How interacting anthropogenic pressures alter the plasticity of breeding time in two common songbirds.

Phenological adjustment is the first line of adaptive response of vertebrates when seasonality is disrupted by climate change. The prevailing response is to reproduce earlier in warmer springs, but habitat changes, such as forest degradation, are expected to affect phenological plasticity, for example, due to loss of reliability of environmental cues used by organisms to time reproduction. Relying on a two-decade, country-level capture-based monitoring of common songbirds' reproduction, we investigated how habitat anthropization, here characterized by the rural-urban and forest-farmland gradients, affected the average phenology and plasticity to local temperature in two common species, the great tit Parus major and the blue tit Cyanistes caeruleus. We built a hierarchical model that simultaneously estimated fledging phenology and its response to spring temperatures based on the changes in the proportion of juveniles captured over the breeding season. Both species fledge earlier in warmer sites (blue tit: 2.94 days/°C, great tit: 3.83 days/°C), in warmer springs (blue tit: 2.49 days/°C, great tit: 2.75 days/°C) and in most urbanized habitats (4 days for blue tit and 2 days for great tit). The slope of the reaction norm of fledging phenology to spring temperature varied across sites in both species, but this variation was explained by habitat anthropization only in the deciduous forest specialist, the blue tit. In this species, the responses to spring temperature were shallower in agricultural landscapes and slightly steeper in more urban areas. Habitat anthropization did not explain variation in the slope of the reaction norm in the habitat-generalist species (great tit), for which mean fledgling phenology and plasticity were correlated (i.e., steeper response in later sites). The effects of habitat change on phenological reaction norms provide another way through which combined environmental degradations may threaten populations' persistence, to an extent depending on species and on the changes in their prey phenology and abundance.

Modifications of the rain forest frugivore community are associated with reduced seed removal at the community level.

Tropical rain forests worldwide are under increasing pressure from human activities, which are altering key ecosystem processes such as plant-animal interactions. However, while the direct impact of anthropogenic disturbance on animal communities has been well studied, the consequences of such defaunation for mutualistic interactions such as seed dispersal remains chiefly understood at the plant species level. We asked whether communities of endozoochorous tree species had altered seed removal in forests affected by hunting and logging and if this could be related to modifications of the frugivore community. At two contrasting forest sites in French Guiana, Nouragues (protected) and Montagne de Kaw (hunted and partly logged), we focused on four families of animal-dispersed trees (Sapotaceae, Myristicaceae, Burseraceae, and Fabaceae), which represent 88% of all endozoochorous trees that were fruiting at the time and location of the study. We assessed the abundance of the seed dispersers and predators of these four focal families by conducting diurnal distance sampling along line transects. Densities of several key seed dispersers such as large-bodied primates were greatly reduced at Montagne de Kaw, where the specialist frugivore Ateles paniscus is probably extinct. In parallel, we estimated seed removal rates from fruit and seed counts conducted in 1-m2 quadrats placed on the ground beneath fruiting trees. Seed removal rates dropped from 77% at Nouragues to 47 % at Montagne de Kaw, confirming that the loss of frugivores associated with human disturbance impacts seed removal at the community level. In contrast to Sapotaceae, whose seeds are dispersed by mammals only, weaker declines in seed removal for Burseraceae and Myristicaceae suggest that some compensation may occur for these bird- and mammal-dispersed families, possibly because of the high abundance of Toucans at the disturbed site. The defaunation process currently occurring across many tropical forests could dramatically reduce the diversity of entire communities of animal-dispersed trees through seed removal limitation.

State transitions: a major mortality risk for seasonal species.

