The evolution and stability of helping behaviour has attracted great research efforts across disciplines. However, the field is also characterized by a great confusion over terminology and a number ...of disagreements, often between disciplines but also along taxonomic boundaries. In an attempt to clarify several issues, we identify four distinct research fields concerning the evolution of helping: (1) basic social evolution theory that studies helping within the framework of Hamilton's inclusive fitness concept, i.e. direct and indirect benefits, (2) an ecological approach that identifies settings that promote life histories or interaction patterns that favour unconditional cooperative and altruistic behaviour, e.g. conditions that lead to interdependency or interactions among kin, (3) the game theoretic approach that identifies strategies that provide feedback and control mechanisms (protecting from cheaters) favouring cooperative behaviour (e.g. pseudo-reciprocity, reciprocity), and (4) the social scientists' approach that particularly emphasizes the special cognitive requirements necessary for human cooperative strategies. The four fields differ with respect to the 'mechanisms' and the 'conditions' favouring helping they investigate. Other major differences concern a focus on either the life-time fitness consequences or the immediate payoff consequences of behaviour, and whether the behaviour of an individual or a whole interaction is considered. We suggest that distinguishing between these four separate fields and their complementary approaches will reduce misunderstandings, facilitating further integration of concepts within and across disciplines.
Pairs of unrelated individuals face a prisoner’s dilemma if cooperation is the best mutual outcome, but each player does best to defect regardless of his partner’s behaviour. Although mutual ...defection is the only evolutionarily stable strategy in one‐shot games, cooperative solutions based on reciprocity can emerge in iterated games. Among the most prominent theoretical solutions are the so‐called bookkeeping strategies, such as tit‐for‐tat, where individuals copy their partner’s behaviour in the previous round. However, the lack of empirical data conforming to predicted strategies has prompted the suggestion that the iterated prisoner’s dilemma (IPD) is neither a useful nor realistic basis for investigating cooperation. Here, we discuss several recent studies where authors have used the IPD framework to interpret their data. We evaluate the validity of their approach and highlight the diversity of proposed solutions. Strategies based on precise accounting are relatively uncommon, perhaps because the full set of assumptions of the IPD model are rarely satisfied. Instead, animals use a diverse array of strategies that apparently promote cooperation, despite the temptation to cheat. These include both positive and negative reciprocity, as well as long‐term mutual investments based on ‘friendships’. Although there are various gaps in these studies that remain to be filled, we argue that in most cases, individuals could theoretically benefit from cheating and that cooperation cannot therefore be explained with the concept of positive pseudo‐reciprocity. We suggest that by incorporating empirical data into the theoretical framework, we may gain fundamental new insights into the evolution of mutual reciprocal investment in nature.
The last decade has seen lots of studies on ‘animal personality’ (i.e. the study of consistent between‐individual behavioural differences). As timely and promising as this field is, its development ...has come with a diversity of research questions. As an unfortunate consequence, it now suffers from substantial confusion about what ‘animal personality’ is, and how relevant related research frameworks are. Here, we stress the current inconsistencies and sources of confusion pertaining to the field, and their consequences on terminology used and miscommunication between researchers. In an attempt to unravel and clarify the concepts underlying the field, we identify two distinct, but complementary, theory‐driven conceptual frameworks: the intra‐individual variability (IIV) approach and the life‐history (LH) approach, which we believe encompass the vast majority of existing ‘personality studies’. Finally, we argue in favour of theory‐driven studies of consistent behavioural differences and state that the integrative statistical properties of random regression models should not override the merit of alternative conceptual frameworks. We then provide brief guidelines and warnings for a parsimonious and sound use of terminology.
