Determinants of Clutch Size Variation in Pentatomid Bugs.

AbstractMany animals lay their eggs in clusters. Eggs on the periphery of clusters can be at higher risk of mortality. We asked whether the most commonly occurring clutch sizes in pentatomid bugs could result from geometrical arrangements that maximize the proportion of eggs in the cluster's interior. Although the most common clutch sizes do not correspond with geometric optimality, stink bugs do tend to lay clusters of eggs in shapes that protect increasing proportions of their offspring as clutch sizes increase. We also considered whether ovariole number, an aspect of reproductive anatomy that may be a fixed trait across many pentatomids, could explain observed distributions of clutch sizes. The most common clutch sizes across many species correspond with multiples of ovariole number. However, there are species with the same number of ovarioles that lay clutches of widely varying size, among which multiples of ovariole number are not overrepresented. In pentatomid bugs, reproductive anatomy appears to be more important than egg mass geometry in determining clutch size uniformity. In addition, our analysis demonstrates that groups of animals with little variation in ovariole number may nonetheless lay a broad range of clutch shapes and sizes.

Egg size variation in the context of polyandry: a case study using long-term field data from snowy plovers.

Gamete size variation between the sexes is central to the concept of sex roles, however, to what extent gamete size variation within the sexes relates to sex role variation remains unclear. Comparative and theoretical studies suggest that, when clutch size is invariable, polyandry is linked to a reduction of egg size, while increased female-female competition for mates favors early breeding when females cannot monopolize multiple males. To understand whether and how breeding phenology, egg size, and mating behavior are related at the individual level, we studied the reproductive histories of 424 snowy plover females observed in the wild over a 15-year period. Egg size, but not polyandry, were highly repeatable for individual females. Consistent with theoretical predictions, we found that polyandrous females were the earliest breeders and that early clutches contained smaller eggs than clutches initiated later. Neither egg size nor mating behavior showed clear signs of an age-related deterioration, on the contrary, prior experience acquired either through age or local recruitment enabled females to nest early. Taken together, these results suggest that gamete size variation is not linked to mating behavior at the individual level, and, consequently, the adaptive potential of such variation appears to be limited.

Age- and trait-dependent breeding responses to environmental variation in a short-lived songbird.

Breeding responses of organisms to environmental changes may profoundly depend on an individual's age, as an age-environment interaction may be expected to affect the expression of reproductive traits. However, little is known about how this interaction affects short-lived species that experience various environmental conditions in adulthood. Here, we used a 32-year dataset from the collared flycatcher, Ficedula albicollis, population to test whether and how the environment interacts with age to shape female age-specific reproduction. To characterise environmental variation, we applied the remotely sensed normalised difference vegetation index (NDVI), estimating vegetation productivity, and used it as a surrogate for habitat quality. Then, we analysed how within-individual age and NDVI determine patterns in laying date, clutch size, offspring production, and recruitment. We found that young and old females, but not middle-aged females, breeding under low NDVI started to lay eggs later and produced smaller clutches than females of the same age breeding under higher NDVI. No such effects were observed for offspring production or recruitment. Our study provides evidence that both an individual's age and the environmental variation experienced during adulthood may be crucial for shaping reproductive patterns in short-lived avian species, as has been found in long-lived birds.

Nest attendance, incubation constancy, and onset of incubation in dabbling ducks.

In birds, parents must provide their eggs with a safe thermal environment suitable for embryonic development. Species with uniparental incubation must balance time spent incubating eggs with time spent away from the nest to satisfy self-maintenance needs. Patterns of nest attendance, therefore, influence embryonic development and the time it takes for eggs to hatch. We studied nest attendance (time on the nest), incubation constancy (time nests were at incubation temperatures), and variation in nest temperature of 1,414 dabbling duck nests of three species in northern California. Daily nest attendance increased from only 1-3% on the day the first egg was laid to 51-57% on the day of clutch completion, and 80-83% after clutch completion through hatch. Variation in nest temperature also decreased gradually during egg-laying, and then dropped sharply (33-38%) between the day of and the day after clutch completion because increased nest attendance, particularly at night, resulted in more consistent nest temperatures. During the egg-laying stage, nocturnal nest attendance was low (13-25%), whereas after clutch completion, nest attendance was greater at night (≥87%) than during the day (70-77%) because most incubation recesses occurred during the day. Moreover, during egg-laying, nest attendance and incubation constancy increased more slowly among nests with larger final clutch sizes, suggesting that the number of eggs remaining to be laid is a major driver of incubation effort during egg-laying. Although overall nest attendance after clutch completion was similar among species, the average length of individual incubation bouts was greatest among gadwall (Mareca strepera; 779 minutes), followed by mallard (Anas platyrhynchos; 636 minutes) and then cinnamon teal (Spatula cyanoptera; 347 minutes). These results demonstrate that dabbling ducks moderate their incubation behavior according to nest stage, nest age, time of day, and clutch size and this moderation likely has important implications for egg development and overall nest success.

