BEHAVIORAL ECOLOGY

Impact of human presence and activity on urban Eurasian red squirrels' innovative problem-solving
Chow PKY, Loukola OJ and Solvi C
Humans impact wildlife positively and negatively, and increasing evidence shows that humans potentially play a major role in shaping urban wildlife cognition. However, it remains unclear which, and how specific anthropogenic factors, shape animal cognitive performance. Here, across 15 urban areas in Oulu, Finland, we investigated how varied levels of human presence nearby, types of human activity (walking, dog-walking, cycling, and playground activities), and distance to the nearest footpaths influenced 64 squirrels' innovative problem-solving ability-measured as the proportion of solving success at the site level, solving outcome at the individual level as well as individuals' first-success latency. Higher mean human presence nearby and all measured human activities significantly decreased the proportion of success at the site level. Playground activity showed the highest negative impact on both the first- and subsequent-visit success rate at the site level. Increased mean human presence and walking decreased the likelihood of a squirrel successfully solving the novel food-extraction problem. When examining the problem-solving latency of individual squirrels, increased human presence also decreased squirrels' first-success latency, and dog-walking was the outstanding factor affecting first-success latency. These results show the negative effects of specific human-related factors on an important cognitive trait, problem-solving ability. These factors may also potentially exert selective pressure on shaping urban wildlife cognition.
Boat noise alters individual behaviors but not communication between partners in a fish-shrimp mutualism
Manera JL, Martin JM, Palacios MM, Mason RT, McCormick MI and Wong BBM
Persistent noise pollution produced by boat traffic is reshaping marine soundscapes globally. Despite growing ecological concern, most studies to date have focused on individual-level effects under laboratory conditions, leaving major gaps in our understanding of how boat noise shapes species interactions in the wild. Using field-based behavioral assays, we investigate how boat noise from different engine types (4-stroke and 2-stroke) affects the mutualistic partnership between Steinitz's goby () and snapping shrimp ( spp.). Across 123 partnerships, we recorded behavioral responses before, during, and after noise exposure. Gobies increased burrow use during 4-stroke boat noise exposure, while shrimp responded stronger to 2-stroke noise-reflecting taxon-specific sensitivities to different noise spectra. Despite these shifts, tactile partner communication was not affected by boat noise. These findings highlight divergent vulnerabilities between species tied to different engine acoustics and emphasize the need for targeted research to inform strategies for mitigating marine noise pollution.
Good guardian, bad parent: tradeoffs between territory defense and parental care in Darwin's finches
Katsis AC, Common LK, Akçay Ç and Kleindorfer S
Although defending a territory may benefit individuals by allowing them to retain important resources, the time and energy costs associated with territory defense may lead territory owners to neglect other reproductively important behaviors. In this study, we assessed the potential tradeoff between territory defense and parental care in 4 Darwin's finch species on Floreana Island, Galápagos. Using song playback, we simulated territory intrusions to measure male aggressiveness across multiple stages of the breeding cycle (unpaired, paired, incubating, and chick feeding). To quantify parental care at each nest, we conducted 1-h observations to record the frequency of male food deliveries and the duration of female incubation and brooding. Male aggressiveness toward a perceived intruder did not change across the breeding cycle and was highly repeatable (adjusted = 0.597), although responses became less vocal across the breeding cycle. A male's aggressiveness did not predict his frequency of food deliveries during incubation or chick feeding, although females paired with more aggressive males spent significantly less time incubating. This finding provides weak evidence for a tradeoff between territory defense and parental care, although the behavioral mechanisms mediating this relationship remain uncertain. Finally, males with aggressive and nonaggressive behavioral phenotypes did not differ in their hatching success, although additional work is needed to assess other measures of individual fitness.
