Does fitness erode in the absence of selection? An experimental test with Tribolium

In the absence of natural selection, average fitness in the population is expected to decline due to the accumulation of deleterious mutations. Replicate populations of flour beetles (Tribolium confusum) were maintained for 22 generations in the virtual absence of selection (random mating, favorable environment, excess of food, and mortality of only 3%). Larva-to-adult survivorship rates were similar in the stock population and selection-free populations. In contrast, starvation resistance of adult beetles from selection-free populations was significantly reduced (by more than 2% per generation). When tested in the favorable environment, beetles in one selection-free population had significantly slower development and smaller sizes of females than beetles from the stock population. Since such changes in these fitness components are usually maladaptive, they indicate possible erosion of fitness under relaxed selection at the rate of 0.1-0.2% per generation. No fitness erosion was detectable in the second selection-free population.     

Does a trade‐off between current reproductive success and survival affect the honesty of male signalling in species with male parental care?

Recent theory predicted that male advertisement will reliably signal investment in paternal care in species where offspring survival requires paternal care and males allocate resources between advertisement and care. However, the predicted relationship between care and advertisement depended on the marginal gains from investment in current reproductive traits. Life history theory suggests that these fitness gains are also subject to a trade‐off between current and future reproduction. Here, we investigate whether male signalling remains a reliable indicator of parental care when males allocate resources between current advertisement, paternal care and survival to future reproduction. We find that advertisement is predicted to remain a reliable signal of male care but that advertisement may cease to reliably indicate male quality because low‐quality males are predicted to invest in current reproduction, whereas higher‐quality males are able to invest in both current reproduction and survival to future reproduction.     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

Does multiple paternity affect seed mass in angiosperms? An experimental test in Dalechampia scandens

Flowers fertilized by multiple fathers may be expected to produce heavier seeds than those fertilized by a single father. However, the adaptive mechanisms leading to such differences remain unclear, and the evidence inconsistent. Here, we first review the different hypotheses predicting an increase in seed mass when multiple paternity occurs. We show that distinguishing between these hypotheses requires information about average seed mass, but also about within‐fruit variance in seed mass, bias in siring success among pollen donors, and whether siring success and seed mass are correlated. We then report the results of an experiment on Dalechampia scandens (Euphorbiaceae), assessing these critical variables in conjunction with a comparison of seed mass resulting from crosses with single vs. multiple pollen donors. Siring success differed among males when competing for fertilization, but average seed mass was not affected by the number of fathers. Furthermore, paternal identity explained only 3.8% of the variance in seed mass, and siring success was not correlated with the mass of the seeds produced. Finally, within‐infructescence variance in seed mass was not affected by the number of fathers. These results suggest that neither differential allocation nor sibling rivalry has any effect on the average mass of seeds in multiply sired fruits in D. scandens. Overall, the limited paternal effects observed in most studies and the possibility of diversification bet hedging among flowers (but not within flowers), suggest that multiple paternity within fruits or infructescence is unlikely to affect seed mass in a large number of angiosperm species.     

Does competition with wind‐pollinated species alter Echium plantagineum's attractiveness to a common pollinator Bombus terrestris?

1. In insect‐pollinated plants, pollinator attraction is influenced by flowers (e.g. number, size) and their associated rewards (e.g. pollen, nectar). These traits can depend on plant interactions. Indeed, below‐ground competition between plants can lead to a decrease in flower or reward production in insect‐pollinated species. 2. Wind‐pollinated plants, in particular, which are almost never studied in plant–pollinator networks, can alter insect‐pollinated plants' attractiveness through competition for nutrients. The response of pollinators to such changes has never been investigated. 3. A pot experiment was carried out in which an insect‐pollinated species, Echium plantagineum, was grown in binary mixture with three wind‐pollinated species selected to exert a panel of competitive interactions. Below‐ground competition was controlled using dividers limiting interspecific root competition. Floral traits of E. plantagineum (i.e. flower production, floral display size, flower size and nectar production) were measured. For each species mixture, the visits (i.e. first visit, number of visits, 10‐min sequences) of Bombus terrestris individuals released in a flight cage containing two pots were followed, one with and one without below‐ground competition. 4. Below‐ground competition significantly affected nectar's sucrose concentration but did not influence flower and nectar production. Likewise, pollinator visits were not influenced by below‐ground competition. Competitor identity significantly influenced flower and reward production of E. plantagineum, with a decrease in the presence of the most competitive wind‐pollinated species. A tendency for faster flower visitation events was also detected in the presence of the least competitive competitor. This study raises new questions regarding the influence of wind‐pollinated plants on plant–pollinator interactions.     

