Cooperation prevails when individuals adjust their social ties

Conventional evolutionary game theory predicts that natural selection favours the selfish and strong even though cooperative interactions thrive at all levels of organization in living systems. Recent investigations demonstrated that a limiting factor for the evolution of cooperative interactions is the way in which they are organized, cooperators becoming evolutionarily competitive whenever individuals are constrained to interact with few others along the edges of networks with low average connectivity. Despite this insight, the conundrum of cooperation remains since recent empirical data shows that real networks exhibit typically high average connectivity and associated single-to-broad–scale heterogeneity. Here, a computational model is constructed in which individuals are able to self-organize both their strategy and their social ties throughout evolution, based exclusively on their self-interest. We show that the entangled evolution of individual strategy and network structure constitutes a key mechanism for the sustainability of cooperation in social networks. For a given average connectivity of the population, there is a critical value for the ratio W between the time scales associated with the evolution of strategy and of structure above which cooperators wipe out defectors. Moreover, the emerging social networks exhibit an overall heterogeneity that accounts very well for the diversity of patterns recently found in acquired data on social networks. Finally, heterogeneity is found to become maximal when W reaches its critical value. These results show that simple topological dynamics reflecting the individual capacity for self-organization of social ties can produce realistic networks of high average connectivity with associated single-to-broad–scale heterogeneity. On the other hand, they show that cooperation cannot evolve as a result of “social viscosity” alone in heterogeneous networks with high average connectivity, requiring the additional mechanism of topological co-evolution to ensure the survival of cooperative behaviour.     

Strength of social tie predicts cooperative investment in a human social network

Social networks--diagrams which reflect the social structure of animal groups--are increasingly viewed as useful tools in behavioural ecology and evolutionary biology. Network structure may be especially relevant to the study of cooperation, because the action of mechanisms which affect the cost:benefit ratio of cooperating (e.g. reciprocity, punishment, image scoring) is likely to be mediated by the relative position of actor and recipient in the network. Social proximity could thus affect cooperation in a similar manner to biological relatedness. To test this hypothesis, we recruited members of a real-world social group and used a questionnaire to reveal their network. Participants were asked to endure physical discomfort in order to earn money for themselves and other group members, allowing us to explore relationships between willingness to suffer a cost on another's behalf and the relative social position of donor and recipient. Cost endured was positively correlated with the strength of the social tie between donor and recipient. Further, donors suffered greater costs when a relationship was reciprocated. Interestingly, participants regularly suffered greater discomfort for very close peers than for themselves. Our results provide new insight into the effect of social structure on the direct benefits of cooperation.     

Interactions among social monitoring,anti-predator vigilance and group size in eastern grey kangaroos

Group size is known to affect both the amount of time that prey animals spend in vigilance and the degree to which the vigilance of group members is synchronized. However, the variation in group-size effects reported in the literature is not yet understood. Prey animals exhibit vigilance both to protect themselves against predators and to monitor other group members, and both forms of vigilance presumably influence group-size effects on vigilance. However, our understanding of the patterns of individual investment underlying the time sharing between anti-predator and social vigilance is still limited. We studied patterns of variation in individual vigilance and the synchronization of vigilance with group size in a wild population of eastern grey kangaroos (Macropus giganteus) subject to predation, in particular focusing on peripheral females because we expected that they would exhibit both social and anti-predator vigilance. There was no global effect of group size on individual vigilance. The lack of group-size effect was the result of two compensating effects. The proportion of time individuals spent looking at other group members increased, whereas the proportion of time they spent scanning the environment decreased with group size; as a result, overall vigilance levels did not change with group size. Moreover, a degree of synchrony of vigilance occurred within groups and that degree increased with the proportion of vigilance time peripheral females spent in anti-predator vigilance. Our results highlight the crucial roles of both social and anti-predator components of vigilance in the understanding of the relationship between group size and vigilance, as well as in the synchronization of vigilance among group members.     