Prey movement, size, and glossiness interact to impact praying mantid attack behaviors
Wang Y, Stuart-Fox D, Henríquez-Piskulich P and Franklin AM
When moving, animals are vulnerable to predation because movement can rapidly attract the attention of a predator. To reduce the risk of predation while moving, animals can use a variety of different strategies (eg erratic movement, coloration). These strategies often work in combination or may be influenced by other prey characteristics (eg size), but few studies have explored these interactions. Here, we investigate how movement trajectory (linear or erratic), prey size (small or large) and prey coloration (glossy or matte) interact to impact the attack behavior of giant rainforest mantids (). We presented mantids with animations of moving targets and filmed their response with a high-speed camera. As expected, mantids were more likely to track large than small targets and targets moving linearly than erratically. Counterintuitively, however, mantids were quicker to strike at erratically moving targets, perhaps because they more closely resembled preferred prey. When mantids attacked the target, their accuracy was influenced by the interaction of target trajectory and glossiness. As predicted, mantids had larger attack errors (ie lower accuracy) toward erratically moving glossy targets compared with linearly moving glossy targets or erratically moving matte targets. However, contrary to our prediction that linearly moving matte targets would be easiest to capture, these targets also elicited large attack errors, similar to those recorded for erratically moving glossy targets. Together, our results demonstrate that anti-predator tactics for prey in motion may interact in complex ways, and simple experimental scenarios may overlook context-dependent effects that emerge when multiple factors interact.
Site-level variation in field of view is associated with altered anti-predator responses in farming damselfish
Boon JS, Stratford JE, Lynch J, Yesson C, Field R, Exton DA and Keith SA
The three-dimensional (3D) structure of habitats influences how prey detect and respond to predators, but the specific roles of different aspects of structural complexity remain poorly understood, particularly in coral reef ecosystems. We used 3D models of 3 Caribbean reef sites to quantify 3 structural metrics at site level: field of view (the extent of observable area), refuge density (density of holes), and rugosity (reef surface roughness). We then observed the anti-predator behavior of damselfish, parrotfish, and wrasses at each site. Territorial damselfish showed species-specific responses to habitat structure, especially in relation to field of view. , for example, exhibited shorter flight initiation distances (FIDs) at the site with the highest field of view, consistent with expectations from optimal escape theory. In contrast, wrasse and parrotfish species showed little variation in behavior across sites, though larger individuals tended to have longer FIDs and flight distances. Refuge density was similar across sites, likely reflecting long-term regional loss of fine-scale complexity in the Caribbean. While rugosity is widely used as a proxy for reef complexity, our results suggest that field of view may be more strongly associated with differences in anti-predator behavior, particularly in damselfish. These findings highlight the need to assess multiple dimensions of habitat structure, as even closely related species may exhibit distinct behavioral adaptations to their 3D environment.
Honeybees show an increased preference for dietary alcohol when parasitized
Ostap-Chec M, Antoł W, Bajorek D, Stec D and Miler K
Parasitic infections often alter host behavior, including foraging and the consumption of bioactive substances. In honeybees (), infection with the common gut parasite causes metabolic disruption and increased mortality. Ethanol is a naturally occurring bioactive compound found in nectar, and honeybees exhibit high tolerance and resilience to chronic exposure. However, whether honeybees actively use ethanol during infection remains unclear. Here, we investigated whether -infected honeybees alter their ethanol consumption. In a feeding experiment, infected and uninfected honeybees were given a choice between plain sucrose solution and ethanol-spiked food (0.5% or 1% ethanol). We measured food consumption, survival, and spore load. Although overall food intake did not differ between groups, infected honeybees consumed a significantly higher proportion of ethanol-spiked food. Survival analysis showed that a diet containing 1% ethanol caused higher mortality than a diet containing 0.5% ethanol; however, among honeybees on a 1% ethanol diet, this negative effect was less pronounced in infected individuals than in controls. Spore load did not differ between treatments. These results suggest that infection induces a shift in feeding behavior toward increased ethanol intake, which may benefit infected honeybees by reducing mortality. This may reflect a self-medication response, although alternative explanations remain possible. Further research into ethanol's effects on spores is needed. Nonetheless, our findings provide insights into honeybee interactions with bioactive compounds and suggest that ethanol may be a behaviorally relevant dietary substance.