Investing in a nest egg: intraspecific variation in the timing of egg laying across a latitudinal gradient.

Avian reproductive strategies vary widely, and many studies of life-history variation have focused on the incubation and hatching stages of nesting. Birds make proximate decisions regarding reproductive investment during the laying stage, and these decisions likely constrain and tradeoff with other traits and subsequent behaviors. However, we know relatively little about egg-laying stage behaviors given the difficulty of locating and monitoring nest sites from the onset of laying. We used non-invasive continuous video recording to quantify variation in the egg-laying behaviors of burrowing owls (Athene cunicularia) along a 1400-km latitudinal gradient in western North America. Burrowing owls laid eggs disproportionately in the morning hours, and that tendency was strongest among first eggs in a clutch. However, selection appeared to act more strongly on laying intervals (the time between laying of consecutive eggs) than on the diel time of laying, and laying intervals varied widely among and within clutches. Laying intervals declined seasonally and with increasing clutch size but increased with increasing burrow temperature and as a function of laying stage nest attentiveness, which may be a strategy to preserve egg viability. Laying interval was positively correlated with the duration of hatching intervals, suggesting that laying interval duration is one mechanism (along with timing of incubation onset) that generates variation in hatching asynchrony. Our results lend support to two general hypotheses to explain laying schedules; selection favors laying eggs in the morning, but other selective pressures may override that pattern. These conclusions indicate that allocation decisions during laying are an important part of avian life-history strategies which are subject to energetic constraints and tradeoffs with other traits.

Coloniality and development impact intraclutch consistency of avian eggs: a comparative analysis of the individual repeatability of eggshell size and shape metrics.

In oviparous animals, egg morphology is considered an aspect of the extended phenotype of the laying mother and, thus, can be directly assessed for consistency both within and between individual females. Despite a recently renewed interest in the evolution and mechanics of avian eggshell morphology, we still lack a large-scale, comparative understanding of which egg traits are individually plastic and whether individual consistency is shaped by ecological and life history traits at the species level. Here, we aimed to understand whether intraclutch repeatability per se of different eggshell metrics is an evolving trait that responds to selection pressures from socio-ecological contexts across a diverse group of avian species for which clutch-level eggshell morphology data were available to us. Coloniality, ontogeny, and incubation period had significant impacts on the comparative patterns of relative individual repeatability among two egg metrics (i.e., size and shape), whereas other life history traits (including adult size, clutch size, nest type, migration, breeding latitude, host status of brood parasitism) did not have statistical impacts. Our results also demonstrate that individual consistency has a more widespread phylogenetic distribution than expected by evolutionary contingency across avian diversity. Future analyses should also incorporate the effects of intra- and interspecific covariation in other morphological and physiological traits on the evolution of individual consistency, especially those relevant to egg recognition, including eggshell color and maculation.

Sex-Specific Effects of Blood Serotonin on Reproductive Effort in a Small Passerine.

AbstractLaboratory animal models have shown that blood serotonin levels reflect consistent individual differences in behavioral decision-making and maternal behavior. Serotonin could also help to understand intraspecific variation in reproductive strategies, although the mechanisms are poorly understood. In this study, the relationships of plasma serotonin with breeding parameters and parental behavior were examined in wild great tits (Parus major). Females who laid eggs earlier had higher levels of serotonin in the second half of the nestling period, while no significant relationship of serotonin with clutch size, brood size, and body size was detected. In males, serotonin levels were negatively related to clutch size and brood size and positively related to body size. The association of serotonin with provisioning behavior was sex specific, and acute fear stress induced by a predator presentation did not change this relationship. Food provisioning was positively related to size-corrected serotonin levels in females and negatively related to size-corrected serotonin levels in males. These results suggest that peripheral serotonin is a sensitive marker of parental behavior and reproductive effort in wild birds, while the mechanisms linking this neurotransmitter to reproduction are probably mediated by interplay between the serotonergic system, sex hormones, and other neurotransmitters.

