Extended incubation recesses in sanderlings are impacted by temperature and body condition.

Complex incubation strategies have evolved to solve the trade-off between parent survival and care for their eggs with often brief departures (recesses) that maximize egg survival, and infrequent extended recesses maximizing adult condition. Here we examined incubation behaviour of sanderlings (Calidris alba), a species that exhibits both biparental and uniparental incubation behaviour. During 11 breeding seasons in Greenland, we have quantified incubation variability with thermologgers placed in nests. We estimated the impact of environmental conditions and individual characteristics on the occurrence and the duration of recesses. We found that extended recesses are a unique feature of uniparentals, and their frequency and duration increased in colder temperatures. The relationship was mediated by body condition, with individuals in poor condition performing longer extended recesses in colder temperatures. This suggests that extended recesses may represent a shift towards self-maintenance at the expense of the egg care, allowing birds to continue incubating under unfavourable conditions. Our study illustrates how extended recesses may be a key breeding strategy to overcome high energetic costs associated with incubation. Quantifying such behavioural flexibility paves the way for tracking future behavioural responses of individuals in the face of changing environments.

Consistent variations in personality traits and their potential for genetic improvement in biocontrol agents: Trichogramma evanescens as a case study.

Improvements in the biological control of agricultural pests require improvements in the phenotyping methods used by practitioners to select efficient biological control agent (BCA) populations in industrial rearing or field conditions. Consistent inter-individual variations in behaviour (i.e. animal personality) probably affect BCA efficiency, but have never been taken into account in the development of phenotyping methods, despite having characteristics useful for phenotyping: repeatable (by definition), often heritable, etc. We developed a video-tracking method targeting animal personality traits and evaluated the feasibility of its use for genetic improvement in the BCA Trichogramma evanescens, by phenotyping 1,049 individuals from 24 isogenic lines. We found consistent individual variations in boldness, activity and exploration. Personality differences between the 24 isogenic lines suggested a genetic origin of the variations in activity and exploration (broad-sense heritability estimates of 0.06 to 0.11) and revealed a trade-off between exploration and fecundity.

Behavioural responses of breeding arctic sandpipers to ground-surface temperature and primary productivity.

Most birds incubate their eggs, which requires time and energy at the expense of other activities. Birds generally have two incubation strategies: biparental where both mates cooperate in incubating eggs, and uniparental where a single parent incubates. In harsh and unpredictable environments, incubation is challenging due to high energetic demands and variable resource availability. We studied the relationships between the incubation behaviour of sandpipers (genus Calidris) and two environmental variables: temperature and a proxy of primary productivity (i.e. NDVI). We investigated how these relationships vary between incubation strategies and across species among strategies. We also studied how the relationship between current temperature and incubation behaviour varies with previous day's temperature. We monitored the incubation behaviour of nine sandpiper species using thermologgers at 15 arctic sites between 2016 and 2019. We also used thermologgers to record the ground surface temperature at conspecific nest sites and extracted NDVI values from a remote sensing product. We found no relationship between either environmental variables and biparental incubation behaviour. Conversely, as ground-surface temperature increased, uniparental species decreased total duration of recesses (TDR) and mean duration of recesses (MDR), but increased number of recesses (NR). Moreover, small species showed stronger relationships with ground-surface temperature than large species. When all uniparental species were combined, an increase in NDVI was correlated with higher mean duration, total duration and number of recesses, but relationships varied widely across species. Finally, some uniparental species showed a lag effect with a higher nest attentiveness after a warm day while more recesses occurred after a cold day than was predicted based on current temperatures. We demonstrate the complex interplay between shorebird incubation strategies, incubation behaviour, and environmental conditions. Understanding how species respond to changes in their environment during incubation helps predict their future reproductive success.

Fluctuating Dynamics of Mate Availability Promote the Evolution of Flexible Choosiness in Both Sexes.