Latent layers in social networks and their implications for comparative analyses
De Moor D, Hart JDA, Franks DW, Brent LJN, Silk MJ and Brask JB
Animal social systems are remarkably diverse, ranging from solitary individuals to well-connected cooperative groups. Understanding the drivers of this variation is a key question in behavioral ecology and has been the focus of numerous studies linking social structure to ecological, demographic, and life history patterns within groups, population, and species. Equipped with this information, researchers are now turning to investigations that are comparative in nature. However, comparing social networks remains a considerable logistical and analytical challenge. Here, we present the , which outlines how are linked to the 2 underlying latent networks that are of interest for most research questions: the (the actual pattern of social interactions), and the driving these interactions. This conceptual framework provides a clear and unified approach to understand when and why differences in network properties and sampling protocols can introduce discrepancies between observed and latent networks, potentially biasing or confounding statistical inference. We then use this conceptual framework to outline some of the central challenges to comparing animal social networks, describe why and how they create challenges for comparative analyses, and suggest potential directions for solutions. The can help researchers to identify networks they can (or cannot) compare. In doing so, this framework facilitates advances in comparative social network studies with the potential to generate new and important insights into the ecological and evolutionary drivers of variation in social structure across the animal kingdom.
Complex dynamics of social learning in groups of wild Arabian babblers
Aljadeff N, Keynan O and Lotem A
We studied the effect of a demonstrator on the learning of a novel foraging task in 12 groups of free-living cooperative breeding Arabian babblers (). We allowed naïve babblers to forage jointly on a foraging grid with a demonstrator previously trained to solve a task in one of 2 possible methods: lifting covers of 1 color or pecking through covers of another color. We found that most group members learned to solve the task using one of the methods, and persisted with it even when later tested with covers of a third (neutral) color that could be opened by both lifting and pecking. However, the method learned by group members did not necessarily follow the method used by the pre-trained demonstrator. Instead, learners within each group tended to use the same method (significantly more than expected by chance), and the extent to which groups differed from the demonstrator was correlated with the extent to which the demonstrator occasionally (and quite rarely) exhibited also the alternative method. These results, together with further analysis of the sequence of events in each group, suggest that both naïve birds and demonstrators learn socially from each other, as well as through individual trial-and-error learning, which enables naïve individuals to become demonstrators themselves and influence the pattern of social transmission. This process mostly leads to a homogenous group behavior, but one that cannot be predicted by the seeded demonstration.
Background color matching affects sexual behavior, growth, and mortality rate in an African cichlid
Moore TI, Bright WG, Bell WE, Solomon-Lane TK, Alvarado SG and Dijkstra PD
Phenotypic plasticity allows organisms to adapt to changing environments within their lifetimes. However, environmentally induced changes in the plastic trait of interest may influence a range of fitness-related traits due to trade-offs, pleiotropy, linkage, or epistasis of genes regulating the plastic trait. These correlated responses may constrain or facilitate the evolution of plasticity, but their evolutionary implications are often poorly understood due to a lack of data on their direction and magnitude. Males in the African cichlid are blue or yellow, and males are able to adjust their body coloration to the color of the background, presumably to increase crypsis. To test whether background color influences fitness-related traits, we raised mix-sex groups of juvenile to adulthood in yellow or blue tanks. We found that more males adopted the blue phenotype in blue tanks while more males adopted the yellow phenotype in the yellow tank, though the degree of background color matching decreased with age. Males, but not females, from blue tanks showed earlier sexual maturation than those held in yellow tanks. However, across the duration of the experiment, there was a higher occurrence of breeding in females housed in yellow tanks than those that were housed in blue tanks. In addition, fish in blue tanks exhibited reduced growth rate but higher survivorship relative to their yellow-reared counterparts. Our data suggest that background color affects important fitness-related traits in a color polymorphic cichlid, which may influence the evolution of phenotypic plasticity.
