Functional Ecology

The mechanisms driving host-symbiont associations across space and time in contemporary mutualisms can give insight into the capacity for symbiotic organisms to respond to environmental change. High specificity between partners can increase cooperation and facilitate efficient holobiont selection, whereas low specificity could reduce host benefit, but facilitate adaptive associations across heterogeneous environments. The present study explores specificity in natural populations of a cnidarian-algal model, Exaiptasia diaphana, across a latitudinal gradient to understand the genetic and environmental effects driving host-symbiont associations, and their relation to heritable and/or environmental symbiont acquisition. We found that symbiotic associations were extremely flexible in E. diaphana, regardless of transmission mode. E. diaphana were capable of associating with diverse symbiont communities across genetically identical hosts seeded with vertically transmitted symbionts, as well as across highly connected host populations which acquire symbionts horizontally. Host population connectivity was complex and unrelated to geographic distance, whereas symbiont community composition tracked the thermal gradient, potentially due to context dependent biotic interactions. These results indicate that in a flexible symbiosis, symbiont communities are environmentally-determined, suggesting the future of this symbiosis will likely depend on climate adaptation of symbionts.

Stingless bees are key pollinators in tropical ecosystems, yet their ecological dynamics remain poorly understood in highly seasonal environments such as the seasonally dry tropical forests of Ecuador. These ecosystems experience pronounced climatic seasonality, with sharp transitions between dry and wet periods that strongly affect floral resource availability. Understanding interspecific competition and niche partitioning in such systems is critical, particularly given the global decline of pollinators. We investigated resource use and niche dynamics in two native stingless bees, Melipona mimetica and Scaptotrigona sp., by quantifying pollen, nectar, and resin collection across seasons. Log-linear models were used to test the effects of species, season, and their interaction on resource use, while non-metric multidimensional scaling (NMDS) assessed niche overlap. Contrary to the expectation that niche overlap increases under resource scarcity, we found greater overlap during the wet season, when resources are more abundant. This suggests that both species converge on high-quality floral resources during peak availability, reflecting an adaptive response to strong environmental seasonality. Pollen use remained stable across seasons, consistent with generalist foraging behavior. In contrast, nectar collection increased significantly during the wet season, while resin exhibited a shared seasonal peak, likely associated with synchronized nest construction or maintenance. These findings reveal context-dependent competition dynamics and highlight the role of environmental seasonality in shaping pollinator interactions. Our study provides new insights into the ecology of threatened stingless bees and contributes to their conservation in tropical dry forest ecosystems.

Winter survival in dormant animals depends on conserving finite energy reserves, yet winter temperatures fluctuate around shifting means. In ectotherms, metabolic rate increases exponentially with temperature, so thermal variability is expected to accelerate energy loss, with important consequences for overwinter survival and population persistence under climate change. However, it remains unclear whether dormant ectotherms can compensate physiologically for thermal variability. We overwintered Bombus impatiens queens under constant (2, 3, 4{degrees}C) or variable (2 {+/-} 6{degrees}C or 4 {+/-} 6{degrees}C) regimes for six weeks, then measured metabolic rates across a range of temperatures. The temperature dependence of metabolic rate shifted in response to thermal experience, but the direction of compensation depended on mean temperature: variability centered on 2{degrees}C elevated metabolic rate and increased thermal sensitivity relative to all other conditions, whereas variability centered on 4{degrees}C reduced metabolic rate and dampened thermal sensitivity relative to constant 4{degrees}C. We used these metabolic responses to simulate rates of lipid depletion and found that survival trajectories echoed physiological shifts: experiencing variability around 2{degrees}C would reduce subsequent survival time, whereas experiencing variability around 4{degrees}C would preserve subsequent survival even under variable future conditions. Thus, identical thermal variance produced opposite energetic outcomes depending on the mean temperature around which fluctuations occurred. Integrating both temperature means and variability is, therefore, essential for predicting overwintering survival in a changing world.