AbstractThe evolution of choosiness has a strong effect on sexual selection, as it promotes variance in mating success among individuals. The context in which choosiness is expressed, and therefore the associated gain and cost, is highly variable. An overlooked mechanism by current models is the rapid fluctuations in the availability and quality of partners, which generates a dynamic mating market to which each individual must optimally respond. We argue that the rapid fluctuations of the mating market are central to the evolution of optimal choosiness. Using a dynamic game approach, we investigate this hypothesis for various mating systems (characterized by different adult sex ratio and latency period combinations), allowing feedback between the choosiness and partner availability throughout a breeding season while taking into account the fine variation in individual quality. Our results indicate that quality-dependent and flexible choosiness usually evolve in both sexes for various mating systems and that a significant amount of variance in choosiness is observed, especially in males, even when courtship is costly. Accounting for the fluctuating dynamics of the mating market therefore allows envisioning a much wider range of choosiness variation in natural populations and may explain a number of recent empirical results regarding choosiness in the less common sex or its variance within sexes.

Reproductive senescence and parental effects in an indeterminate grower.

Reproductive senescence is the decrease of reproductive performance with increasing age and can potentially include trans-generational effects as the offspring produced by old parents might have a lower fitness than those produced by young parents. This negative effect may be caused either by the age of the father, mother or the interaction between the ages of both parents. Using the common woodlouse Armadillidium vulgare, an indeterminate grower, as a biological model, we tested for the existence of a deleterious effect of parental age on fitness components. Contrary to previous findings reported from vertebrate studies, old parents produced both a higher number and larger offspring than young parents. However, their offspring had lower fitness components (by surviving less, producing a smaller number of clutches or not reproducing at all) than offspring born to young parents. Our findings strongly support the existence of trans-generational senescence in woodlice and contradict the belief that old individuals in indeterminate growers contribute the most to recruitment and correspond thereby to the key life stage for population dynamics. Our work also provides rare evidence that the trans-generational effect of senescence can be stronger than direct reproductive senescence in indeterminate growers.

Magnitude and direction of parasite-induced phenotypic alterations: a meta-analysis in acanthocephalans.

Several parasite species have the ability to modify their host's phenotype to their own advantage thereby increasing the probability of transmission from one host to another. This phenomenon of host manipulation is interpreted as the expression of a parasite extended phenotype. Manipulative parasites generally affect multiple phenotypic traits in their hosts, although both the extent and adaptive significance of such multidimensionality in host manipulation is still poorly documented. To review the multidimensionality and magnitude of host manipulation, and to understand the causes of variation in trait value alteration, we performed a phylogenetically corrected meta-analysis, focusing on a model taxon: acanthocephalan parasites. Acanthocephala is a phylum of helminth parasites that use vertebrates as final hosts and invertebrates as intermediate hosts, and is one of the few parasite groups for which manipulation is predicted to be ancestral. We compiled 279 estimates of parasite-induced alterations in phenotypic trait value, from 81 studies and 13 acanthocephalan species, allocating a sign to effect size estimates according to the direction of alteration favouring parasite transmission, and grouped traits by category. Phylogenetic inertia accounted for a low proportion of variation in effect sizes. The overall average alteration of trait value was moderate and positive when considering the expected effect of alterations on trophic transmission success (signed effect sizes, after the onset of parasite infectivity to the final host). Variation in the alteration of trait value was affected by the category of phenotypic trait, with the largest alterations being reversed taxis/phobia and responses to stimuli, and increased vulnerability to predation, changes to reproductive traits (behavioural or physiological castration) and immunosuppression. Parasite transmission would thereby be facilitated mainly by changing mainly the choice of micro-habitat and the anti-predation behaviour of infected hosts, and by promoting energy-saving strategies in the host. In addition, infection with larval stages not yet infective to definitive hosts (acanthella) tends to induce opposite effects of comparable magnitude to infection with the infective stage (cystacanth), although this result should be considered with caution due to the low number of estimates with acanthella. This analysis raises important issues that should be considered in future studies investigating the adaptive significance of host manipulation, not only in acanthocephalans but also in other taxa. Specifically, the contribution of phenotypic traits to parasite transmission and the range of taxonomic diversity covered deserve thorough attention. In addition, the relationship between behaviour and immunity across parasite developmental stages and host-parasite systems (the neuropsychoimmune hypothesis of host manipulation), still awaits experimental evidence. Most of these issues apply more broadly to reported cases of host manipulation by other groups of parasites.

Pair-bonding influences affective state in a monogamous fish species.