Integrating space, time, and culture in animal conservation practice
Oestreich WK, Barlow DR and Hersh TA
Recent theoretical integration of the spatiotemporal and cultural elements of animal behavior has led to increasing calls to incorporate animal culture into conservation. Implementation of this idea remains sparse due to disconnects between the theoretical concept of considering culture in animal conservation and the spatiotemporal approaches typically employed in conservation practice. Here we propose that this gap can be bridged by (1) clarifying that spatiotemporal conservation interventions inherently interact with culture regardless of whether this connection is acknowledged; and (2) strategically considering feasible "entry points" for considering animal culture in conservation practice. Recent advances in dynamic management strategies indicate the capacity for modern conservation approaches to integrate additional dimensions of animal behavior, and could serve as a particularly fruitful space for considering culture. Drawing on instructive examples from cetaceans, we examine instances where protection in space and time can facilitate the conservation of culture, and where focusing on conserving culturally distinct groups can yield protection in space and time. Human interventions that explicitly consider these interwoven dimensions in practice are achievable and can enable more holistic protections for diverse taxa.
Plan ahead, or wing it? How storm-petrel parents adjust food delivery to young chicks
Mauck RA, Taylor LU, Neirink SC, Lichtner KE, Chapman SE, Veitch JH, Kyle IT, Haussmann MF and Jones PL
Parents must decide how to allocate energy gained from foraging between self and offspring. Storm-petrels (Procellariiformes: Hydrobatidae) are pelagic seabirds that travel hundreds of kilometers across multiple days before returning to the nesting burrow to feed a dependent chick. Parents return to the nest with food stored in the proventriculus, a portion of which is regurgitated to their offspring. As the chick grows, provisioning demands increase. However, it is unknown whether parents meet this increasing demand by (1) altering their foraging strategies to acquire more food or (2) allocating a greater proportion of their intake to the chick. We designed, validated, and implemented a new technology-the Burrow Scale Monitor-to measure Leach's storm-petrels () as they entered and exited the nesting burrow. We monitored breeding adults over the first 30 d of chick rearing to determine whether storm-petrel parents adjust their foraging intake to the age of the chick or simply adjust energy allocation at the nest. Food delivery increased with chick age, but this increase was driven to a much greater extent by parents delivering a greater proportion of their body mass as food (ie, a shift in parental allocation) rather than by adults adjusting their foraging strategy to match chick age. Only by measuring adult body mass on arrival and exit at the nesting burrow could we understand how parents adapt their provisioning strategy to the increasing demands of the growing chick.
Perch choice and substrate matching to the dorsal patterns of lizards
Salisbury JW and Peters RA
The backgrounds that cryptic animals choose will affect the efficacy of their camouflage. Most animals use a range of microhabitats consisting of a variety of substrates, vegetation and lighting conditions. As some of these will be better suited to facilitating camouflage than others, we expect cryptic animals to consider their conspicuousness when choosing a background to occupy. If the availability of backgrounds varies between populations of cryptic animals, then selective pressure on their coloration may also vary, resulting in intraspecific variation and presumably animals being better suited to the backgrounds locally available to them than those at other locations. In this study we investigate how backgrounds available to Jacky dragons () vary across their range, whether these lizards are occupying backgrounds that match well to their dorsal patterns, and how backgrounds compare to their dorsal patterns. Wild lizards were located and photographed along with the background they were found on, and other options available nearby. We compared lizards and backgrounds within their microhabitat as well as all backgrounds across all microhabitats. We found that lizards were not occupying the backgrounds that best matched their own patterns, that background options varied between locations, and that lizards from certain locations were occupying backgrounds closer matching to their own pattern than those from other locations. These outcomes provide interesting insight into the variance of local factors that influence the pattern phenotype, as well as how the relative need for camouflage might vary and be balanced with other needs.