O_LIMany organisms alter phenotypically plastic traits in response to environmental cues to match their phenotypes with variable environments. In larval amphibians, development and growth rates respond to spatiotemporally variable mortality risk from predation, wetland drying, or resource limitation. However, these rates are also temperature-dependent for ectotherms. Although wild animals experience these factors simultaneously (e.g., thermal regimes, predation risk, resource limitation), most studies investigate their impacts in isolation, limiting our understanding of how they interact across ecological contexts. C_LIO_LIHere we simultaneously exposed larval Plains Leopard Frogs (Lithobates blairi) to varying resource levels and predation risk treatments across a thermal regime to investigate the joint effects of these ecological drivers on growth and development rates and their consequences for size and vagility after metamorphosis. We crossed two predation treatments (waterborne cues from Procambarus gracilis fed L. blairi larvae, control water) with three food resource levels (5%, 25%, 50% of body mass) and six thermal regimes (diel {+/-} 3{degrees}C cycles of 15, 20, 22, 24, 26, 28{degrees}C), replicating each combination five times for a total of 180 individuals. We recorded growth and development rates and completion of metamorphosis, then measured juvenile body size and jumping performance. C_LIO_LIThe number of larvae completing metamorphosis was primarily determined by temperature and temperature-dependent effects of resource limitation. Percent metamorphosis peaked at intermediate temperatures when resources were high and were higher in predation-risk treatments at the warmest temperatures. Under high resources, development and growth rates showed unimodal thermal responses that were absent when resources were constrained. Higher resources increased development rates, but proportional increases in growth maintained constant body size across temperatures. Post-metamorphic body size differed only by predation treatment, with predator-exposed individuals being smaller. Juvenile jumping performance increased with body size and individuals raised with high resources without predator cues exhibited the highest performance. C_LIO_LIThe absence of temperature effects on size at metamorphosis reflected unexpected coupling of growth and development rates across treatments, producing uniform body sizes. This pattern contrasts with the temperature-size rule and suggests that plastic responses may exhibit selection for a minimum viable size at metamorphosis. C_LI

Most plants require animal pollination to reproduce, prompting concern that pollinator declines immediately threaten plant populations. This concern is warranted if pollinator-mediated seed losses cause declines in plant population growth rates ({lambda}). However, demographic trade-offs might reduce the risk of population decline if seed loss improves performance elsewhere in the life cycle. We conducted a multi-year pollination manipulation on four species and measured how demographic vital rates and {lambda} responded. Seed responses did not predict net changes in {lambda}. Reduced pollination decreased seed production, but only caused a net decrease in {lambda} in one species; in the others, improved survival buffered {lambda}. Increased pollination boosted seed production, but at a cost to survival that caused a net reduction in {lambda} in three species. Our results highlight the importance of demographic trade-offs for understanding the impacts of pollinator declines on plant biodiversity and, more broadly, the population-level impacts of changing mutualisms.

Species in stable social groups engage in diverse social interactions, where social competence - the ability to adjust behaviour using social information - can influence fitness. Yet, whether adaptive behavioural flexibility is expressed across contexts within individuals remains relatively untested. To address this, we exposed cooperatively breeding cichlids (Neolamprologus pulcher) to a role-reversal paradigm. In this species, the early social environment shapes social competence, with more competent individuals adjusting behaviour flexibly to social challenges, while individuals also show consistent differences in traits such as aggression. In the present study, individuals were successively assigned to two contrasting roles, smaller territory owners (TOs) and larger intruders (INTs). We predicted role-specific social competence metrics based on behaviours facilitating shelter acquisition. Social competence metrics correlated within, but not across the two roles. Competent TOs showed shorter latencies to submit, more submissive responses to received aggression, and low aggression after initial submission. Competent INTs escalated quickly and relied more on overt aggression rather than displays, allowing faster shelter acquisition. Across roles, individuals competent as TOs were not competent as INTs. In contrast, consistent individual differences in aggression across social roles suggest that stable behavioural tendencies ( animal personalities) may constrain how social competence shapes behavioural strategies.