In humans, affective states are a key component in pair-bonding, particularly in the early stage of a relationship. Pairing with a high-quality partner elicits positive affective states which, in turn, validate and reinforce the mate choice. Affective states thus strongly affect pair stability and future reproductive success. We propose generalizing the link between affective states and pair-bonding to encompass other monogamous species exhibiting biparental care, chiefly where the reproductive success of the pair critically depends on the coordination between partners. The convict cichlid Amatitlania siquia is a monogamous fish species that forms long-lasting pairs with strong cooperation between parents for parental care. In this species, we showed that females paired with their non-preferred male had lower reproductive success than those paired with their preferred male. We then transposed the judgement bias paradigm, previously used in other animal species, to assess objectively affective states in fishes. Females that were assigned their non-preferred partner exhibited pessimistic bias, which indicates a negative affective state. By contrast, females that were assigned their preferred partner did not exhibit changes in their affective state. Our results highlight that the influence of pair-bonding on affective states is not human-specific and can also be observed in non-human species.

Risk of predation makes foragers less choosy about their food.

Animals foraging in the wild have to balance speed of decision making and accuracy of assessment of a food item's quality. If resource quality is important for maximizing fitness, then the duration of decision making may be in conflict with other crucial and time consuming tasks, such as anti-predator behaviours or competition monitoring. Individuals facing the risk of predation and/or competition should adjust the duration of decision making and, as a consequence, their level of choosiness for resources. When exposed to predation, the forager could either maintain its level of choosiness for food items but accept a reduction in the amount of food items consumed or it could reduce its level of choosiness and accept all prey items encountered. Under competition risk, individuals are expected to reduce their level of choosiness as slow decision making exposes individuals to a higher risk of opportunity costs. To test these predictions, the level of choosiness of a seed-eating carabid beetle, Harpalus affinis, was examined under 4 different experimental conditions of risk: i) predation risk; ii) intraspecific competition; iii) interspecific competition; and, iv) control. All the risks were simulated using chemical cues from individual conspecifics or beetles of different species that are predatory or granivorous. Our results show that when foraging under the risk of predation, H. affinis individuals significantly reduce their level of choosiness for seeds. Reductions in level of choosiness for food items might serve as a sensible strategy to reduce both the total duration of a foraging task and the cognitive load of the food quality assessment. No significant differences were observed when individuals were exposed to competition cues. Competition, (i.e opportunity cost) may not be perceived as risk high enough to induce changes in the level of choosiness. Our results suggest that considering the amount of items consumed, alone, would be a misleading metric when assessing individual response to a risk of predation. Foraging studies should therefore also take in account the decision making process.

Personality, immune response and reproductive success: an appraisal of the pace-of-life syndrome hypothesis.

The pace-of-life syndrome (POLS) hypothesis is an extended concept of the life-history theory that includes behavioural traits. The studies challenging the POLS hypothesis often focus on the relationships between a single personality trait and a physiological and/or life-history trait. While pathogens represent a major selective pressure, few studies have been interested in testing relationships between behavioural syndrome, and several fitness components including immunity. The aim of this study was to address this question in the mealworm beetle, Tenebrio molitor, a model species in immunity studies. The personality score was estimated from a multidimensional syndrome based of four repeatable behavioural traits. In a first experiment, we investigated its relationship with two measures of fitness (reproduction and survival) and three components of the innate immunity (haemocyte concentration, and levels of activity of the phenoloxidase including the total proenzyme and the naturally activated one) to challenge the POLS hypothesis in T. molitor. Overall, we found a relationship between behavioural syndrome and reproductive success in this species, thus supporting the POLS hypothesis. We also showed a sex-specific relationship between behavioural syndrome and basal immune parameters. In a second experiment, we tested whether this observed relationship with innate immunity could be confirmed in term of differential survival after challenging by entomopathogenic bacteria, Bacillus thuringiensis. In this case, no significant relationship was evidenced. We recommend that future researchers on the POLS should control for differences in evolutionary trajectory between sexes and to pay attention to the choice of the proxy used, especially when looking at immune traits.

Hormonal and fitness consequences of behavioral assortative mating in the convict cichlid (Amatitlania siquia).

