Evolutionary Ecology

Arctic and Boreal raptor communities will continue to be affected by borealization and other climate change related processes, providing a challenge for ecologists predicting future sates. However, by using community assembly theory and species traits, future communities may be predictable. In this study, we analyzed variation in reproduction traits as a consequence of diet specialization for 29 raptors, 2 skuas and 3 corvids. We assessed and implemented foraging traits for specialists and generalists into predator-prey models from which successful invasion conditions were derived. Specialist raptors produced larger clutch sizes, had a higher proportion of fledged per clutch and also expressed more variation compared to generalist raptors. These results suggest a relationship between diet specialization and reproductive traits which was also observed within phylogenetic orders. Specialist owls (Strigiformes) produced higher clutch sizes with a larger clutch range compared to generalist owls. The same pattern was observed for falcons (Falconiformes). No clear difference in reproduction was observed for specialist and generalist hawks, kites and eagles (Accipitriformes). Corvids expressed clutch sizes similar to that of specialist raptors while having the lowest proportion of fledged per clutch. Differences in foraging traits between specialists and generalists could be distinguished using functional response curves. A predator-prey model parameterized with foraging trait data showed that a generalist can coexist with a resident specialist if it has access to prey unavailable to the resident specialist. Otherwise, the native specialist outcompetes the invading generalist due to foraging efficiency. The combined empirical and theoretical findings in this study show how diet specialization affects both reproduction and the potential invasion success of raptors.

Many animals have camouflage appearances that correspond to the habitat where they live, termed phenotype-environment matching. However, this has generally been tested in only a limited number of systems and natural settings, and often not to the relevant vision of key receivers (predators) across different spatial scales. Here, we measured the plumage attributes and putative camouflage used by ground-nesting bird species specialist to particular biomes, specifically tropical rainforest, taiga forest, dry forest, grassland, desert, and tundra from museum specimens. Digital photography and image analysis were used to quantify colour patterns to models of predator vision to understand how different colour patterns may correspond with biome type. With this information, we next created bird models that were photographed in situ in the Valdivian temperate rainforest and Patagonian grassland biomes of Chile to quantify the extent to which plumage coloration and pattern traits provide effective camouflage at different spatial scales. In general, we find that specialist ground-nesting birds express a phenotype that better matches the substrate composition and vegetation structure across large spatial scales of their own biome. This study reveals how animal camouflage works across biomes, relative to the visual system of raptor predators, and at the appropriate distance at which detection may occur.

Animal colouration evolves under multiple, often conflicting, selective pressures. Conspicuous, non-aposematic colour-patterns that enhance conspecific communication may simultaneously increase detectability by predators. Such trade-offs can be resolved by optimising colour-patterns to match the perceptual abilities of different receivers. We tested whether dorsal colour-patterns of the Lusitanian wall lizard (Podarcis lusitanicus) are optimised for ecologically relevant receivers across relevant viewing distances, while accounting for the visual acuity of conspecifics and predators. Conspecifics and snakes detected chromatic information at shorter distances, whereas achromatic and luminance information were detected at longer distances. Birds showed a uniform decline in detectability across the colour-pattern components with increasing viewing distance. Larger males retained high chromatic detectability across all receivers despite the general distance-related decline, whereas females and smaller individuals exhibited less salient colour patterns, consistent with predator avoidance strategy. Our results show that lizards resolve the trade-off between conspecific communication and predator detection through distance-dependent colour-pattern perceptibility across receivers. This resolution breaks down in large males, for whom the benefits of salient chromatic patterns for intraspecific communication may outweigh increased detectability to predators.