Factors varying along elevational gradients impose strong aerodynamic and physiological constraints on powered flight, yet the internal anatomical correlates of flight performance in animals under such conditions remain poorly understood. In hummingbirds, sustained hovering requires extreme muscular power output, making the pectoralis muscle a key interface between environmental constraint and performance. We tested whether elevation is associated with variation in pectoralis microanatomy across three hummingbird assemblages spanning a [~]1500 m gradient in the Colombian Andes. Using tissue morphometry of trichrome-stained transverse sections of the pectoralis, we measured interstitial collagen fraction as a proxy for extracellular matrix investment and quantified fiber cross-sectional area, packing density, and size heterogeneity. Collagen investment varied across elevational bands, peaking at mid elevation ([~]1750 m) and declining toward high elevation ([~]2600 m). In contrast, muscle fibers were smaller and more densely packed at higher elevations. Variation among species was small relative to differences among elevational assemblages. Formal model comparisons provided limited support for non-linear responses to elevation, indicating that patterns across traits are better explained by interacting constraints than by a single monotonic response to factors varying along elevational gradients. These results show that hummingbird flight muscle microanatomy varies with elevation in a trait-specific manner, with the strongest evidence in fiber geometry. More broadly, our findings highlight that multiple components of muscle microarchitecture, including the extracellular matrix, vary in a context-dependent manner across elevational gradients in an extreme volant system.

O_LISaproxylic community assembly is structured both by deadwood and forest habitat gradients, as well as biotic interactions such as competition, predation, and parasitism. However, covariation between abiotic and biotic conditions in natural systems have limited our ability to disentangle these mechanisms. Furthermore, focus towards beetles and fungi in temperate or boreal forests has led to important taxonomic and geographic knowledge gaps. C_LIO_LIHere, we tested how experimentally-manipulated tree diversity, deadwood position (lying vs. standing), and biotic interactions with a dominant antagonist (ant exclusion) structure the community assembly of deadwood-cavity-nesting bees, wasps, and their parasitoids in a subtropical forest. C_LIO_LIOur findings reveal that lying deadwood supports a nested subset of the communities occurring in standing deadwood, with less diversity and abundance of hosts and parasitoids. We found that increased moisture, rather than ant activity, was the primary mechanism filtering Hymenoptera communities, as deadwood in contact with the forest floor retained twice as much moisture as standing substrate. Moreover, moisture gradients within each substrate type further reduced host abundance - likely due to reduced brood cell production and survival. In contrast, forest habitat (tree species richness, canopy cover, and coarse woody debris) had comparatively minor roles in shaping cavity-nesting community assembly. C_LIO_LIOur results provide a mechanism for the positive association between cavity-nesting Hymenoptera and standing deadwood in forests. Because standing deadwood is typically scarce in many managed forests, these findings support the retention and enhancement of such substrates to promote these ecologically-important insects. C_LI

Interactions between cleaner fish Labroides dimidiatus and client fish, from which cleaners remove ectoparasites and mucus, represent a textbook example of mutualism involving sophisticated strategic decision-making. However, cleaners must also face intraspecific social challenges within a size-based hierarchy, where the largest females may eventually change sex and become males with higher reproductive rates. Following 540 individuals over 11 months, we found that, contrary to expectations, slow-growing females spent more time cleaning and cheated more frequently, without causing more negative client responses than fast-growing females did. Instead, variation in growth was best explained by social factors: fast-growing individuals experienced reduced social control, while slow growers spent more time in proximity to dominant individuals. As there was no evidence that spawning activity affected growth patterns, it appears that fast growth as a viable strategy for becoming a male largely depends on the lack of control by dominants.