In monogamous species, partner compatibility is a key factor influencing pairing and reproductive success. In pairs with biparental care, studies have mostly focused on behavioral compatibility because it is likely to encourage the coordination of parental care within pairs, leading to a better reproductive success. Behavior modulation, throughout the breeding season and as a function of the social context, is under the regulatory feedback control of endocrine mechanisms. From this link, the idea of hormonal partner compatibility as a key component of pair cohesion and maintenance has recently emerged. Here, we investigated the link between partner behavioral assortment and their hormonal response to the pairing context. We formed assortative and disassortative pairs of convict cichlids based on their behavioral type (proactive or reactive) and took hormone and fitness measurements. Testosterone, 11-ketotestosterone, 17ß-estradiol and cortisol levels were measured from fish-holding water before and after pair formation. We found no relationship between the behavioral type of individuals and their pre-pairing hormone levels. Only the level of cortisol was affected by the partner but independently of its behavioral type. Reproductive success was not affected by the level of hormonal similarity within pairs, but we found that the variation in 11-ketotestosterone similarity between the isolated context and the pairing context was related to spawning size, and the variation in cortisol similarity to the number of fry. Behavioral compatibility does not reflect hormonal compatibility in the convict cichlid, but the relationship between reproductive success and the flexibility of hormonal similarity between partners suggests hormonal adjustment within pairs in this species.

Mismatched partners that achieve postpairing behavioral similarity improve their reproductive success.

Behavioral similarity between partners is likely to promote within-pair compatibility and to result in better reproductive success. Therefore, individuals are expected to choose a partner that is alike in behavioral type. However, mate searching is very costly and does not guarantee finding a matching partner. If mismatched individuals pair, they may benefit from increasing their similarity after pairing. We show in a monogamous fish species-the convict cichlid-that the behavioral similarity between mismatched partners can increase after pairing. This increase resulted from asymmetrical adjustment because only the reactive individual became more alike its proactive partner, whereas the latter did not change its behavior. The mismatched pairs that increased their similarity not only improved their reproductive success but also raised it up to the level of matched pairs. While most studies assume that assortative mating results from mate choice, our study suggests that postpairing adjustment could be an alternative explanation for the high behavioral similarity between partners observed in the field. It also explains why interindividual behavioral differences can be maintained within a given population.

Distribution and abundance of hematophagous flies (Glossinidae, Stomoxys, and Tabanidae) in two national parks of Gabon.

In order to minimize risks of pathogen transmission with the development of ecotourism in Gabon, a seasonal inventory has been performed in five contrasted biotopes in Ivindo (INP) and Moukalaba-Doudou (MDNP) National Parks. A total of 10,033 hematophagous flies were captured. The Glossinidae, with six different species identified, was the most abundant group and constitutes about 60% of the captured flies compared to the Stomoxys (6 species also identified) and Tabanidae with 28% and 12%, respectively. The Glossinidae showed a higher rate of capture in primary forest and in research camps. In INP, the Stomoxys showed a higher rate of capture in secondary forest and at village borders, whereas in MDNP the Stomoxys were captured more in the savannah area. Thus, each fly group seemed to reach maximum abundance in different habitats. The Glossinidae were more abundant in primary forest and near research camps while Stomoxys were more abundant in secondary forest and savannah. The Tabanidae did not show a clear habitat preference.

Female mate choice in convict cichlids is transitive and consistent with a self-referent directional preference.

INTRODUCTION: One of the most important decisions that an animal has to make in its life is choosing a mate. Although most studies in sexual selection assume that mate choice is rational, this assumption has not been tested seriously. A crucial component of rationality is that animals exhibit transitive choices: if an individual prefers option A over B, and B over C, then it also prefers A over C. RESULTS: We assessed transitivity in mate choice: 40 female convict cichlids had to make a series of binary choices between males of varying size. Ninety percent of females showed transitive choices. The mean preference index was significantly higher when a female chose between their most preferred and least preferred male (male 1 vs. male 3) compared to when they chose between males of adjacent ranks (1 vs. 2 or 2 vs. 3). The results are consistent with a simple underlying preference function leading to transitive choice: females preferred males about one third larger than themselves. This rule of thumb correctly predicted which male was preferred in 67% of the cases and the ordering in binary choices in 78% of cases. CONCLUSIONS: This study provides the first evidence for strong stochastic transitivity in a context of mate choice. The females exhibited ordinal preferences and the direction and magnitude of these preferences could be predicted from a simple rule. The females do not necessarily compare two males to choose the best; it is sufficient to use a self-referent evaluation. Such a simple decision rule has important implications for the evolution of the mating strategies and it is consistent with patterns of assortative mating repeatedly observed at population level.