Anthelmintic usage on the Reproductive Parameters in Captive reared Agoutis (Dasyprocta leporina) in Trinidad and Tobago, West Indies / Uso de anti-helmíntico nos parâmetros reprodutivos em cutias criadas em cativeiro (Dasyprocta leporina) em Trinidad e Tobago, Índias Ocidentais

The agouti (Dasyprocta leporina) is a rodent that is found in the Neo-tropical region. This animal is hunted for its meat but has recently been reared in captivity as a source of meat protein in rural communities. A 20-month experiment was carried out to evaluate the effect of an anthelmintic on the reproductive performance of the agouti (Dasyprocta leporina) reared in captivity. This experiment was conducted in the humid tropics of Trinidad and Tobago. Sixteen animals (15 females, 1 male) placed in each of the two treatment groups in a completely randomized study design. In treatment 1 (T1) animals were given subcutaneous injections of Endovet Ces® (Ivermectin/Praziquantel) at 0.2 mg/kg every three months. Treatment 2 (T2) was the negative control group where animals were not exposed to an anthelmintic. Reproductive data were collected at parturition which included birth weight, litter weight, litter size and gender of offspring. The results showed that there was no statistical difference (p > 0.05) between the treatment groups with respect to birth weight, litter weight, litter size and gender. However, agoutis that were dewormed had a higher birth weight (220.24 g vs 209.1 g) and litter weight (369.8 g vs 343 g). The same values were obtained for the litter size (1.7 vs 1.7) and animals that were dewormed had a higher female offspring to male offspring (2.41:1 vs 1.11:1). This experiment demonstrated that the use of an anthelmintic strategically in the management of captive reared agoutis had no statistical effect (p > 0.05) on the reproductive parameters. Therefore, these animals can be kept in captive conditions without being dewormed and produce efficiently with proper feeding and housing management.

A cutia (Dasyprocta leporina) é um roedor que se encontra na região neo-tropical. Esse animal é caçado por sua carne, mas recentemente foi criado em cativeiro como fonte de proteína de carne em comunidades rurais. Um experimento de 20 meses foi realizado para avaliar o efeito de um anti-helmíntico no desempenho reprodutivo de cutias (Dasyprocta leporina) criadas em cativeiro. Esse experimento foi conduzido nos trópicos úmidos de Trinidad e Tobago. Dezesseis animais (15 fêmeas, 1 macho) colocados em cada um dos dois grupos de tratamento em um desenho de estudo completamente randomizado. No tratamento 1 (T1) os animais receberam injeções subcutâneas de Endovet Ces® (Ivermectina / Praziquantel) na dose de 0,2 mg / kg a cada três meses. O tratamento 2 (T2) foi o grupo de controle negativo onde os animais não foram expostos a um anti-helmíntico. Os dados reprodutivos foram coletados no parto, incluindo peso ao nascer, peso da ninhada, tamanho da ninhada e sexo da prole. Os resultados mostraram que não houve diferença estatística (p > 0,05) entre os grupos de tratamento com relação ao peso ao nascer, peso da ninhada, tamanho da ninhada e sexo. No entanto, cutias desparasitadas tiveram maior peso ao nascer (220,24 g vs. 209,1 g) e peso da ninhada (369,8 g vs. 343 g). Os mesmos valores foram obtidos para o tamanho da ninhada (1,7 vs. 1,7) e os animais que foram desparasitados tiveram uma prole feminina maior do que a prole masculina (2,41: 1 vs. 1,11: 1). Esse experimento demonstrou que o uso de anti-helmíntico estrategicamente no manejo de cutias criadas em cativeiro não teve efeito estatístico (p > 0,05) sobre os parâmetros reprodutivos. Portanto, esses animais podem ser mantidos em cativeiro sem serem vermifugados e produzir de forma eficiente com alimentação adequada e manejo do alojamento.

A global meta-analysis reveals higher variation in breeding phenology in urban birds than in their non-urban neighbours.