Predation on parasites is an important ecological process, but few experimental studies have examined the long-term impacts on the prey. Cleaner fish prey upon large numbers and selectively feed on ...the larger individuals of the ectoparasitic stage of gnathiid isopods. Removal of cleaner fish Labroides dimidiatus for 1.5–12.5 years negatively affects coral reef fishes, but the mechanism is unclear. A reduction in local parasite populations or the size of individual parasites would benefit all susceptible fishes. We tested whether cleaner presence reduces local gnathiid populations using 18 patch-reefs distributed between two sites (both at Lizard Island, Great Barrier Reef) which were maintained cleaner-free or undisturbed for 12 years. Using emergence traps (1 m²), free-living gnathiid stages were sampled before and after cleaner fish were removed during the day and night, up to 11 times over the course of the experiment. There were effects of the removal in the predicted direction, driven largely by the response at one site over the other involving 200% more gnathiids, but manifested only in the daytime sampling after 4 months. There was also a main effect (36%) for the shared sample dates at both sites after 12 years. Gnathiid size occasionally differed with cleaner presence, but in no consistent way over time. Contrary to our predictions, changes in free-living gnathiid population numbers and their size structure rarely reflected the changes in fish populations and individuals observed on cleaner-free reefs. Therefore, evidence that this predator alone regulates gnathiids remains limited, suggesting other contributing processes are involved.
Mutualisms are pivotal in shaping ecological communities. Iconic images of cleaner fish entering the mouths of predatory fish clients to remove ectoparasites epitomize their mutual benefit. ...Experimental manipulations of cleaner wrasse reveal declines in fish size and growth, and population abundance and diversity of client fishes in the absence of cleaner wrasse. Fishes grow more slowly and are less abundant and diverse on reefs without cleaner wrasse, both for larger species that are regularly cleaned and have high ectoparasite loads (“attractive species”), and for those smaller species that are rarely cleaned and are rarely infested with parasites (“unattractive species”). We therefore considered whether these previously observed declines in individual and population parameters on reefs without cleaners were related to increased ectoparasite infestation using an attractive species (
Hemigymnus melapterus
, Labridae) and an unattractive species (
Pomacentrus amboinensis
, Pomacentridae). Traps with these fish as a form of bait were deployed to sample blood-sucking gnathiid ectoparasites (Gnathiidae: Isopoda) on reefs from which cleaners (
Labroides dimidiatus
, Labridae) have been removed for 13 yr. Cleaner fish could not enter traps to access the clients/hosts, but gnathiids could enter the traps to infest hosts; thus, this method sampled the indirect effect of cleaners on gnathiid infestation of fish. Infestation was higher on reefs without cleaners than on those with them. The effect was only detected during the daytime when cleaners are active and only on the attractive species (
H. melapterus
). Thus, cleaner presence indirectly reduced fish exposure to parasites in a species that is highly susceptible to parasites, but not in one that is rarely infested with parasites. This suggests that cleaner presence indirectly reduces exposure of a common fish species to harmful parasites, which may explain some observed benefits in fishes at this location.
Joint group membership is of major importance for cooperation in humans, and close ties or familiarity with a partner are also thought to promote cooperation in other animals. Here, we present the ...opposite pattern: female cleaner fish, Labroides dimidiatus, behave more cooperatively (by feeding more against their preference) when paired with an unfamiliar male rather than with their social partner. We propose that cooperation based on asymmetric punishment causes this reversed pattern. Males are larger than and dominant to female partners and are more aggressive to unfamiliar than to familiar female partners. In response, females behave more cooperatively with unfamiliar male partners. Our data suggest that in asymmetric interactions, weaker players might behave more cooperatively with out-group members than with in-group members to avoid harsher punishment.
Woodpecker finches are famous for their spontaneous tool use behaviour in the wild. They use twigs or cactus spines to pry arthropods out of crevices and use this ability more than any other ...tool-using species known. We experimentally investigated the cognitive abilities related to tool use. We chose three experimental designs that have been used to test several primate species (trap tube task and modification task) and New Caledonian crows (tool length task). One of six woodpecker finches was able to solve the trap tube task, and several individuals modified tools and chose twigs of appropriate length. Most subjects mastered these new tasks quickly, but we found no evidence that they were able to assess the problems in advance. These findings resemble those obtained for primates in these tasks.