Heterozygosity-fitness correlations in adult and juvenile Zenaida Dove, Zenaida aurita.

Understanding how fitness is related to genetic variation is of crucial importance in both evolutionary ecology and conservation biology. We report a study of heterozygosity-fitness correlations in a wild, noninbred population of Zenaida Doves, Zenaida aurita, based on a sample comprising 489 individuals (382 adults and 107 juveniles) typed at 13 microsatellite loci, resulting in a data set comprising 5793 genotypes. In both adults and juveniles, and irrespective of sex, no evidence was found for an effect of either multilocus or single-locus heterozygosity on traits potentially related to fitness such as foraging tactic, competitive ability, and fluctuating asymmetry. In contrast, a significant negative correlation between body condition and multilocus heterozygosity, indicative of outbreeding depression, was found in juveniles, whereas no such trend was observed in adults. However, the frequency distribution of heterozygosity did not differ between the two age classes, suggesting compensatory growth by heterozygous juveniles. We discuss our results in relation to some practical limitations associated with studies of heterozygosity-fitness correlations, and suggest that tropical bird species with allopatric divergence between island populations may provide a good biological model for the detection of outbreeding depression.

Handling stress does not reflect personality in female zebra finches (Taeniopygia guttata).

Although increasing attention is given to both the causes and consequences of variation in animal personality, the measurement of personality in captive or free-ranging individuals remains an issue. In particular, one important question concerns whether personality should be established from the existence of complex behavioral syndromes (a suite of correlated behavioral traits) or could be more easily deduced from a single variable. In that context, it has recently been suggested that handling stress, measured through breathing rate during handling, could be a good descriptor of personality, at least in passerine birds. The authors experimentally investigated to what extent handling stress was correlated with personality in female zebra finches (Taeniopygia guttata), as assessed from a suite of repeatable behavioral traits, including activity, exploratory behavior, neophobia, and reaction to startle. Although breathing rate was repeatable across individuals, it was not related to any behavioral trait, suggesting that it cannot be used to quickly predict personality, at least in zebra finches. Breathing rate during handling, in addition, was related to morphology, questioning the fact that breathing rate during handling reflects personality irrespective of individual state. The authors suggest that inference on global personality from a reduced number of traits should be performed with caution.

Rapid assessment of female preference for male size predicts subsequent choice of spawning partner in a socially monogamous cichlid fish.

Although size-assortative mating in convict cichlids, Amatitliana nigrofasciata, is supposed to result from mutual mating preference for larger individuals, female choice in relation to male size remains ambiguous. We revisited the evidence for directional preference for larger males in female convict cichlids using a classical two-way choice apparatus in which each female could decide to spend time in front of a small male or a large one. We found evidence for female preference for large males, as assessed from association preference during a 4-hour period following encounter. Furthermore, females decided to spawn in front of the initially preferred male more often than expected by chance. Our results thus confirm the existence of a directional preference for large males in female convict cichlids, and indicate that association preference measured over a short period of time can provide a quick and reliable proxy for reproductive preference in this species.

Does foreplay matter? Gammarus pulex females may benefit from long-lasting precopulatory mate guarding.

Precopulatory mate guarding (PCMG) is generally assumed to be costly for both sexes. However, males may gain by displaying long-lasting mate guarding under strong male-male competition. Surprisingly, the potential for females to benefit from being held by males has been largely overlooked in previous studies. In Gammarus pulex, an amphipod crustacean, PCMG lasts several weeks, yet females are described as bearing only cost from such male mating strategy. We investigated potential female benefits by assessing the effect of mate guarding on her intermoult duration. Unpaired females had longer intermoult duration than paired females. Intermoult duration clearly decreased when paired females engaged in early and long-lasting mate guarding. In addition, short intermoults and long-lasting mate guarding had no effect on egg number. These results highlight a potential benefit associated with PCMG for G. pulex females, suggesting that the strength of an intersexual conflict over its duration may be overestimated.