Ageing results from the accumulation of multifactorial damage over time. However, the temporal distribution of this damage remains unknown. In seasonal species, transitions between seasons are critical periods of massive physiological remodelling. We hypothesised that these recurrent peaks of physiological remodelling are costly in terms of survival. We tested whether captive small primates exposed to an experimentally increased frequency of seasonal transitions die sooner than individuals living under natural seasonality. The results show that experiencing one additional season per year increases the mortality hazard by a factor of 3 to 4, whereas the expected number of seasons lived is only slightly impacted by the seasonal rhythm. These results demonstrate that physiological transitions between periods of high and low metabolic activity represent a major mortality risk for seasonal organisms, which has been ignored until now.

When to initiate torpor use? Food availability times the transition to winter phenotype in a tropical heterotherm.

Timing of winter phenotype expression determines individual chances of survival until the next reproductive season. Environmental cues triggering this seasonal phenotypic transition have rarely been investigated, although they play a central role in the compensation of climatic fluctuations via plastic phenotypic adjustments. Initiation of winter daily torpor use-a widespread energy-saving phenotype-could be primarily timed according to anticipatory seasonal cues (anticipatory cues hypothesis), or flexibly fine-tuned according to actual energy availability (food shortage hypothesis). We conducted a food supplementation experiment on wild heterothermic primates (grey mouse lemurs, Microcebus murinus) at the transition to the food-limited dry season, i.e. the austral winter. As expected under the food shortage hypothesis, food-supplemented individuals postponed the seasonal transition to normal torpor use by 1-2 month(s), spent four times less torpid, and exhibited minimal skin temperature 6 °C higher than control animals. This study provides the first in situ experimental evidence that food availability, rather than abiotic cues, times the launching of torpor use. Fine-tuning of the timing of seasonal phenotypic transitions according to actual food shortage should provide heterotherms with a flexible adaptive mechanism to survive unexpected environmental fluctuations.

Bending properties of a macroalga: Adaptation of Peirce's cantilever test for in situ measurements of Laminaria digitata (Laminariaceae).

• Premise of the study: The mechanical properties of a plant are key variables governing the interaction between the plant and its environment. Thus, measuring variables such as the flexural rigidity (bending) of a plant element is necessary to understand and predict the plant-flow interaction. However, plant elements such as macrophyte blades can be relatively thin and flexible, thus difficult to characterize. Different adaptations of the classical 3-point bending tests can also affect the interpretation of the flexural rigidity of an element. A simple, robust, method is newly applied to a biomaterial and validated here as an alternative to measure flexural rigidity of thin, flexible plant elements.• Methods: Based on a bending test procedure developed for the textile industry, an apparatus for in-situ measurements was developed and compared with other normalized methods, then used in a field test on the blade of a marine macroalga (Laminaria digitata) to assess its suitability to measure the bending modulus of a biomaterial.• Key results: Results of the presented method on selected surrogate materials agree with a normalized cantilever method (ISO 9073-7:1998) and 3-point bending test (ISO 178:2010). Values determined for the bending moduli for blades of L. digitaria were in the typical range for algal material. The range of validity of the method is discussed.• Conclusion: By validating this method with existing norms, this study suggests a better approach to measure bending properties of different biomaterials in the field compared with more traditional bending tests and opens new possibilities.

Shallow hypothermia depends on the level of fatty acid unsaturation in adipose and liver tissues in a tropical heterothermic primate.

Optimal levels of unsaturated fatty acids have positive impacts on the use of prolonged bouts of hypothermia in mammalian hibernators, which generally have to face low winter ambient temperatures. Unsaturated fatty acids can maintain the fluidity of fat and membrane phospholipids at low body temperatures. However, less attention has been paid to their role in the regulation of shallow hypothermia, and in tropical species, which may be challenged more by seasonal energetic and/or water shortages than by low temperatures. The present study assessed the relationship between the fatty acids content of white adipose and liver tissues and the expression of shallow hypothermia in a tropical heterothermic primate, the gray mouse lemur (Microcebus murinus). The adipose tissue is the main tissue for fat storage and the liver is involved in lipid metabolism, so both tissues were expected to influence hypothermia dependence on fatty acids. As mouse lemurs largely avoid deep hypothermia (i.e. torpor) use under standard captive conditions, the expression of hypothermia was triggered by food-restricting experimental animals. Hypothermia depth increased with time, with a stronger increase for individuals that exhibited higher contents of unsaturated fatty acids suggesting that they were more flexible in their use of hypothermia. However these same animals delayed the use of long hypothermia bouts relative to individuals with a higher level of saturated fatty acids. This study evidences for the first time that body fatty acids unsaturation levels influence the regulation of body temperature not only in cold-exposed hibernators but also in tropical, facultative heterotherms.