Can plants evolve tolerance mechanisms to heterospecific pollen effects? An experimental test of the adaptive potential in Clarkia species

Flowering plants do not occur alone and often grow in mixed‐species communities where pollinator sharing is high and interactions via pollinators can occur at pre‐ and post‐pollination stages. While the causes and consequences of pre‐pollination interactions have been well studied little is known about post‐pollination interactions via heterospecific pollen (HP) receipt, and even less about the evolutionary implications of these interactions. In particular, the degree to which plants can evolve tolerance mechanisms to the negative effects of HP receipt has received little attention. Here, we aim to fill this gap in our understanding of post‐pollination interactions by experimentally testing whether two co‐flowering Clarkia species can evolve HP tolerance, and whether tolerance to specific HP ‘genotypes’ (fine‐scale local adaptation to HP) occurs. We find that Clarkia species vary in their tolerance to HP effects. Furthermore, conspecific pollen performance and the magnitude of HP effects were related to the recipient's history of exposure to HP in C. xantiana but not in C. speciosa. Specifically, better conspecific pollen performance and smaller HP effects were observed in populations of C. xantiana plants with previous exposure to HP compared to populations without such exposure. These results suggest that plants may have the potential to evolve tolerance mechanisms to HP effects but that these may occur not from the female (stigma, style) but from the male (pollen) perspective, a possibility that is often overlooked. We find no evidence for fine‐scale local adaptation to HP receipt. Studies that evaluate the adaptive potential of plants to the negative effects of HP receipt are an important first step in understanding the evolutionary consequences of plant–plant post‐pollination interactions. Such knowledge is in turn crucial for deciphering the role of plant–pollinator interactions in driving floral evolution and the composition of co‐flowering communities.     

Does interference with mucoperiosteum and palatal bone affect craniofacial growth? An experimental study in beagles

This study was designed to assess the effects of raising mucoperiosteal flaps and exposing palatal bone at the time of palatoplasty. Using 62 beagle puppies as subjects, we tested the hypothesis that raising mucoperiosteal flaps does not interfere with craniofacial growth. We further hypothesized that the size of the area of bone exposed following palatoplasty does affect subsequent craniofacial growth. The animals were divided into four groups: two control groups (unoperated and unrepaired) and two experimental groups. In the first experimental group, two-flap palatoplasty was used to close the surgically induced palatal defect, leaving narrow strips (0 to 2.5 mm) of bone exposed lateral to the flaps. In the second group, one flap was raised to close the defect, leaving a wide area (5 to 6 mm) of palatal bone exposed on one side. Thirty-four direct craniometric measurements were analyzed. Animals that had elevation of both mucoperiosteal flaps with narrow strips of denuded bone on both sides had less severe craniofacial growth aberrations than those in which the defect was left unrepaired or was repaired with one mucoperiosteal flap leaving a wider area of bare bone exposed. These findings suggest that raising mucoperiosteal flaps is less detrimental to craniofacial growth than leaving large areas of exposed palatal bone.     

Does coping style predict optimization? An experimental test in a wild passerine bird

A number of studies have suggested that avian brood size is individually optimized. Yet, optimal reproductive decisions likely vary owing to among-individual differences in environmental sensitivity. Specifically, ‘proactive’ individuals who do not track environmental changes may be less able to produce optimal brood sizes than ‘reactive’ individuals who have more precise local environmental knowledge. To test this, we quantified exploratory behaviour (a proxy for proactivity) in a great tit (Parus major) population, manipulated brood sizes (reduced, control, enlarged) and evaluated whether individuals of dissimilar coping style differed in their level of optimization. If reactive females behaved optimally, any deviation from their original brood size should lower fitness, whereas this should not be the case for proactive females. Reactive females indeed performed best at their natural brood size, whereas proactive females performed best when raising an enlarged brood. These findings imply that proactive females produced sub-optimal brood sizes. We speculate that proactive females might (i) take decisions based on biased perception of their environment, (ii) face energetic constraints in offspring production and/or (iii) be more willing to invest into current reproduction when given the option. Our findings provide experimental evidence for coping style-related differences in optimal reproductive decisions and life-history strategies.     