High spatial pair cohesion during and after breeding in a socially monogamous territorial passerine
Speelman FJD, Tyson CW, Naguib M and Griffith SC
Long-term social monogamy, a prevalent mating system in avian species, is often associated with increased cooperation and coordination as well as reduced sexual conflict. Although many studies have highlighted the benefits of long-term partnerships for individuals, there remains a lack of insight into how closely partners associate with one another behaviorally. To date, studies investigating pair cohesion in seasonal and long-term partnerships are typically restricted to arrivals at the nest or feeding sites during the breeding season. Using fine-scale automated tracking data on chirruping wedgebills (), a territorial socially monogamous species, we characterized how partners coordinate their movement during and after the breeding season. We used 12 pair-bonded individuals with consistently high localization rates that were tracked for a period between 32 and 69 days, with an average of 260,000 localizations per individual. We demonstrate that pairs (1) had extremely similar home ranges with a similarity index of 0.93 versus 0.18 for non-pairs, (2) maintained consistently closer proximity than expected from movement without paying attention to a partner, and (3) followed each other as they moved, with individuals following their moving partner in 42% of cases during and in 47% of cases after breeding. Our findings show that pair cohesion in socially monogamous territorial species can be very high in both a breeding and non-breeding context, illustrating that strong coordination among partners has important functions beyond reproduction and parental care.
Comparative evaluation over time during mate choice in the green swordtail
Bullough K, Kuijper B and Kelley LA
During mate choice, choosers are exposed to a variety of sexual signals varying in both magnitude and the environment in which they are experienced. Previous work assumes that choosers evaluate signal variation from potential mates that are simultaneously viewed and compared. However, this is an extreme scenario, and it is important to also consider sequential presentation of mates, as most animals likely experience both scenarios during mate choice. Using green swordtail fish (), we assessed whether female preferences for larger males differed when males of different sizes were experienced simultaneously or sequentially. We also investigated the perceptual mechanisms of comparison in both contexts, given recent research suggesting that female preferences are often nonlinear. We found that females consistently preferred larger males, irrespective of whether males were experienced simultaneously or sequentially. However, female preferences were stronger for a male of a given size when viewed under simultaneous, compared with sequential, conditions. During sequential presentation, females compared information from both previously and currently presented males, and interest did not decay with subsequent presentations. Previous research has demonstrated that female green swordtails assessing males simultaneously attend to the relative size difference between males, but we found no evidence of any comparative size assessment. Our study demonstrates that when designing mate choice experiments, it is important to consider how females encounter potential mates in the wild, highlighting that there are clear differences in preferences due to methods of mate presentation and that stochastic adjusting of internal standards of quality frequently occur.
Larger frogs are better mimics but are more risk-averse in a nontoxic poison frog
McEwen BL, Yeager J, Veneat A and Barnett JB
Aposematic species signal to potential predators with salient and recognizable coloration. Predators learn to associate these warning signals with secondary defenses (eg toxins) and will subsequently avoid attacking aposematic prey. Warning signals can therefore reduce the need to hide and/or flee and alleviate some of the energetic/opportunity costs of predator avoidance. Consequently, aposematic species are frequently active and bold in behavior. Batesian mimics replicate the colors, and often the behavior, of aposematic species and may benefit from a similar reduction in predation risk and energetic/opportunity costs. (Aromobatidae) is a nontoxic Batesian mimic of the chemically defended poison frog (Dendrobatidae). However, the efficacy of mimicry appears to change throughout ontogeny as develops from a seemingly cryptic juvenile to the mimetic adult. We examined how morphological mimicry (ie color) and the propensity to explore a novel environment (ie boldness) changed throughout ontogeny. We predicted that mimicry would improve with increasing size and that better mimics would engage in more exploratory behavior. We found that larger mimics more closely matched their model however they were less likely to be active than were smaller frogs. These data suggest that larger size, and more accurate mimicry, do not necessarily correspond to increases in behavioral boldness. This result may arise from limitations in Batesian mimicry but factors including foraging requirements or social/reproductive behavior cannot be discounted. More research is needed to understand the relationship between behavior, color, body size, and maturity in these frogs and across Batesian mimics more widely.