Synchrony in adult survival is remarkably strong among common temperate songbirds across France.

Synchronous variation in demographic parameters across species increases the risk of simultaneous local extinction, which lowers the probability of subsequent recolonization. Synchrony therefore tends to destabilize meta-populations and meta-communities. Quantifying interspecific synchrony in demographic parameters, like abundance, survival, or reproduction, is thus a way to indirectly assess the stability of meta-populations and meta-communities. Moreover, it is particularly informative to identify environmental drivers of interspecific synchrony because those drivers are important across species. Using a Bayesian hierarchical multisite multispecies mark-recapture model, we investigated temporal interspecific synchrony in annual adult apparent survival for 16 common songbird species across France for the period 2001-2016. Annual adult survival was largely synchronous among species (73%, 95% credible interval [47%-94%] of the variation among years was common to all species), despite species differing in ecological niche and life history. This result was robust to different model formulations, uneven species sample sizes, and removing the long-term trend in survival. Synchrony was also shared across migratory strategies, which suggests that environmental forcing during the 4-month temperate breeding season has a large-scale, interspecific impact on songbird survival. However, the strong interspecific synchrony was not easily explained by a set of candidate weather variables we defined a priori. Spring weather variables explained only 1.4% [0.01%-5.5%] of synchrony, while the contribution of large-scale winter weather indices may have been stronger but uncertain, accounting for 12% [0.3%-37%] of synchrony. Future research could jointly model interspecific variation and covariation in breeding success, age-dependent survival, and age-dependent dispersal to understand when interspecific synchrony in abundance emerges and destabilizes meta-communities.

Birds introduced in new areas show rest disorders.

All colonizing individuals have to settle in a novel, conspecific-free environment. The introduction process should be poorly compatible with a good rest. We compared the resting behaviour of radio-tagged house sparrows (Passer domesticus) experimentally translocated into new, conspecific-free areas (introduced individuals, n = 10), with that of translocated sparrows that settled in naturally established populations (controls, n = 5). Resting habits of introduced sparrows markedly differed from those of control birds: they did not vocalize before going to roost, they changed their roosting habitat and they roosted 24 ± 7 min later and departed 13 ± 4 min earlier from the roost, resulting in a 5% rest debt. Because colonizing a new environment is expected to require heightened cognitive and physical activities, which in turn are constrained by the quality and duration of rest, we hypothesize that rest disorders and resulting cognitive impairments of newly released individuals could functionally contribute to the low post-release survival observed in (re)introduction attempts.

Modeling of senescent cell dynamics predicts a late-life decrease in cancer incidence.