Does low nutritional quality act as a plant defence? An experimental test of the slow‐growth,high‐mortality hypothesis

Abstract.  1. The slow-growth, high-mortality hypothesis was experimentally tested in this study by investigating the effects of plant quality and natural enemies on leaf-miner growth, performance, and survivorship. Two leaf miners ( Acrocercops albinatella and Brachys tesselatus ) occurring on the turkey oak Quercus laevis were studied using a factorial design that manipulated predation/parasitism pressure and plant nutritional quality.
2. Forty trees were randomly divided into four treatments: (1) control plants (nutrients and natural enemies unaltered); (2) nutrients added, natural enemies unaltered; (3) nutrients unaltered, natural enemies reduced; and (4) nutrients added and natural enemies reduced. Water content, leaf toughness, tannin concentration, and foliar nitrogen were quantified monthly for each plant, and mine growth and survivorship were assessed by tracing mines on a 2–3-day interval and by following the fates of 50 mines per species per treatment combination.
3. Fertilised plants exhibited significantly higher amounts of nitrogen, but no significant differences among treatments were observed for water content, leaf toughness, and tannin concentration. These results only partially support the slow-growth, high-mortality hypothesis, as mines were significantly smaller and developed faster on fertilised plants, but neither fertilisation nor natural enemy exclusion significantly affected mine survivorship or mortality caused by natural enemies.     

Does Neighborhood Floral Display Matter? Fruit Set in Carpenter Bee‐pollinated Heterophragma quadriloculare and Beetle‐pollinated Lasiosiphon eriocephalus

Fruit set is pollen‐limited in the self‐incompatible tree Heterophragma quadriloculare (Bignoniaceae), pollinated by long‐distance flying carpenter bees, and in the self‐compatible shrub Lasiosiphon eriocephalus (Thymdeaceae), pollinated by weak‐flying, sedentary beetles. We studied a single H. quadriloculare population over high and low flowering years and found no difference in bee visitation rates between these years. For H. quadriloculare, neighborhood floral display did not make a significant contribution to reproductive success. We investigated dense and sparse L. eriocephalus populations in the same year. In the low density L. eriocephalus population, individual floral displays were higher than in the dense population, yet reproductive success was low, indicating that plant isolation was a major factor influencing fruit set. This result was due to the extremely low number of beetles per plant and per flower in this population. In the dense L. eriocephalus population, although the displays of individual neighbors were smaller and plants were closer, neighborhood floral display did not contribute significantly to reproductive success, whereas the effect of individual floral display was ambiguous. Species with self‐incompatible rather than self‐compatible breeding systems are expected to experience neighborhood effects on reproductive success; however, at the spatial scale and floral display levels of plants in this study, only individual floral display affected fruit set in H. quadriloculare, whereas neither individual nor neighborhood display influenced fruit set in L. eriocephalus. Therefore, pollinator type, pollinator behavior, and plant and population isolation, rather than breeding system alone, will determine if neighborhood floral display affects fruit set.     