Data on the shape of a group of organisms can be conceptualized as forming a point cloud in the multivariate space of measurement. This is literally true for traditional linear measures, while in a geometric morphometric context the cloud resides in Kendalls shape space, tangent to the true shape space. Regardless of the method of construction, the topology of this point cloud, or phenotypic (hyper)ellipse, is a beguiling target for evolutionary analysis. Reordination of the axes will not change the geometry of this phenotypic ellipse, and the notion that its geometry carries a meaningful biological signal is an old idea; but the character of this signal is often elusive. This paper explores the application of the most commonly used parameter designed to summarize differences in phenotypic ellipse geometry (relative eigenvalue variance, or Vrel), and demonstrates that it is incapable of differentiating between several plausible ways in which phenotypic ellipse geometry might differ among species, because it confounds three separate parameters necessary to describe the ellipse. Two example data sets are analyzed to illustrate variability in phenotypic ellipse geometry and draw conclusions about observed differences. The first case compares wolves to domestic dogs, and replicates previous findings of much greater variance yet tighter integration in dogs. This calls into question the simple model of a single peak in the fitness landscape of dogs. The second example comprises geometric morphometric landmarks from the jaws of a clade of sigmodontine rodents, and allows comparison of ellipse geometry in a phylogenetically controlled setting with qualitative ecological categories. Three parameters are found to vary in concert along a grade of most to least ecologically specialized: the phenotypic variance, the effective rank (dimensionality), and the degree of covariance (Vrel and related metrics). Use of all three of these quantities to characterize the geometry of the phenotypic ellipse is advocated, as all are necessary to characterize how variance is distributed in the ellipse in different taxa. The phenotypic ellipse geometries illustrated here appear to reflect something of the geometry of the adaptive peak upon which each taxon sits; or that they represent (aspects of) the mapping function of adaptive peak to phenotypic ellipse, in ways first predicted by Simpson in the twentieth century.

Humans are efficient and deadly predators, yet they may also interact with wildlife in non-lethal ways. This study explores how interactions with lethal and non-lethal human "superpredators" alter predator-prey dynamics using an agent-based modelling approach. Our model incorporates both the consumptive (lethal) and non-consumptive (behavioural) effects of humans, as well as of predators on prey. We explored how the replacement of apex predators by humans affects mesopredator-prey dynamics, with particular emphasis on trophic targeting and differences between lethal and non-lethal interactions. We found that human superpredators have a greater effect on model outcomes than apex predators. When superpredators consume mesopredators alone or with prey, the probability of mesopredator-prey coexistence increases to a greater extent than when apex predators consume mesopredators. In contrast, superpredators consuming only prey slightly increases overall extinction risks and reduces coexistence. Non-lethal superpredators, despite eliciting anti-predator responses in mesopredators and prey, had a negligible effect on population dynamics. Our findings demonstrate that human superpredators may functionally replace apex predators when they are lethal. However, non-lethal interactions with humans may not be as ecologically significant as lethal interactions, even when humans induce anti-predator responses.

Empirical studies have demonstrated that mutation rates may change with individual condition, such as in the case of stress-induced mutagenesis. This has led to the hypothesis that condition-dependent (or "plastic") mutation rates could be selectively favoured, as the increased production of new mutants in times of maladaptation enhances evolvability, the ability to undergo adaptive evolution. However, while empirical evidence for condition-dependent mutation rates is accumulating, theoretical models studying their evolution are lacking. Here, we employ an individual-based simulation approach to examine the evolution of condition-dependent mutation rates in a changing environment. We find that condition-dependent mutation rates consistently evolve when the environment changes at an intermediate pace. Furthermore, populations with condition-dependent mutation rates are substantially better adapted to their (changing) environment. Finally, the evolutionary dynamics of condition-dependent mutation rates are both accelerated and destabilised when the mutation rate is self-referential (i.e., when mutator loci affect their own mutation rate). We conclude that condition-dependent mutation rates (and thus evolvability) can readily evolve in changing environments. Significance statementMutation provides the raw material for evolution. Mutation rates thus tune evolvability, the ability to undergo adaptive evolution: if mutation rates are too low, evolution is impeded; if mutation rates are too high, adaptive traits cannot be maintained. Using a theoretical model, we explore the evolution of plastic mutation rates that systematically depend on the condition of the organism and its environment. An example is stress-induced mutagenesis in bacteria, which is implicated in the evolution of antibiotic resistance. We show that plastic mutation rates readily evolve, providing "well-timed" variation specifically when organisms are poorly adapted. Such plastic mutation rates thus facilitate better adaptation to changing environments, and their evolution provides an example of evolvability itself evolving.

The evolutionary patterns of trait diversification provide insights into the function of the trait. Early burst of trait evolution is often associated with adaptive radiation, rapidly diversifying the trait in response to vacant niches followed by the slowdown of the diversification with niche filling, whereas late burst is more likely to be associated with sexual selection, possibly contributing to reproductive barriers between closely related species. Here, we studied the diversification of tail fork depth through time in hirundines to infer its function, which remains unclear due to the competing two alternative hypotheses: the sexual selection hypothesis, which is a classic explanation of deeply forked tails, proposed that this trait has evolved via sexual selection, which was then challenged by the viability selection hypothesis, which proposed that deeply forked tails have mainly evolved via viability selection for enhancing aerodynamic performance during aerial foraging on large prey. We found a late burst of tail fork depth, but not of bill length, i.e., an index of prey size. The observed pattern is consistent with the sexual selection hypothesis but not with the viability selection hypothesis.