Cities pose a major ecological challenge for wildlife worldwide. Phenotypic variation, which can result from underlying genetic variation or plasticity, is an important metric to understand eco-evolutionary responses to environmental change. Recent work suggests that urban populations might have higher levels of phenotypic variation than non-urban counterparts. This prediction, however, has never been tested across species nor over a broad geographical range. Here, we conducted a meta-analysis of the avian literature to compare urban versus non-urban means and variation in phenology (i.e. lay date) and reproductive effort (i.e. clutch size, number of fledglings). First, we show that urban populations reproduce earlier and have smaller broods than non-urban conspecifics. Second, we show that urban populations have higher phenotypic variation in laying date than non-urban populations. This result arises from differences between populations within breeding seasons, conceivably due to higher landscape heterogeneity in urban habitats. These findings reveal a novel effect of urbanisation on animal life histories with potential implications for species adaptation to urban environments (which will require further investigation). The higher variation in phenology in birds subjected to urban disturbance could result from plastic responses to a heterogeneous environment, or from higher genetic variation in phenology, possibly linked to higher evolutionary potential.

Experimentally manipulated food availability affects offspring quality but not quantity in zebra finch meso-populations.

Food availability modulates survival, reproduction and thereby population size. In addition to direct effects, food availability has indirect effects through density of conspecifics and predators. We tested the prediction that food availability in isolation affects reproductive success by experimentally manipulating food availability continuously for 3 years in zebra finches (Taeniopygia guttata) housed in outdoor aviaries. To this end, we applied a technique that mimics natural variation in food availability: increasing the effort required per food reward without affecting diet. Lower food availability resulted in a slight delay of start of laying and fewer clutches per season, but did not affect clutch size or number of offspring reared per annum. However, increasing foraging costs substantially reduced offspring growth. Thus, food availability in isolation did not impact the quantity of offspring reared, at the expense of offspring quality. Growth declined strongly with brood size, and we interpret the lack of response with respect to offspring number as an adaptation to environments with low predictability, at the time of egg laying, of food availability during the period of peak food demand, typically weeks later. Manipulated natal brood size of the parents did not affect reproductive success. Individuals that were more successful reproducers were more likely to survive to the next breeding season, as frequently found in natural populations. We conclude that the causal mechanisms underlying associations between food availability and reproductive success in natural conditions may be more complex than usually assumed. Experiments in semi-natural meso-populations can contribute to further unravelling these mechanisms.

Hold on for One More Day: Energetic Costs of Oviductal Egg Retention in Eastern Musk Turtles (Sternotherus odoratus).

AbstractIn oviparous reptiles, parental care is often limited to the energy allocated to embryos before oviposition. Reproducing females can allocate energy toward vitellogenesis, determining the number and size of eggs, fertilization, eggshell calcification, retention of eggs within the oviduct after fertilization (oviductal egg retention), and nesting activities. Oviductal egg retention in turtles ranges from 2 wk to half a year, permitting flexibility in the timing of oviposition. The energetic cost of oviductal egg retention in eastern musk turtles (Sternotherus odoratus) was investigated by measuring the metabolism of females before and after oviposition. Gravid female metabolic rates were elevated relative to male and nongravid female metabolic rates, indicating an associated energetic cost for egg retention. Metabolism of gravid females was 40% higher before oviposition than after oviposition, and it was relatively constant across the period of oviductal egg retention. Metabolic costs associated with egg retention were correlated with clutch mass and female body mass but not with clutch size or the number of days leading up to oviposition. These results suggest that the strategy of oviductal egg retention has considerable energetic costs for eastern musk turtles but that it likely provides critical flexibility in nesting phenology.

Brood reduction caused by sibling cannibalism in Isodontia harmandi (Hymenoptera: Sphecidae), a solitary wasp species building communal brood cells.