Current oncogenic theories state that tumors arise from cell lineages that sequentially accumulate (epi)mutations, progressively turning healthy cells into carcinogenic ones. While those models found some empirical support, they are little predictive of intraspecies age-specific cancer incidence and of interspecies cancer prevalence. Notably, in humans and lab rodents, a deceleration (and sometimes decline) of cancer incidence rate has been found at old ages. Additionally, dominant theoretical models of oncogenesis predict that cancer risk should increase in large and/or long-lived species, which is not supported by empirical data. Here, we explore the hypothesis that cellular senescence could explain those incongruent empirical patterns. More precisely, we hypothesize that there is a trade-off between dying of cancer and of (other) ageing-related causes. This trade-off between organismal mortality components would be mediated, at the cellular scale, by the accumulation of senescent cells. In this framework, damaged cells can either undergo apoptosis or enter senescence. Apoptotic cells lead to compensatory proliferation, associated with an excess risk of cancer, whereas senescent cell accumulation leads to ageing-related mortality. To test our framework, we build a deterministic model that first describes how cells get damaged, undergo apoptosis, or enter senescence. We then translate those cellular dynamics into a compound organismal survival metric also integrating life-history traits. We address four different questions linked to our framework: can cellular senescence be adaptive, do the predictions of our model reflect epidemiological patterns observed among mammal species, what is the effect of species sizes on those answers, and what happens when senescent cells are removed? Importantly, we find that cellular senescence can optimize lifetime reproductive success. Moreover, we find that life-history traits play an important role in shaping the cellular trade-offs. Overall, we demonstrate that integrating cellular biology knowledge with eco-evolutionary principles is crucial to solve parts of the cancer puzzle.

Convenience polyandry or convenience polygyny? Costly sex under female control in a promiscuous primate.

Classic sex roles depict females as choosy, but polyandry is widespread. Empirical attempts to understand the evolution of polyandry have often focused on its adaptive value to females, whereas 'convenience polyandry' might simply decrease the costs of sexual harassment. We tested whether constraint-free female strategies favour promiscuity over mating selectivity through an original experimental design. We investigated variation in mating behaviour in response to a reversible alteration of sexual dimorphism in body mass in the grey mouse lemur, a small primate where female brief sexual receptivity allows quantifying polyandry. We manipulated body condition in captive females, predicting that convenience polyandry would increase when females are weaker than males, thus less likely to resist their solicitations. Our results rather support the alternative hypothesis of 'adaptive polyandry': females in better condition are more polyandrous. Furthermore, we reveal that multiple mating incurs significant energetic costs, which are strikingly symmetrical between the sexes. Our study shows that mouse lemur females exert tight control over mating and actively seek multiple mates. The benefits of remating are nevertheless not offset by its costs in low-condition females, suggesting that polyandry is a flexible strategy yielding moderate fitness benefits in this small mammal.

Torpor use during gestation and lactation in a primate.

Torpor is an energy-saving mechanism that allows endotherms to overcome energetic challenges. Torpor should be avoided during reproduction because of potential incompatibility with offspring growth. To test if torpor can be used during gestation and lactation to compensate for food shortage, we exposed reproductive female grey mouse lemurs (Microcebus murinus), a heterothermic primate, to different levels of food availability. Torpor use was characterised by daily skin temperature profiles, and its energetic outcome was assessed from changes in body mass. Food shortage triggered torpor during the end of the gestation period (n = 1), ranging from shallow in response to 40% food restriction to deep daily torpor in response to 80% restriction. During the early period of lactation, females fed ad libitum (n = 2) or exposed to a 40% restriction (n = 4) remained normothermic; but 80% food restricted females (n = 5) gave priority to energy saving, increasing the frequency and depth of torpor bouts. The use of torpor was insufficient to compensate for 80% energetic shortage during lactation resulting in loss of mass from the mother and delayed growth in the pups. This study provides the first evidence that a heterothermic primate can use torpor to compensate for food shortages even during reproduction. This physiological flexibility likely evolved as a response to climate-driven fluctuations in food availability in Madagascar.

Mutual mate choice in a female-dominant and sexually monomorphic primate.