Electric organ discharge (EOD) waveform diversity in African elephantfish is often attributed to sexual selection, yet EODs also mediate active electrolocation during prey detection, raising the possibility that natural selection on foraging ecology contributes to waveform divergence. Paramormyrops kingsleyae exhibits an intraspecific polymorphism where certain populations emit biphasic EODs whereas other populations emit triphasic waveforms. The genes underlying this polymorphism show signatures of selection; the polymorphism persists despite gene flow and is behaviorally discriminable by the fish themselves. If waveform differences influence prey detection during active electrolocation, biphasic and triphasic fish should consume systematically different prey. We tested this prediction using DNA metabarcoding of gut contents from 186 mormyrids representing 16 species across eight sites in Gabon, employing two independent COI primer sets for cross-validation and pairing dietary data with environmental invertebrate sampling to distinguish active prey preference from passive availability. At the community level in the diverse Bale Creek mormyrid assemblage, species identity was the dominant predictor of diet composition (R{superscript 2} {approx} 24%), consistent with phylogenetic signal in foraging ecology. Within P. kingsleyae, waveform type was the strongest independent predictor of dietary composition (R{superscript 2} = 5-6%), explaining variance independently of geographic region, sex, body size, and parasitism status -- a result concordant across both primer sets. Dietary differences were driven by prey species turnover rather than differential abundance of shared prey, and prey selectivity analyses confirmed that waveform types differ in which prey they actively prefer, not merely in what is locally available. These findings are consistent with natural selection on foraging ecology contributing to the maintenance of EOD waveform polymorphism, though the sensory mechanisms linking subtle waveform differences to prey detection remain an open question.

Resource competition and parasite exposure both present common density-dependent fitness costs for wild animals. Because launching effective immune responses is costly in terms of resources, parasites fitness costs should be further exacerbated in high-density, resource-depleted areas. To disentangle these relationships, we related density, parasitism, and resource availability to survival and fecundity across lifespan in a long-term study of wild red deer. All fitness measures declined with a combination of parasite count, greater density, and reduced resource availability. Beyond these relationships, as expected, local density and resource scarcity exacerbated survival costs of parasitism in calves, effectively undermining tolerance of infection. However, these synergistic relationships faded in yearlings and then reversed in adults, likely through age-structured selection biases. These findings emphasize that the costs of parasites and resource scarcity can be synergistic and intertwined with density in wild populations, accentuating the value of incorporating resource competition when examining parasite-dependent population regulation.

O_LINitrogen (N) is limiting for terrestrial herbivores, particularly over winter. Caribou (Rangifer tarandus) have adapted to seasonal scarcity of N by accruing muscle mass during the growing season when N is more abundant. C_LIO_LINitrogen stored in muscle tissue is then relied upon during winter to compensate for dietary deficits. Once their diet shifts from N-rich vascular plants to N-poor lichen over winter, caribou can lose [~]30% of their muscle mass. As catabolized N is shed in urine on wintering grounds, caribou could act as elemental transport across seasons and landscapes. Furthermore, if deposited N is taken up by lichen or other winter forage, it might enrich the nitrogen-poor winter diet of caribou in the future. C_LIO_LIWe tested this potential transport via three steps. We analysed Cladonia spp. lichen and vascular plants upon which caribou forage across Fogo Island, Newfoundland, using %N content as our metric of forage quality. We then compared seasonal habitat selection responses to forage quality by caribou using integrated step selection analyses. In summer, caribou selected areas with higher vascular plant %N but did not select nor avoid Cladonia quality. In contrast, caribou selected sites with higher quality Cladonia in winter but responded neutrally to vascular plant quality. C_LIO_LIWe compared seasonal distributions of caribou to determine whether nitrogen consumed in summer and deposited in winter would occur in spatially discrete locations. Population-level kernel density estimates for summer and winter in this island herd were mostly non-overlapping, lending credence to the potential landscape effects of N transport. C_LIO_LIWhen viewed together with established seasonal changes in woodland caribou physiology, sociality, and forage preferences, the shifts in habitat selection and seasonal ranges we observe here could serve as an adaptive strategy for caribou to recycle N and mitigate winter nutrient scarcity. C_LI