Does rainfall or temperature influence antipredator vigilance in a hibernating mammal?
Bobb K, Adler KA, Martin JGA and Blumstein DT
As the global climate changes, temperatures are rising, snow is melting earlier, and rainfall is becoming more variable, and these climatic changes may create an ecological mismatch. While prior work has shown how animals respond to these changes physiologically and behaviorally, few have specifically investigated antipredator behavior, an essential activity. In many species, there are direct fitness tradeoffs between allocating time and energy to antipredator vigilance and foraging. To discover how these tradeoffs are affected by climate change, we studied how temperature, snowmelt date, and rainfall affected the proportion of time yellow-bellied marmots () allocated to vigilance during bouts of foraging. While snowmelt and temperature did not explain variation in vigilance, rainfall did. Higher rainfall in the week prior to a focal observation was associated with higher vigilance, possibly reflecting more abundant food that affords the luxury of increasing antipredator vigilance while foraging. Such an effect might be consequential at the population level given the importance of foraging and antipredator behaviors for a highly time restrictive hibernating species. Further research is necessary to determine consequences at the population level and whether and how these findings extend to other species.
Double brooding in house wrens is repeatable but constrained by time-of-season
Leischner L, Sakaluk SK, Thompson CF and Nietlisbach P
Birds in some species produce 2 broods in a season to maximize reproductive success, but in species such as the northern house wren (), not all individuals produce a second brood even if they have enough time to do so. We investigated whether variation in individual quality, in addition to time-of-season, explains some of the variation in the likelihood of producing a second brood. We examined the effects of individual age, body condition, and territory quality, and estimated the repeatability of producing a second brood using breeding records from a long-term dataset. We also cross-fostered eggs between earlier-nesting females (presumed high quality) and later-nesting females (presumed low quality) to delay or advance the natural nesting date, exposing an effect of quality if present. There was little evidence that the individual traits measured affected the production of a second brood. Cross-fostered, delayed females did not differ from the control in their likelihood of producing a second brood, but advanced females were more likely to produce a second brood compared with the control. However, the production of a second brood was significantly repeatable, and this repeatability was substantially higher after accounting for variation in the timing of the first brood. This suggests that individual characteristics, in addition to time-of-season, have a substantial effect on the likelihood of producing a second brood, but that further studies are needed to identify the specific traits responsible for this effect.
Long-term fitness effects of the early-life environment in a wild bird population
Sun Y, Burke TA, Dugdale HL and Schroeder J
Environmental conditions and experiences during development can have long-term fitness consequences, including a reduction of adulthood survival and reproduction. These long-term fitness consequences may play an important role in shaping the evolution of life history. We tested two hypotheses on the long-term fitness effects of the developmental environment-the silver spoon hypothesis and the internal predictive adaptive response (PAR) hypothesis. We compared the change in annual survival and annual reproductive output with age for adult birds hatched and/or reared in poor--impacted by anthropogenic noise, and/or high sibling competition--and good--not impacted by anthropogenic noise, and/or low sibling competition--environments. We used a 23-year longitudinal dataset from a wild house sparrow () population inhabiting an isolated island, which enabled near-complete monitoring and unusually accurate lifetime fitness estimates. We used a cross-fostering setup to disentangle environmental effects experienced postnatally from those experienced prenatally. We found that adults that, as chicks experienced more within-brood competition had a stronger increase in early-life annual survival, but also a stronger decrease in late-life annual survival. Females that hatched in a noisy environment produced fewer genetic recruits annually, supporting a sex-specific silver spoon hypothesis. Males reared in a noisy environment had accelerated reproductive schedules, supporting a sex-specific internal PAR hypothesis. Our results highlight that anthropogenic noise (∼68 dB from power generators) can have long-term fitness consequences in wild animals, altering their life-history strategies, and that effects may be sex-specific.