Animal personality is the expression of consistent among-individual variation in a behavioural trait across time and context. The application of this theory to behavioural research provides a valuable framework to investigate the among- and within-individual variation in behaviour. The animal personality theory is particularly relevant to understand and diagnostic fish welfare. It can be integrated within the nature-based welfare framework, which emphasizes the expression of species-specific behaviours and the maintenance of consistent behavioural patterns over time. Because behavioural trait such as boldness may be closely linked to other functional phenotypes, such associations reflect the broader concept of animal personality, whereby a behavioural trait can covary with another phenotypic trait to support an adaptive responses to environmental conditions. Although these relationships are both species and context dependent, they are consistently shaped by environmental conditions and environmental complexity tend to promote species specific behaviours and reduce maladaptive traits. Here we examined how structural environmental complexity shapes personality in Arctic charr and their covarying functional phenotypes, specifically growth rate and brain size. We propose that environmental complexity promotes alternative behavioural and functional phenotypes through multivariate phenotypic plasticity. We found that environmental complexity did not influence mean boldness between both treatments, but repeatability of boldness in the complex environment was remarkably consistent over a longer-term period, while estimates collapse after seven days in the plain treatment. Our findings are a clear evidence that environmental complexity foster stable behavioural trait expression and that a plain environment may suppress personality. Our results provide compelling evidence that behavioural structure and dynamics are embedded within patterns of behavioural variance. Although we found no support for behavioural covariation or associations with growth rate and brain size, we suggest that the animal personality framework may offer a valuable approach for diagnosing fish welfare issues through the partitioning of behavioural variance.

Land-bridge islands are former mainland areas isolated by post-glacial sea-level rise (<15,000 years) and the most common island type. Because of their recurrent connectivity with continents, it is unclear whether species on land-bridge islands can undergo evolutionary changes associated with the more isolated oceanic islands ( island syndrome). Here, we test the hypothesis that the selective environment on land-bridge islands exerts predictable and consistent evolutionary shifts in morphological traits of songbirds. We apply Bayesian hierarchical models to a morphological dataset of 6,917 individuals comprising 185 species of songbirds from four land-bridge islands (Bioko, Sri Lanka, Taiwan and Trinidad) and adjacent continents. Across all 185 species, we find that occurrence on a land-bridge island has clear directional effects on five morphological traits related to beak, wing, and tarsus, as well as a general increase in body size. At the species level, 57 out of 90 tested species exhibit significant morphological divergence between land-bridge island and mainland, yet for only 20 of these are the land-bridge island populations recognised as distinct endemic subspecies. Our results show that occurrence on land-bridge islands has a detectable effect on passerine morphology consistent with the island syndrome, and suggest these islands harbour previously unrecognized unique biodiversity.

The impacts of a changing abiotic environment on fitness and performance arise not only from low tolerance to new environmental conditions, but also from changes in the abundance and interaction intensity with other species. The strength of the interaction may itself depend on how well each species performs across environments, but there is a dearth of studies investigating how intrinsic fitness and interaction intensity covary across an abiotic environmental gradient. We addressed this question in a hypersaline consumer-resource system: the microalga Dunaliella spp. grazed by the brine shrimp Artemia franciscana. We exposed four Dunaliella strains to a range of salinities above seawater, with or without brine shrimps, and tracked their population sizes over time and the survival of their predators, to estimate basic parameters of a Lotka-Volterra model. We found that the intrinsic growth rate of algae, the survival rate of predators, and the per-capita predation rate, all varied with salinity and algal strain. Significant interactions between strain and salinity further revealed that these ecological responses to salinity are evolvable. Together with correlations between demographic parameters across salinity, this suggests that predation may influence the evolution of salinity tolerance curves, blurring the line between the fundamental and realized niches.