Sexual dimorphism is common in polygynous species, where intrasexual competition is often thought to drive the evolution of large male body size, and in turn, male behavioral dominance over females. In Madagascar, the entire lemur radiation, which embraces diverse mating systems, lacks sexual dimorphism and exhibits frequent female dominance over males. The evolution of such morphological and behavioral peculiarities, often referred to as "the lemur syndrome," has proven difficult to understand. Among other hypotheses, a potential role of intersexual selection has been repeatedly proposed but hardly ever tested. Here, we investigate whether female choice favors small and compliant males, and whether male choice favors large females in captive gray mouse lemurs (Microcebus murinus). Detailed analysis of a combination of behavioral observations and hormonal data available for both sexes shows that (1) females accept more matings from males with higher fighting abilities, (2) males adjust their investment in intrasexual competition to female fertility, and (3) both male and female strategies are weakly influenced by the body mass of potential partners, in directions contradicting our predictions. These results do not suggest a prominent role of intersexual selection in the evolution and maintenance of the lemur syndrome but rather point to alternative mechanisms relating to male-male competition, specifically highlighting an absence of relationship between male body mass and fighting ability. Finally, our findings add to the growing body of evidence suggesting flexible sex roles, by showing the expression of mutual mate choice in a female-dominant, sexually monomorphic and promiscuous primate.

Research effort on birds' reservoir host potential for Lyme borreliosis: A systematic review and perspectives.

Zoonotic tick-borne diseases threat human and animal health. Understanding the role of hosts in the production of infected ticks in an epidemiological system is essential to be able to design effective measures to reduce the exposure of humans and animals to infectious tick bites. The reservoir host potential, that is, number of infected ticks produced by a host species, depends on three components: tick production, realized reservoir competence and host density. The parameters and factors that determine the reservoir host potential need to be characterized to achieve a robust understanding of the dynamics of pathogen-tick-host systems, and thus to mitigate the acarological risk of emerging infections. Few studies have investigated the role of birds in the local spread of Lyme borreliosis Borrelia. Knowledge of the research effort on the reservoir host potential of birds in Lyme borreliosis Borrelia circulation is necessary to prioritize future research on this topic. We provide a systematic review of the research effort on components of the reservoir host potential of wild birds for Lyme borreliosis Borrelia circulation, and factors that modulate these components in the European epidemiological system. Our review of 242 selected publications showed that tick production has been 1.4 and 21 times more studied than realized reservoir competence and bird density respectively. Only one study achieved to characterize the global host reservoir potential of birds in a given epidemiological system. Investigated factors were mostly related to bird species identity, individual characteristics of birds and tick characteristics, whereas the influence of bird life-history traits have been largely under-investigated. Because simultaneous characterization of all parameters is notoriously complex, interdisciplinary research is needed to combine and accumulate independent field and laboratory investigations targeting each parameter on specific epidemiological system or host species. This can help gain an integrated appraisal of the functioning of the studied system at a local scale.

Diversity of Tick-Borne Pathogens in Tick Larvae Feeding on Breeding Birds in France.

Birds play a role in maintaining tick-borne diseases by contributing to the multiplication of ticks and pathogens on a local scale during the breeding season. In the present study, we describe the diversity of tick and pathogen species of medical and veterinary importance in Europe hosted by 1040 captured birds (56 species) during their breeding season in France. Of the 3114 ticks collected, Ixodes ricinus was the most prevalent species (89.5%), followed by I. frontalis (0.8%), I. arboricola (0.7%), Haemaphysalis concinna (0.5%), H. punctata (0.5%), Hyalomma spp. (0.2%), and Rhipicephalus spp. (0.06%). Because they may be representative of the bird infection status for some pathogen species, 1106 engorged tick larvae were screened for pathogens. Borrelia burgdorferi sensu lato was the most prevalent pathogen genus in bird-feeding larvae (11.7%), followed by Rickettsia spp. (7.4%), Anaplasma spp. (5.7%), Babesia spp. (2.3%), Ehrlichia spp. (1.4%), and B. miyamotoi (1%). Turdidae birds (Turdus merula and T. philomelos), Troglodytes troglodytes, and Anthus trivialis had a significantly higher prevalence of B. burgdorferi s.l.-infected larvae than other pathogen genera. This suggests that these bird species could act as reservoir hosts for B. burgdorferi s.l. during their breeding season, and thus play an important role in acarological risk.

Physiological flexibility and acclimation to food shortage in a heterothermic primate.