Sibling rivalry or brood reduction prevailing within bird nests is effectively avoided in solitary aculeate nests because the larvae of wasps and bees usually develop in each brood cell. However, a solitary wasp species, Isodontia harmandi, allows us to study brood reduction in a communal brood cell, where up to a dozen larvae develop in a group relying on prey provisioned by a female wasp. To demonstrate brood reduction in this species, we collected nests at various developmental brood stages from fields for five years (2010-2015). There was a significant decrease in the brood size between the nests sampled at the egg or hatchling stages and those at later stages when analyzing only data excluding nests that were parasitized, attacked by predators, or containing deteriorated prey. In whole brood-rearing experiments, we also confirmed that brood reduction occurred in 30 of 39 nests during larval stages and in 23 nests after cocoon stage. Larval survival was affected positively by total prey weight and negatively by brood size, though cocoon survival was not affected. A third-quarter (76%) of larval death was identified as sibling cannibalism through observation by time-lapse recording on multi-larvae rearing experiments. Therefore, we conclude that brood reduction routinely occurs as a result of sibling cannibalism in I. harmandi. Additionally, as we could not detect any positive effects of clutch size on the amount of provision, female wasps might overproduce offspring due to the unpredictability of available prey resources. Differences in brood size and reduction among sex categories were undetected, except for parental provisions. Thus, sibling cannibalism may efficiently regulate brood size in communal brood cells under prey shortage.

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.

An intergenerational approach to parasitoid fitness determined using clutch size.

Parasitoids, as important natural enemies, occur in high numbers and help maintain balance in natural ecosystems. Their fitness is traditionally studied as fertility based on the number of offspring in the F1 generation. Here, using gregarious parasitoids as models, we show that this traditional approach omits one important parameter: the clutch size-body size-fertility correlation among offspring. As a result of this correlation, when females adjust the number of offspring laid in a host, they determine not only the number of offspring produced but also the body size and reproductive potential of those offspring. Although parasitoid fertility has been determined several times from clutch size, here we use Anaphes flavipes to demonstrate the use of this relationship in an upgraded intergenerational approach to parasitoid fitness. We show that with a range of hosts simultaneously utilized by female parasitoids, identical fertility in the F1 generation can lead to distinctly different fertility values in the F2 generation. Even with the same number of hosts, lower fertility in the F1 generation can generate higher fertility in the F2 generation. Our approach provides an intergenerational perspective for determining individual fitness of gregarious parasitoids and new possibilities for the modelling of parasitoid population density.

The evolution of reproductive strategies in turtles.

Optimal egg size theory assumes that changes in the egg and clutch are driven by selection, resulting in adjustments for the largest possible production of offspring with the highest fitness. Evidence supports the idea that large-bodied turtles tend to produce larger clutches with small and round eggs, while smaller species produce small clutches with large and elongated eggs. Our goals were to investigate whether egg and clutch size follow the predictions of egg size theory, if there are convergent reproductive strategies, and identify ecological factors that influence clutch and egg traits across all clades of living turtles. Using phylogenetic methods, we tested the covariance among reproductive traits, if they are convergent among different turtle lineages, and which ecological factors influence these traits. We found that both egg shape and size inversely correlate with clutch size, although with different evolutionary rates, following the predictions of the egg size theory. We also present compelling evidence for convergence among different turtle clades, over at least two reproductive strategies. Furthermore, climatic zone is the only ecological predictor to influence both egg size and fecundity, while diet only influences egg size. We conclude that egg and clutch traits in Testudines evolved independently several times across non-directly related clades that converged to similar reproductive strategies. Egg and clutch characteristics follow the trade-offs predicted by egg size theory and are influenced by ecological factors. Climatic zone and diet play an important role in the distribution of reproductive characteristics among turtles.

Differences in the temporal scale of reproductive investment across the slow-fast continuum in a passerine.

Life-history strategies differ with respect to investment in current versus 'future' reproduction, but when is this future? Under the novel 'temporality in reproductive investment hypothesis', we postulate variation should exist in the time frame over which reproductive costs are paid. Slow-paced individuals should pay reproductive costs over short (e.g. inter-annual) time scales to prevent reproductive costs accumulating, whereas fast-paced individuals should allow costs to accumulate (i.e. senescence). Using Fourier transforms, we quantify adjustments in clutch size with age, across four populations of blue tits (Cyanistes caeruleus). Fast populations had more prevalent and stronger long-term changes in reproductive investment, whereas slower populations had more prevalent short-term adjustments. Inter-annual environmental variation partly accounted for short-, but not long-term changes in reproductive investment. Our study reveals individuals differ in when they pay the cost of reproduction and that failure to partition this variation across different temporal scales and environments could underestimate reproductive trade-offs.