An animal's musculature must support its specific biomechanical needs, so muscle morphology and volume allocation may adapt when locomotor strategies diversify. We examined muscle size and morphology in two sister families of bombycoid moths, wild silkmoths (Saturniidae) and hawkmoths (Sphingidae), that have diverged in wingbeat frequency, wing morphology, and behavior. Although both families rely on the same muscles to power and steer flight, they may distribute muscle volume differently to prioritize distinct functions. We hypothesized that flight power muscle proportions are larger in hawkmoths and increase with wingbeat frequency, helping meet inertial power demands of high-frequency maneuverable flight. We also hypothesized that some individual muscles diverge in proportional volume and area to support distinct wing control strategies. To test our hypotheses, we took CT scans of twenty bombycoid species and quantified volumes and geometries of six flight muscle pairs. As expected, flight power muscle proportions positively correlate with wingbeat frequency and are generally greater in hawkmoths. Two of three steering muscles diverge substantially in relative volume and area between families. Most muscles exhibit greater length in silkmoths and greater cross-sectional area in hawkmoths. Finally, the dorsal oblique(DO) muscle diverges exceptionally in size and morphology, being highly developed in hawkmoths and smaller or absent in silkmoths. This unexpected difference supports the DO having an underappreciated role in flight control, possibly via shaping indirect strain propagation in the elastic thorax. We show that muscle volume distribution parallels bombycoids' divergent flight strategies, demonstrating how muscle allocation can adapt for specialized functional goals.

Brain size and structure are hypothesized to be adaptively designed to satisfy the behavioral requirements of securing food and living socially. The importance of these socioecological and sociobiological selective forces in brain evolution is constantly debated. Socioecological divergence is striking in the Neotropical ant genus Neoponera: N. apicalis is a generalist solitary predator forming small colonies of ~100 whereas N. commutata colonies are approximately 10 times as large and workers pheromonally organize cooperatively raids only on Syntermes termite colonies. We interspecifically compared the size and structure of the compound eyes, size and number of antennal glomeruli, mosaic brain scaling and synaptic processing (microglomeruli-MG). Our results indicate that N. apicalis workers have a larger number of ommatidia, antennal lobe glomeruli, and allometrically larger antennal and optic lobes than N. commutata. These sensory traits were associated with differences in higher-order processing architectures in the mushroom body (MB) microglomeruli (MG). N. commutata workers had an allometrically larger MB, perhaps due to their socially complex chemical foraging communication, although MG density in N. apicalis was higher in both the MB lip and collar, regions associated with processing olfactory and visual information, respectively. The increase in MG density in N. apicalis may be associated with higher demands for navigation, learning, and memory, as well as a higher density of antennal lobe glomeruli to support prey odor discrimination. In contrast, N. commutata workers had larger ommatidia and antennal lobe glomeruli. Larger ommatidia correlate with their diurnal/nocturnal habits and a larger MB Our findings indicate that differences in behavioral performance demands associated with socioecological differentiation are reflected in variation in visual and olfactory system structure, brain size, mosaicism, and synaptic organization. Our results support both social and ecological brain hypothesis as drivers of mosaic brain evolution.

Phenotypic differences among populations can arise through heritable genetic divergence, phenotypic plasticity, or both, making it difficult to determine whether trait-environment correlations observed in nature reflect adaptive evolution. Within threespine stickleback (Gasterosteus aculeatus) studies, numerous document morphological differences among allopatric-, parapatric-, and even sympatric populations. These phenotypic differences among populations are often correlated with diet and lake habitat (e.g., lake size), suggesting an adaptive value to the population differences. However, many studies of ecomorphological divergence in stickleback use wild-caught stickleback, which may differ due to evolution or plasticity. Although common garden experiments have confirmed that population differences can be heritable, such experiments typically entail small numbers of populations. Consequently, we still do not know to what extent well-known trait-environment correlations in stickleback are a result of evolution. To address this gap, we reared stickleback embryos from 27 lake populations on Vancouver Island, in a laboratory environment. Morphological differences among populations persist in common-garden fish, confirming a large role for divergent evolution. These heritable differences were associated with environmental variation among lakes, implying an adaptive value. However, some well-known trait-environment relationships in stickleback did not persist in common-garden fish and may be primarily plastic.