Does the badge of status influence parental care and investment in house sparrows? An experimental test

Theory predicts that traits which signal parental quality might evolve in males of species with biparental care. In avian species, male ornaments may be the most likely candidates for such signals. Male house sparrows (Passer domesticus) possess a black throat patch often referred to as a “badge” or a “badge of status”. By assuming a trade-off between male attractiveness (reflected in male ornaments) and parental care under the differential allocation hypothesis, we predicted that badge size would be negatively correlated with male parental investment. An experiment in which the badge was enlarged in one group and unchanged in a control group was conducted. Our manipulation was predicted to affect female as well as male parental investment. However, we found that eight variables associated with parental investment—the start date for breeding, clutch size, male and female incubation time, male and female food provisioning rate, and average chick weight and the number of fledglings—barely differed between treatments. Also, little evidence for correlations between natural variation in badge size and any of these eight variables was found. Instead, the start date for breeding and the number of fledglings were significantly correlated with both male and female age, while clutch size increased with female age. Female condition was a positive predictor of clutch size and number of fledglings. Female tarsus length, unexpectedly, is related to both male and female incubation time. Badge size was also positively correlated with male age. However, parental age (male or female) was not related to parental care. We conclude that badge size does not signal parental quality, but that the ages of both sexes and the condition of the female play significant roles in the reproductive performance of this species.     

Are plasticity in functional traits and constancy in performance traits linked with invasiveness? An experimental test comparing invasive and naturalized plant species

The role of phenotypic plasticity in plant invasions is among the most often discussed relationships in invasion ecology. However, despite the large number of studies on this topic, there is little consistency. Reconsideration of the role of plasticity by distinguishing two substantially distinct trait-groups, performance traits (contributing directly to fitness) and functional traits (influencing fitness indirectly), could form a more operative framework for comparative studies. In the current study we expect that invasive species benefit from being plastic in functional traits, which allows them to maintain a more constant performance across different environmental conditions compared to non-invasive alien species. We compared invasive and naturalized non-invasive alien plant species by their germination (20 species), their vegetative (10 species) and their reproductive (four species) responses to three different levels of water, light and nutrient availability in a common garden experiment. Used traits were classified into performance (germination ratio, total biomass, seed number) and functional traits (time to germination, root:shoot ratio, specific leaf area, reproductive allocation). We found that invasive and non-invasive species responded similarly to environmental factors, except for example that invasive species germinated earlier with decreasing light conditions or, surprisingly, non-invasive species reacted more intensely to increased nitrogen availability by having a superior ability to achieve greater biomass. The two groups were equally plastic in all the germination and vegetative traits measured but the reproductive traits, since higher plasticity in relative reproductive allocation and higher constancy in reproductive performance showed a pronounced relation with invasiveness.     

Does allometry of a sexually selected ornamental trait vary with sexual selection intensity? A multi‐species test in damselflies

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Does fire affect the temporal pattern of trophic resource supply to pollinators and seed-dispersing frugivores in a Brazilian savanna community?

In tropical savannas, such as the campo cerrado in Brazil, fire plays an important role, affecting plant species’ life history. Since fire has the potential to modify the structure of savanna communities as a whole, it is expected that it may influence the resource supply for mutualists by altering the pattern of investment in sexual reproduction. We used an experimental approach to test if fire alters trophic resource availability to pollinators (nectar, pollen, and oil) and seed-dispersing frugivores (fleshy fruits) by altering the seasonality of reproductive phenophases in a savanna community. We sampled all individuals of 60 species that were common to both control and experimental fire treatments. Each month we recorded the number of reproductive individuals to test whether fire affected the temporal resource offered by the plant assemblage as a whole, and by each specific plant group supporting distinct groups of pollinators and seed-dispersing frugivores. We noticed that fire advanced the nectar, pollen, and fleshy fruit offered by the whole assemblage. Additionally, fire affected the temporal pattern of nectar and pollen available to various pollinator groups, and of fleshy fruits available to all seed-dispersing frugivores. In general, fire seems to have a neutral or even a positive effect on resource availability to mutualists. Nevertheless, there were differences in the availability of the resource utilized by each guild of mutualists.     

Does resource availability affect offspring begging and parental provisioning in a partially begging species?