Intraspecific cooperation and interspecific mutualism often feature a marked asymmetry in the scope for exploitation. Cooperation may nevertheless persist despite one-sided opportunities for ...cheating, provided that the partner vulnerable to exploitation has sufficient control over the duration of interaction. The effectiveness of the threat of terminating an encounter, however, depends upon the ease with which both the potential victim and the potential exploiter can find replacement partners. Here, we extend a simple, game-theoretical model of this form of partner control to incorporate variation in the relative abundance of potential victims and exploiters, which leads to variation in the time required for individuals of each type to find a new partner. We show that such market effects have a dramatic influence on the stable level of exploitation (and consequent duration of interaction). As the relative abundance of victims decreases, they become less tolerant to exploitation, terminating encounters earlier (for a given level of exploitation), whereas exploiters behave in a more cooperative manner. As a result, the stable duration of interaction actually increases, despite the decreasing tolerance of the victims. Below a critical level of relative victim abundance, the model suggests that the cost of finding a replacement partner becomes so great that it does not pay to exploit at all.
Animals choose a course of action countless times each day. To do so, they need to prioritise their behaviour within a set of alternative actions and decide which of these actions to perform at any ...one time and for how long, that is, determine when the behaviour has reached its desired effect. This process has classically been called the proximate behavioural control mechanism. Several aspects contribute to this process: internal and external stimuli, the emotions that they elicit, motivation (wants), behavioural goals, valuation, decision‐making and its modulation by mood, and the assessment of behavioural outcomes (liking). I will address all these aspects in the form of an integrated conceptual model. In the process of behavioural control, options need to be valued, and I will refer to evidence showing that an affective hedonic process in respect to (future) reward and punishment heavily affects this value. Moreover, I view motivation, the force that finally drives a specific behavioural output, as being primarily influenced by affective states or even corresponding fully to them. Given the feedback in behavioural control by (dis‐)liking outcomes of behaviour, I reason that in respect to welfare it is more important for animals to reach proximate goals than to avoid negative stimuli. All behavioural choices are modulated, and I show how mood, a long‐term affective state, can cause such modulation. Proximate control of behaviour takes place in the brain, and I will briefly discuss how current and future brain research may elucidate how the brain computes these processes. Furthermore, the inclusion of affective states in the conceptual model raises the question of the subjective experience of animals, and I will address some of the important open questions in this area of research. I will conclude that neural studies cannot currently provide a detailed and general theory on the algorithms of proximate behavioural control. In parallel to further developing these approaches, I propose to strengthen a refined ethological approach with a focus on the states of “wanting” and “liking” in ecologically meaningful circumstances and with a strong ontogenetic (within species) and comparative (between species) component. I consider this ethological approach to be a highly promising step in understanding proximate behavioural control.
The cleaner wrasse Labroides dimidiatus often touches 'client' reef fish dorsal fin areas with its pelvic and pectoral fins. The relative spatial positions of cleaner and client remain constant and ...the cleaner's head points away from the client's body. Therefore, this behaviour is not compatible with foraging and the removal of client ectoparasites. As clients seek such 'tactile stimulation', it can be classified as an interspecific socio-positive behaviour. Our field observations on 12 cleaners (observation time of 112h) suggest that cleaners use tactile stimulation in order to successfully (i) alter client decisions over how long to stay for an inspection, and (ii) stop clients from fleeing or aggressive chasing of the cleaner in response to a cleaner fish bite that made them jolt. Finally, predatory clients receive tactile stimulation more often than non-predatory clients, which might be interpreted as an extra service that cleaners give to specific partners as pre-conflict management, as these partners would be particularly dangerous if they started a conflict. We therefore propose that cleaner fish use interspecific social strategies, which have so far been reported only from mammals, particularly primates.
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