The spatial configuration of habitat patches is a key driver of metacommunity dynamics, yet the role of network topology remains poorly understood. In this study, I experimentally tested how different aspects of network structure influence metacommunity processes operating at different spatial scales. Using protist microcosms, I assembled metacommunities with patches connected as random or scale-free networks, and quantified occupancy, biomass, and extinction dynamics in relation to local (patch degree) and global (closeness centrality) metrics of connectivity. Scale-free metacommunities supported higher occupancy and biomass than random networks. At local scales, biomass declined with increasing patch degree, suggesting that reduced connectivity may enhance productivity, likely by limiting negative interactions. In contrast, extinction dynamics were not related to degree but strongly associated with patch centrality, with network topology modulating the relationship. These results reveal a decoupling between ecological processes, showing that different components of network structure can regulate dynamics at different spatial scales.

Habitat complexity (HC) in part determines the diversity, stability, and behavior of food webs and can influence predation according to a wide variety of functional relationships. Many aquatic species provide habitat complexity and are also consumed by other species (e.g., macrophytes, corals, mussels). However, food web theory does not readily account for these species that act as edible habitat complexity (EHC). Here, we combine existing theory on predator-prey interactions, HC, and prey switching to describe the role of EHC in benthic food web models. We dissect feedback loops in each model to demonstrate how self-regulation of the prey species, mediated by species densities and HC, drives that food webs behavior. HC can stabilize predator-prey interactions by coupling prey self-regulation with HC self-regulation. EHC can further stabilize predator-prey interactions across a wide variety of "HC functions" that relate HC to predation rates. Significance StatementHabitat complexity (HC) plays a critical role in trophic interactions, population dynamics, and food web stability. However, little theory exists to describe edible habitat complexity (EHC), where a species is both consumed and confers habitat complexity for other species. We provide a series of models demonstrating how HC and EHC alter the population dynamics and stability of simple aquatic food webs. HC is strongly stabilizing in food webs by providing safety in rarity for prey. EHC provides safety in rarity for both prey and the EHC species because their predators are omnivorous. Given the prevalence of EHC species in aquatic systems (e.g., macrophytes, corals, mussels), our models demonstrate the importance of maintaining EHC species in aquatic systems for stable food webs.

Conspicuous coloration in animals is generally thought to evolve and be maintained through inter- or intraspecific interactions such as mate choice, but this might not always be the case. The sight-line hypothesis proposes that conspicuous light-dark contrast in front of the eyes (hereafter, eyeline) evolves and is maintained due to viability selection, enhancing an individual visual acuity and thus evolutionarily associated with a particular foraging behavior that requires accurate aiming. However, empirical evidence that supports the sight-line hypothesis is virtually absent, with no studies demonstrating the key prediction that the direction of eyelines matters. Here, I tested the sight-line hypothesis using macroevolutionary analyses in terns and allies, which are a suitable study system, because they have variation in facial color patterns, including presence/absence and, if any, various angles of eyelines. They also have a large variation in foraging behavior, including picking, plunge diving, and skimming. As predicted by the sight-line hypothesis, tern lineages that require accurate aiming at foraging (e.g., plunge diving) are more likely to have eyelines. In addition, the evolutionary transition to the state with eyelines and these foraging behaviors was more likely to occur than the reverse transition. Furthermore, as expected by the fact that the direction of travel is upwardly deviated from the direction of the bills during skimming, the eyeline angle from bills was evolutionarily positively associated with the occurrence of skimming behavior. To my knowledge, the current study is the first to demonstrate that the direction of the eyeline matters, thereby strongly supporting the sight-line hypothesis.