Variation between populations and local adaptation in acanthocephalan-induced parasite manipulation.

Many trophically transmitted parasites manipulate their intermediate host phenotype, resulting in higher transmission to the final host. However, it is not known if manipulation is a fixed adaptation of the parasite or a dynamic process upon which selection still acts. In particular, local adaptation has never been tested in manipulating parasites. In this study, using experimental infections between six populations of the acanthocephalan parasite Pomphorhynchus laevis and its amphipod host Gammarus pulex, we investigated whether a manipulative parasite may be locally adapted to its host. We compared adaptation patterns for infectivity and manipulative ability. We first found a negative effect of all parasite infections on host survival. Both parasite and host origins influenced infection success. We found a tendency for higher infectivity in sympatric versus allopatric combinations, but detailed analyses revealed significant differences for two populations only. Conversely, no pattern of local adaptation was found for behavioral manipulation, but manipulation ability varied among parasite origins. This suggests that parasites may adapt their investment in behavioral manipulation according to some of their host's characteristics. In addition, all naturally infected host populations were less sensitive to parasite manipulation compared to a naive host population, suggesting that hosts may evolve a general resistance to manipulation.

Speed versus accuracy in decision-making ants: expediting politics and policy implementation.

Compromises between speed and accuracy are seemingly inevitable in decision-making when accuracy depends on time-consuming information gathering. In collective decision-making, such compromises are especially likely because information is shared to determine corporate policy. This political process will also take time. Speed-accuracy trade-offs occur among house-hunting rock ants, Temnothorax albipennis. A key aspect of their decision-making is quorum sensing in a potential new nest. Finding a sufficient number of nest-mates, i.e. a quorum threshold (QT), in a potential nest site indicates that many ants find it suitable. Quorum sensing collates information. However, the QT is also used as a switch, from recruitment of nest-mates to their new home by slow tandem running, to recruitment by carrying, which is three times faster. Although tandem running is slow, it effectively enables one successful ant to lead and teach another the route between the nests. Tandem running creates positive feedback; more and more ants are shown the way, as tandem followers become, in turn, tandem leaders. The resulting corps of trained ants can then quickly carry their nest-mates; but carried ants do not learn the route. Therefore, the QT seems to set both the amount of information gathered and the speed of the emigration. Low QTs might cause more errors and a slower emigration--the worst possible outcome. This possible paradox of quick decisions leading to slow implementation might be resolved if the ants could deploy another positive-feedback recruitment process when they have used a low QT. Reverse tandem runs occur after carrying has begun and lead ants back from the new nest to the old one. Here we show experimentally that reverse tandem runs can bring lost scouts into an active role in emigrations and can help to maintain high-speed emigrations. Thus, in rock ants, although quick decision-making and rapid implementation of choices are initially in opposition, a third recruitment method can restore rapid implementation after a snap decision. This work reveals a principle of widespread importance: the dynamics of collective decision-making (i.e. the politics) and the dynamics of policy implementation are sometimes intertwined, and only by analysing the mechanisms of both can we understand certain forms of adaptive organization.

Simple learning rules to cope with changing environments.

We consider an agent that must choose repeatedly among several actions. Each action has a certain probability of giving the agent an energy reward, and costs may be associated with switching between actions. The agent does not know which action has the highest reward probability, and the probabilities change randomly over time. We study two learning rules that have been widely used to model decision-making processes in animals-one deterministic and the other stochastic. In particular, we examine the influence of the rules' 'learning rate' on the agent's energy gain. We compare the performance of each rule with the best performance attainable when the agent has either full knowledge or no knowledge of the environment. Over relatively short periods of time, both rules are successful in enabling agents to exploit their environment. Moreover, under a range of effective learning rates, both rules are equivalent, and can be expressed by a third rule that requires the agent to select the action for which the current run of unsuccessful trials is shortest. However, the performance of both rules is relatively poor over longer periods of time, and under most circumstances no better than the performance an agent could achieve without knowledge of the environment. We propose a simple extension to the original rules that enables agents to learn about and effectively exploit a changing environment for an unlimited period of time.