In species where parents repeatedly provide their offspring with food, the offspring often communicate their need to the parents. Burying beetles, which breed on a wide size range of carcasses of small vertebrates, are interesting model systems to test theories on begging, because the larvae show partial begging, that is, they obtain food through both signalling to their parents (begging) and feeding directly from the carcass. We manipulated resource availability inNicrophorus vespilloides by providing parents with mouse carcasses spanning a wide size range, and allowing them to rear the larvae that hatched, so that both the amount of resources and the number of siblings varied. Time spent begging by each larva was strongly influenced by the time parents spent near the larvae. Brood size had a nonlinear effect on larval begging, with begging increasing with brood size for relatively smaller broods and decreasing again for larger broods. Carcass size and number of parents present had no effect on begging. Time spent provisioning the larvae by the parents was strongly associated with the time spent begging by each larva. Parents spent more time provisioning under biparental care than under uniparental care, while brood size and carcass size had no significant effect. These findings suggest that the larvae adjust their begging to the behaviour of their parents and the number of siblings, but not to the amount of resources. Furthermore, parents adjust the time spent provisioning to the average amount of begging by each larva in the brood, and not to the availability of resources.     

Are reward polymorphisms subject to frequency‐ and density‐dependent selection? Evidence from a monoecious species pollinated by deceit

Abstract Most deceit‐pollinated species involve floral dimorphisms characterized by the presence of rewarding male flowers and nonrewarding female flowers. It has been proposed that this polymorphism establishes the conditions for the action of frequency dependent selection (FDS). The tendency of foraging animals to aggregate in areas of high resource density suggests that pollination efficiency and fruit production may be positively influenced by flower density (density dependent selection, DDS). In this paper we offer a graphical model describing the effects of FDS and DDS on a monoecious species pollinated by deceit. We test the FDS and DDS assumptions and the predictions of the model using field observations and experimental populations of Begonia gracilis in which population sex ratio and flower density were controlled. We found a marked effect of both FDS and DDS on pollinator visitation, fruit‐set, and on the probability of female flowers to setting fruits. We conclude that these two types of selection have had a strong influence on the evolution of deceit‐pollinated species.     

Does saline water consumption affect feeding and fuel deposition rate of a staging,long‐distance migrating passerine?

To accomplish their enduring journeys, migrating birds accumulate fuel consisting mainly of lipids in stopover sites located throughout their migration routes. Fuel deposition rate (FDR) is considered a key parameter determining the speed of migration and thereby bird fitness, and recent studies have demonstrated the positive effects of fresh water consumption on the FDR of migrating blackcaps Sylvia atricapilla. Sewage water reservoirs, characterized by higher water salinity than fresh water, were extensively built in different parts of the world and are used by birds during their travel, but their effects on wildlife and specifically on migrating birds have been largely overlooked thus far. We experimentally examined the effects of water salinity on blackcap FDR during migration. We captured birds in an autumn stopover site, transported them to the laboratory and provided them with fruits, mealworms and water of different salinity levels (0.3, 4.5 and 9‰ NaCl) for several days. We examined the effects of water salinity on the blackcaps’ diet, water consumption and FDR and found that FDR was mainly affected by fruit consumption rate and not by the water salinity levels. Water salinity nevertheless caused elevated water consumption as the birds consumed almost 3 times more saline water than fresh water per consumed fruit mass. Our work is the first to explore the consequences of saline water consumption on migrating passerines, specifically suggesting that anthropogenic alterations of habitats by sewage water treatment facilities may modulate bird nutrition and diet.     

Should food‐deceptive species flower before or after rewarding species? An experimental test of pollinator visitation behaviour under contrasting phenologies

Many plant species reward their pollinators, whereas some species, particularly among orchids, do not. Similarity of floral cues between co‐flowering species influences how rapidly pollinators learn to avoid deceptive plants. This learning process, which affects the reproductive success of deceptive plants, may additionally depend on relative timing of flowering of sympatric rewarding and deceptive species. We tested the combined effects of corolla colour similarity and flowering order of rewarding and deceptive artificial inflorescences on visitation by naïve bumblebees. When deceptive inflorescences were offered after rewarding inflorescences, bumblebees visited them four times more often if both species were similar compared with when they were dissimilar. Pollinator visitation rate to deceptive inflorescences offered before rewarding inflorescences was intermediate and independent of similarity. Thus, early‐flowering deceptive species avoid the costs of dissimilarity with rewarding species. This mechanism may favour adaptive evolution of flowering phenology in deceptive species and explain why temperate deceptive orchids usually flower earlier than rewarding ones.