Extreme climatic events impose considerable stress on organisms with consequences for key ecological interactions such as pollination. Because temperature directly affects metabolic processes, heat variations may also importantly influence the nutritional needs and feeding choices of animals. Here, we studied the effects of thermal stress on the nutritional choices and performances of bumblebees, using a 3D nutritional geometry design. At optimal temperature for colony development (30{degrees}C), bees successfully balanced carbohydrate, protein, and lipid collection, at levels beneficial for body weight and survival. Under cold stress (20{degrees}C), bees reduced their overall nutrient collection while selecting proportionally more carbohydrates, thereby prioritizing survival over weight gain. Under heat stress (35{degrees}C), nutrient balancing was disrupted and survival dropped. Notably however, across all temperatures, bees maintained stable lipid collection while flexibly adjusting the amount of carbohydrates and proteins, suggesting strong constraints on lipid regulation. Given the pivotal role of bees for pollination, identifying how their nutritional needs change in response to climatic conditions is of prior importance for food safety and the conservation of terrestrial ecosystems.

Spatial heterogeneity of abiotic resources is essential for species coexistence. Ecological theory often assumes predefined heterogeneity of resources that constrains community dynamics, but the recent developments of meta-ecosystem ecology and zoogeochemistry highlight nutrient patterns could result from the interactions between the activities and movements of organisms and their abiotic environment. Here we investigate the mechanisms by which biotic-abiotic feedbacks could generate nutrient spatial heterogeneity in a simple plant-herbivore occupancy model where populations forage, recycle, and disperse in a homogenous landscape. By systematically varying organisms ranges of foraging and dispersal, and recycling levels, we found that limited dispersal of plants plays a key role on the emergence of nutrient patchiness by favoring small clusters of vegetation that shape their environment through consumption and recycling. However, herbivores could also create nutrient spatial heterogeneity when large foraging and dispersal ranges, and high recycling, allow them to efficiently track plant hot spots and to increase population persistence. Unexpectedly, strong aggregation of herbivore populations did not necessarily result in nutrient clustering. Rather than via recycling, herbivores mainly affected nutrient distribution indirectly, through their top-down impact on plant distribution. When evenly spread in the landscape, herbivore populations with large foraging ranges created areas of strong herbivory pressure unfavorable to plant colonization where nutrient can accumulate. These results can help understand the dynamical feedback between biota and abiotic resources. In a context where human activities alter both nutrient distribution and species abundances, a better understanding of this biotic-abiotic feedback will be key to anticipate the response of ecosystems to current perturbations.

1. Biotic insular systems differ from conventional islands because patch attributes change dynamically as patch-forming organisms develop. It therefore remains unclear whether the assembly mechanisms predicted by island biogeography theory (IBT) operate in such systems. Here, using epiphytic birds nest ferns (BNFs, Asplenium nidus) as a model biotic island system, we tested whether fungal and bacterial community diversity conform to species-area relationships predicted by IBT. With a stratified sampling scheme, we further evaluated the underlying mechanisms (passive sampling, disproportionate effects, and environmental heterogeneity) of species-area relationships, and assessed isolation effects using distance-decay patterns in community similarity. 2. We treated each BNF individual as a microbial island and categorized 24 BNFs into three size classes. Microbial and humus samples from multiple litter layers within each BNF individual were collected; microbial communities were characterized using next-generation sequencing, and humus chemical properties (pH and C:N ratio) were measured to characterize microhabitat conditions. To investigate mechanisms underlying species-area relationships, we applied a multi-scale rarefaction framework to partition diversity components. Spatial distances among BNFs were quantified to evaluate isolation effects. 3. Consistent with IBT predictions, both fungal and bacterial communities exhibited positive species-area relationships, indicating that larger BNFs harbored greater microbial richness. Diversity partitioning suggested that fungal richness increased through both disproportionate effects and environmental heterogeneity, whereas bacterial richness was primarily driven by environmental heterogeneity. Within larger ferns, greater heterogeneity in litter pH was associated with increased species turnover across litter layers, suggesting that decomposition-driven pH gradients create diverse microhabitats that promote microbial diversity. In addition, both microbial communities exhibited distance-decay patterns, indicating that isolation contributes to community assembly through dispersal limitation. 4. Synthesis. Our results demonstrate that BNFs function as a biotic insular system, in which both patch size and spatial isolation structure microbial diversity, consistent with predictions from IBT. Furthermore, we show that environmental heterogeneity generated by the growth of the habitatforming BNF mechanistically links island area to microbial diversity. Our study integrates both local habitat heterogeneity and regional spatial structure, highlighting the potential to extend IBT and metacommunity theory to organism-formed habitats.