As ecosystems undergo changes worldwide, physiological flexibility is likely to be an important adaptive response to increased climate instability. Extreme weather fluctuations impose energetical constraints such as unpredictable food shortage. We tested how grey mouse lemurs (Microcebus murinus) could adjust their daily heterothermy and locomotor activity to these 'energetic accidents' with a food restriction experiment. The experimental design consisted of acute calorie restriction (2 weeks, 80% restriction) in the middle of winter, after a fattening season with low (11 weeks, 40% restriction) versus high (ad libitum) food availability. This design aimed at simulating the combined effects of the quality of the fattening season (acclimation effect) and a sudden, severe food shortage during the lean season. Hour of start and duration of torpor were the most flexible components of energy savings, increasing in response to the acute food shortage with facilitation by chronic restriction (acclimation effect). Modulations of locomotor activity did not support the hypothesis of energy savings, as total locomotor activity was not reduced. Nonetheless, acutely restricted individuals modified their temporal pattern of locomotor activity according to former food availability. We provide the first experimental evidence of different temporal levels of flexibility of energy-saving mechanisms in a heterotherm exposed to food shortage. The acclimation effect of past food scarcity suggests that heterothermic organisms are better able to respond to unpredicted food scarcity during the lean season. The flexible control of energy expenditure conferred by heterothermy may facilitate the plastic response of heterothermic species to more frequent climatic hazards.

Flexibility Is Costly: Hidden Physiological Damage From Seasonal Phenotypic Transitions in Heterothermic Species.

Heterothermy allows organisms to cope with fluctuating environmental conditions. The use of regulated hypometabolism allows seasonal heterothermic species to cope with annual resource shortages and thus to maximize survival during the unfavorable season. This comes with deep physiological remodeling at each seasonal transition to allow the organism to adjust to the changing environment. In the wild, this adaptation is highly beneficial and largely overcomes potential costs. However, researchers recently proposed that it might also generate both ecological and physiological costs for the organism. Here, we propose new perspectives to be considered when analyzing adaptation to seasonality, in particular considering these costs. We propose a list of putative costs, including DNA damage, inflammatory response to fat load, brain and cognitive defects, digestive malfunction and immunodeficiency, that should receive more attention in future research on physiological seasonality. These costs may only be marginal at each transition event but accumulate over time and therefore emerge with age. In this context, studies in captivity, where we have access to aging individuals with limited extrinsic mortality (e.g., predation), could be highly valuable to experimentally assess the costs of physiological flexibility. Finally, we offer new perspectives, which should be included in demographic models, on how the adaptive value of physiological flexibility could be altered in the future in the context of global warming.

Advantages of volunteer-based biodiversity monitoring in Europe.

Without robust and unbiased systems for monitoring, changes in natural systems will remain enigmatic for policy makers, leaving them without a clear idea of the consequences of any environmental policies they might adopt. Generally, biodiversity-monitoring activities are not integrated or evaluated across any large geographic region. The EuMon project conducted the first large-scale evaluation of monitoring practices in Europe through an on-line questionnaire and is reporting on the results of this survey. In September 2007 the EuMon project had documented 395 monitoring schemes for species, which represents a total annual cost of about 4 million euro, involving more than 46,000 persons devoting over 148,000 person-days/year to biodiversity-monitoring activities. Here we focused on the analysis of variations of monitoring practices across a set of taxonomic groups (birds, amphibians and reptiles, mammals, butterflies, plants, and other insects) and across 5 European countries (France, Germany, Hungary, Lithuania, and Poland). Our results suggest that the overall sampling effort of a scheme is linked with the proportion of volunteers involved in that scheme. Because precision is a function of the number of monitored sites and the number of sites is maximized by volunteer involvement, our results do not support the common belief that volunteer-based schemes are too noisy to be informative. Just the opposite, we believe volunteer-based schemes provide relatively reliable data, with state-of-the-art survey designs or data-analysis methods, and consequently can yield unbiased results. Quality of data collected by volunteers is more likely determined by survey design, analytical methodology, and communication skills within the schemes rather than by volunteer involvement per se.