Recognizing patterns in functional trait evolution is a necessary step in testing macroevolutionary questions. Quantification of these patterns and interpretation of their generative processes relies on an ever-expanding suite of comparative approaches, but current methods oversimplify the process-to-pattern mapping. This simplification may promote binary classifications of patterns and their drivers, such as adaptive versus non-adaptive or constrained versus unconstrained. A potentially more robust and evolutionarily informative alternative is to fit an expanded suite of evolutionary models at different levels of taxonomic or ecological resolution to reveal how evolutionary drivers imprint simultaneously on observed patterns of trait evolution. Here, we perform a clade-wide analysis of functional trait evolution across Anura (frogs and toads). Focusing on three functionally important traits, we quantify the relative fit of random, directional, and bounded models at the intra- and inter-microhabitat levels to explore how changes in trait function across microhabitats translate into different evolutionary regimes. We recover heterogeneous regime-level distributions of model support that imply complex underlying evolutionary dynamics, while also revealing methodological biases and model identifiability issues. These findings underscore the need to develop more robust tools for evolutionary model fitting and advance beyond binary frameworks for interpreting evolutionary processes.

True crabs (Brachyura) are among the most iconic marine arthropods, representing noteworthy examples of morphological and ecological disparity. A striking feature of brachyurans are their anterior pincer-like appendages: chelipeds. These structures showcase a large diversity of morphologies that reflect ecology and overall multifunctionality. Yet, a comprehensive assessment of appendage functional morphology within phylogenetic and ecological trait contexts has never been attempted. By combining 3D geometric morphometrics, finite element analyses, multilocus molecular phylogeny, and ecological trait data for 80 crab species, including three fossil forms, we unveil a complex evolutionary history for crab chelipeds. Despite extreme shape diversity amongst chelipeds, stress distributions are very similar across taxa and hint a many-to-one pattern. High concentrations of chelipeds within constrained morphospace regions associated with peak pinch forces illustrates that brachyuran morphologies optimised for shell crushing may have arisen in the Cretaceous. Deviations from this morphospace highlight the diversification of non-shell-crushing life modes and the influence of sexual selection on appendages. Neither cheliped shape nor pinch force show phylogenetic signal. Together these results indicate that the evolution of cheliped shape is closely associated with, and inferred to have been strongly influenced by, crab ecology, biomechanical needs and sexual selection. SIGNIFICANCE STATEMENTChelipeds, the pincer-like claws of crabs, are among the most morphologically diverse appendages within Arthropoda, yet the evolutionary forces driving this diversity remain poorly understood. By integrating 3D geometric morphometrics, biomechanical modelling, molecular phylogeny, and ecological data across 80 crab species including fossil forms, we demonstrate that cheliped morphology is driven by ecology, biomechanical demands, and sexual selection rather than phylogenetic relatedness. The multifunctionality of these structures produces strong evidence for many-to-one mapping of form to function. Morphologies optimised for durophagy appear to have originated in the Cretaceous, with subsequent diversification into manipulative and sexually selected forms from a morphologically flexible foundation. These findings demonstrate that cheliped diversity reflects a complex interplay between ecological specialisation, biomechanical optimisation, and sexual selection across Brachyura.

Across species, changing environmental conditions are altering major phenological milestones. In seasonal environments, this can involve shifts in the timing of activity onset, growth, reproduction, and senescence; however, predicting these responses can be challenging and often requires species-specific information. In this study, we investigated phenological patterns of bumble bee colony development using observations from wild nests of two common species, Bombus griseocollis and B. impatiens. Using observations of nest traffic, we documented dates of nest-searching, peak worker activity, first gyne (new queen) production, and colony senescence over three years of field data collection. We also tested whether colony growth was density dependent, because longstanding life history models have shown that colony growth patterns determine the optimal timing of reproduction in constant environments. Here, we present a novel extension of these models, using them to infer that density-independent colony growth should be associated with extended phenology in warmer years, while density-dependent should have consistent phenology. In total, we found 79 wild nests, including 34 reproductive colonies (19 B. griseocolis and 15 B. impatiens). Bombus impatiens colonies were larger, had a longer activity period, and showed density-independent growth and later reproduction in warmer years, while B. griseocollis colonies were smaller, shorter-lived, and showed density-dependent growth and similar dates of reproduction across years. Both species had higher gyne production in warmer years. These results highlight the role of life history theory for predicting interspecific variation in phenological changes, as well as a research framework for exploring possible evolutionary mismatches of existing life histories in changing environments.