Seasonal rainfall shapes biological responses in tropical ecosystems, yet how tropical organisms integrate behavioral and physiological responses to cope with seasonality remains poorly understood. We assessed how four poison frog species with contrasting reproductive strategies respond to dry and wet season environmental conditions. We quantified spatial behavior, microhabitat use, hormone concentrations, and chemical defenses in two seasonal breeders (Allobates femoralis and Ameerega trivittata) and two year-round breeders (Ameerega macero and Ameerega shihuemoy). Seasonal breeders exhibited pronounced sex-specific shifts in space use, where males expanded their space use during the wet season, likely to track reproductive opportunities, while A. femoralis females increased their spatial use during the dry season, likely responding to foraging demands when prey resources are sparse. Year-round breeders maintained similar space use across seasons, likely reflecting their ability to access key resources within the same space to reproduce year-round. Microhabitat use was flexible, as seasonal breeders shifted toward humid refugia during the dry season and reproduction-associated microhabitats during the wet season, whereas year-round breeders selected microhabitats that facilitate continuous reproduction across seasons. Despite these behavioral responses, corticosterone, testosterone, and chemical defenses showed no consistent seasonal variation, suggesting that behavioral flexibility is decoupled from seasonal variation in these measured physiological responses. Our study suggests that poison frogs are able to buffer environmental fluctuations through behavioral flexibility. However, given the increasing unpredictability in rainfall timing and intensity as a result of climate change, how these coping strategies will function in the long term is uncertain.

Protogynous sex change, where individuals first function as females and later as males, is a key life-history strategy among polygynous reef fishes. In haremic systems, sex change is typically socially regulated, with dominants suppressing subordinates sex change through aggression. Females within a harem form a size-based hierarchy that can remain stable in most species through the threat of eviction. We studied a different situation in the cleaner wrasse Labroides dimidiatus, where larger females have incomplete control, as they spend most of their time alone at their own cleaning territory. We tracked over 400 individuals for 12 months, recording growth, behavior, social organization, and sex change. We confirmed earlier reports that both sexes direct aggression primarily at those ranked immediately below them. However, we observed 30 cases where smaller females outgrew larger ones, revealing hierarchy instability. Of 42 sex change events, 43% occurred in presence of the male, and half of these early sex changers were not the largest female, but individuals overlooked by the male. Fast growth relative to harem-mates and harem switching increased the likelihood of sex change. Local population densities also influenced growth and sex change, with individuals in high-density demes growing faster and changing sex at larger sizes. Our findings reveal flexible sex change dynamics in a system with incomplete social dominance. Such incomplete control and observations that becoming male confers both higher reproductive success and survival highlight the need to expand game-theoretical and life-history frameworks to encompass such strategic flexibility. Lay summaryDominant cleaner wrasse cannot fully control subordinates as individuals occupy distinct core areas. Tracking 400 fish for a year, we found that smaller females could outgrow initially larger ones, and early sex change despite a larger male. Fast growth and harem switching increased the chances of becoming male. Population density also shaped these strategies. Our findings reveal flexible sex change dynamics in a system where becoming male confers both higher reproductive success and survival.