Maternal care shapes an aposematic display and provides lifelong protection against predators
Lindstedt C, Boncoraglio G, Cotter SC, Gilbert JDJ and Kilner RM
Parental care can improve early offspring survival against predators by providing protection and resources. However, we have little knowledge of how its effects shape predator-prey interactions later in life. We investigated this with the burying beetle which provides care for offspring and carries warning coloration to advertise its chemical defenses to predators. Warning displays by prey are selected by predators for uniformity and to reliably advertise the extent to which individuals are chemically defended. We investigated whether the strength of the correlation between the conspicuousness of the warning display and the potency of the chemical defenses depends on levels of care received during development by manipulating the level of maternal care received by larvae and tracking the effects into adulthood. We found that individuals that received limited care, developed into smaller adults with less conspicuous warning displays. The correlation between the visual display and the chemical defense was also weaker when broods received little care as larvae. We conclude that maternal care received by burying beetles modulates the information content of aposematic defense: less care makes signals less reliable. Our results further suggest that the prey's social environment could constrain the response to selection from predators on warning signal reliability.
Mating status-dependent "choice" in competitive and noncompetitive arenas
Dugand RJ, Lymbery RA, Dewan N, Kennington WJ and Tomkins JL
To maximize their reproductive fitness, females of many polyandrous species should display mating status-dependent choice, where they mate relatively indiscriminately once to ensure reproductive output, and then become choosy and mate preferentially with higher-quality males. Despite this potential contrast in choosiness, most mate choice experiments use virgin females. Here, using a panel of 20 isofemale strains that originated from wild-caught flies, we allowed virgin and non-virgin females to choose among males from the same panel of strains. We used single-male latency trials and a series of male competition trials to help disentangle female "choices" from male-male competitive effects. Most virgin females mated within 2 h of males being introduced, compared with fewer than half of non-virgin females mating over the same period. However, despite mating more rapidly, virgin females did not mate indiscriminately, and their "choices" strongly aligned with those of previously mated females across both the single-male latency and male-male competition trials. Our results challenge the idea that virgin females mate relatively indiscriminately and show that female choice may be more stable than is generally appreciated.
The interplay between satiation and temptation drives cleaner fish foraging behavior and service quality toward client reef fish
Triki Z, Li Richter XY, Pinto A, Baud A, Binning SA, Aellen M, Emery Y, Staubli V, Raihani N and Bshary R
Supply and demand affect the values of goods exchanged in cooperative trades where high demand typically leads to a higher cost. An exception has been described in the marine cleaning mutualism involving the cleaner fish and its variety of "client" coral reef fishes. Cleaner fish feed on clients' ectoparasites (ie gnathiid isopods) but prefer eating clients' mucus instead, which constitutes cheating. Here, we provide field observations, followed by a set of laboratory experiments with real client fish and Plexiglas feeding plates as surrogates for clients. In the field and in three experiments with real clients, we found that satiated cleaner fish were more cooperative, even though low hunger levels should make them less dependent on cleaning interactions. Similarly, the more abstract version of the cleaner-client experiments using Plexiglas plates offering two food types as stand-ins for client ectoparasites and mucus showed that satiation led cleaner fish to feed more against their preferences-an indicator of cooperative behaviour. However, this outcome occurred only if the temptation to eat the preferred food was low. When temptation to cheat was high, cleaner fish did so. We provide a further general support to these findings with a game-theoretic model. Many mutualisms involve food as a commodity. Thus, identifying foraging decision rules will enhance our understanding of how individuals adjust to variations in market conditions in real-time rather than playing a fixed strategy based on average market conditions.