Resource limitation is a central ecological phenomenon shaping pollinator behavior, reproduction, and community dynamics. Many studies have looked at these interactions, but few have forced competition and explored species-specific responses. Here, we experimentally evaluated behavioral and reproductive responses of honey bees (Apis mellifera), bumble bees (Bombus impatiens), and mason bees (Osmia bruneri) in controlled single and multispecies foraging environments. Across all treatments, each species exhibited distinct forms of behavioral compensation when exposed to increased interspecific competition. Apis mellifera reduced the number of foraging events in mixed species cages yet showed variation in colony growth. B. impatiens increased foraging effort when co-foraging with other species, but this heightened activity did not translate into increased colony growth. O. bruneri maintained consistent foraging effort across treatments but exhibited reduced reproduction and a marked shift in floral host use, switching from preferred species (Phacelia tanacetifolia and Melilotus alba) to less-preferred alternatives (Collinsia grandiflora and Trifolium incarnatum) when competing with social bees. Regardless of compensatory behaviors, all three species demonstrated reduced reproductive success under competitive conditions. Our findings underscore the importance of evaluating interspecific competition when placing managed bees in natural or semi-natural habitats to avoid inadvertently stressing wild or managed bee populations and taking into consideration species-specific responses in addition to community level responses to competition.

For organisms with biparental care, successful reproduction hinges on coordination between partners. Seabirds face an extreme coordination challenge because parents must schedule nest attendance on land with long-distance foraging trips at sea. We present a computational model of incubation schedules for a vulnerable seabird, the Leachs Storm-Petrel (Hydrobates leucorhous). Using only simple energetic rules and parameters, the model recapitulates natural incubation rhythms, exposes a tradeoff between parent energy and egg attendance, and predicts severe reproductive failure in harsh environments. Incubation primarily fails through "schedule breakdown" -- a single point in the season when both parents spend too long foraging and the egg dies from cold. The resilience of the developing offspring to neglect is thus a fundamental adaptation to the uncertainties of biparental care. These results raise new alarms about the indirect causes of reproductive failure in sensitive marine species and provide theoretical foundations for the evolutionary ecology of scheduling behaviors.

Premise of researchIn plant species that depend on pollinators to produce seeds, declines in pollinator populations should strengthen natural selection for floral traits that increase outcross pollen receipt by increasing the probability of pollinator visitation. One such trait is floral longevity, the amount of time that a flower is open and functional. If flowers that are open longer have a higher probability of pollinator visitation and this pollination benefit outweighs the resource cost of maintaining floral tissue, then pollinator decline should strengthen selection for extended floral longevity. MethodologyTo test whether pollinator decline could strengthen selection for extended longevity, we studied the pollinator-dependent species Lobelia siphilitica. We exposed L. siphilitica plants to either an ambient open-pollination treatment or a reduced open-pollination treatment that simulates pollinator decline. Within each pollination treatment, we measured floral longevity and seeds per fruit and used these data to estimate directional selection on longevity. This experiment was repeated twice. Pivotal resultsIn one experiment, there was significant selection within both ambient and reduced pollination treatments for shortened (rather than extended) floral longevity. In the other experiment, selection on floral longevity within both ambient and reduced pollination treatments was not significantly different from zero. ConclusionsWe found no evidence of selection for extended floral longevity, regardless of the pollination environment. These results suggest that pollinator decline will not strengthen selection for extended longevity and thus that pollinator-dependent species such as L. siphilitica may not respond to pollinator decline by evolving extended floral longevity.

Yeasts thriving in floral nectar have a proven ecological engineering capacity, as they modify the internal environment and the rewards of individual flowers in ways that influence how plants and pollinators interact. This floricentric perpective, however, produces only a partial view of the potential role of yeasts in plant-pollinator interactions, and little is known on their possible engineering effects on whole inflorescences. This study assessed yeast ability to shape features of the vertical inflorescences of Gladiolus illyricus (Iridaceae), a Darwins syndrome plant (acropetalous protandrous flowers pollinated by insects foraging from bottom to top). When yeasts were excluded from inflorescences a declining gradient in nectar sugar concentration built up from the oldest, female-stage flowers at bottom up to the youngest, male-stage ones at top. This plant-intrinsic gradient was reversed in inflorescences with yeasts, where sugar concentration increased from the female-stage flowers harboring dense yeasts population to male-stage ones with few yeasts. The two species of yeasts involved (Metschnikowi reukaufii and M. gruessii) differed in their effects. Results add support to the growing consensus that specialized nectar yeasts can obfuscate intrinsic patterns of intra and interspecific variation in flowering features of plants at the level of individual flower, inflorescence, population and plant community.