Does urbanization cause stress in wild birds during development? Insights from feather corticosterone levels in juvenile house sparrows (Passer domesticus).

Urban landscapes are associated with abiotic and biotic environmental changes that may result in potential stressors for wild vertebrates. Urban exploiters have physiological, morphological, and behavioral adaptations to live in cities. However, there is increasing evidence that urban exploiters themselves can suffer from urban conditions, especially during specific life-history stages. We looked for a link between the degree of urbanization and the level of developmental stress in an urban exploiter (the house sparrow, Passer domesticus), which has recently been declining in multiple European cities (e.g., London, UK). Specifically, we conducted a large-scale study and sampled juvenile sparrows in 11 urban and rural sites to evaluate their feather corticosterone (CORT) levels. We found that juvenile feather CORT levels were positively correlated with the degree of urbanization, supporting the idea that developing house sparrows may suffer from urban environmental conditions. However, we did not find any correlation between juvenile feather CORT levels and body size, mass, or body condition. This suggests either that the growth and condition of urban sparrows are not impacted by elevated developmental CORT levels, or that urban sparrows may compensate for developmental constraints once they have left the nest. Although feather CORT levels were not correlated with baseline CORT levels, we found that feather CORT levels were slightly and positively correlated with the CORT stress response in juveniles. This suggests that urban developmental conditions may potentially have long-lasting effects on stress physiology and stress sensitivity in this urban exploiter.

Female reproduction bears no survival cost in captivity for gray mouse lemurs.

The survival cost of reproduction has been revealed in many free-ranging vertebrates. However, recent studies on captive populations failed to detect this cost. Theoretically, this lack of survival/reproduction trade-off is expected when resources are not limiting, but these studies may have failed to detect the cost, as they may not have fully accounted for potential confounding effects, in particular interindividual heterogeneity. Here, we investigated the effects of current and past reproductive effort on later survival in captive females of a small primate, the gray mouse lemur. Survival analyses showed no cost of reproduction in females; and the pattern was even in the opposite direction: the higher the reproductive effort, the higher the chances of survival until the next reproductive event. These conclusions hold even while accounting for interindividual heterogeneity. In agreement with aforementioned studies on captive vertebrates, these results remind us that reproduction is expected to be traded against body maintenance and the survival prospect only when resources are so limiting that they induce an allocation trade-off. Thus, the cost of reproduction has a major extrinsic component driven by environmental conditions.

Ancestrality and evolution of trait syndromes in finches (Fringillidae).

Species traits have been hypothesized by one of us (Ponge, 2013) to evolve in a correlated manner as species colonize stable, undisturbed habitats, shifting from "ancestral" to "derived" strategies. We predicted that generalism, r-selection, sexual monomorphism, and migration/gregariousness are the ancestral states (collectively called strategy A) and evolved correlatively toward specialism, K-selection, sexual dimorphism, and residence/territoriality as habitat stabilized (collectively called B strategy). We analyzed the correlated evolution of four syndromes, summarizing the covariation between 53 traits, respectively, involved in ecological specialization, r-K gradient, sexual selection, and dispersal/social behaviors in 81 species representative of Fringillidae, a bird family with available natural history information and that shows variability for all these traits. The ancestrality of strategy A was supported for three of the four syndromes, the ancestrality of generalism having a weaker support, except for the core group Carduelinae (69 species). It appeared that two different B-strategies evolved from the ancestral state A, both associated with highly predictable environments: one in poorly seasonal environments, called B1, with species living permanently in lowland tropics, with "slow pace of life" and weak sexual dimorphism, and one in highly seasonal environments, called B2, with species breeding out-of-the-tropics, migratory, with a "fast pace of life" and high sexual dimorphism.