Network Diversity and Affect Dynamics: The Role of Personality Traits

People divide their time unequally among their social contacts due to time constraints and varying strength of relationships. It was found that high diversity of social communication, dividing time more evenly among social contacts, is correlated with economic well-being both at macro and micro levels. Besides economic well-being, it is not clear how the diversity of social communication is also associated with the two components of individuals’ subjective well-being, positive and negative affect. Specifically, positive affect and negative affect are two independent dimensions representing the experience (feeling) of emotions. In this paper, we investigate the relationship between the daily diversity of social communication and dynamic affect states that people experience in their daily lives. We collected two high-resolution datasets that capture affect scores via daily experience sampling surveys and social interaction through wearable sensing technologies: sociometric badges for face-to-face interaction and smart phones for mobile phone calls. We found that communication diversity correlates with desirable affect states–e.g. an increase in the positive affect state or a decrease in the negative affect state–for some personality types, but correlates with undesirable affect states for others. For example, diversity in phone calls is experienced as good by introverts, but bad by extroverts; diversity in face-to-face interaction is experienced as good by people who tend to be positive by nature (trait) but bad for people who tend to be not positive by nature. More broadly, the moderating effect of personality type on the relationship between diversity and affect was detected without any knowledge of the type of social tie or the content of communication. This provides further support for the power of unobtrusive sensing in understanding social dynamics, and in measuring the effect of potential interventions designed to improve well-being.     

Do marmosets care to share? Oxytocin treatment reduces prosocial behavior toward strangers

Cooperatively-breeding and socially-monogamous primates, like marmosets and humans, exhibit high levels of social tolerance and prosociality toward others. Oxytocin (OXT) generally facilitates prosocial behavior, but there is growing recognition that OXT modulation of prosocial behavior is shaped by the context of social interactions and by other motivational states such as arousal or anxiety. To determine whether prosociality varies based on social context, we evaluated whether marmoset donors (Callithrix penicillata) preferentially rewarded pairmates versus opposite-sex strangers in a prosocial food-sharing task. To examine potential links among OXT, stress systems, and prosociality, we evaluated whether pretrial cortisol levels in marmosets altered the impact of OXT on prosocial responses. Marmosets exhibited spontaneous prosociality toward others, but they did so preferentially toward strangers compared to their pairmates. When donor marmosets were treated with marmoset-specific Pro⁸-OXT, they exhibited reduced prosociality toward strangers compared to marmosets treated with saline or consensus-mammalian Leu⁸-OXT. When pretrial cortisol levels were lower, marmosets exhibited higher prosociality toward strangers. These findings demonstrate that while marmosets show spontaneous prosocial responses toward others, they do so preferentially toward opposite-sex strangers. Cooperative breeding may be associated with the expression of prosociality, but the existence of a pair-bond between marmoset partners appears to be neither necessary nor sufficient for the expression of spontaneous prosocial responses. Furthermore, high prosociality toward strangers is significantly reduced in marmosets treated with Pro⁸-OXT, suggesting that OXT does not universally enhance prosociality, but, rather OXT modulation of prosocial behavior varies depending on social context.     

Levels of spending and resource allocation to HIV programs and services in Latin America and the Caribbean

Background

An estimated 1.86 million people are living with HIV in Latin America and the Caribbean (LAC). The region is comprised of mainly middle-income countries with steady economic growth while simultaneously there are enormous social inequalities and several concentrated AIDS epidemics. This paper describes HIV spending patterns in LAC countries including analysis of the levels and patterns of domestic HIV spending from both public and international sources.

Methods and Findings

We conducted an extensive analysis of the most recently available data from LAC countries using the National AIDS Spending Assessment tool. The LAC countries spent a total of US$ 1.59 billion on HIV programs and services during the latest reported year. Countries providing detailed information on spending showed that high percentages are allocated to treatment and care (75.1%) and prevention (15.0%). Domestic sources accounted for 93.6 percent of overall spending and 79 percent of domestic funds were directed to treatment and care. International funds represented 5.4 percent of total HIV funding in the region, but they supplied the majority of the effort to reach most-at-risk-populations (MARPs). However, prevalence rates among men who have sex with men (MSM) still reached over 25 percent in some countries.

Conclusions

Although countries in the region have increasingly sustained their response from domestic sources, still there are future challenges: 1) The growing number of new HIV infections and more people-living-with-HIV (PLWH) eligible to receive antiretroviral treatment (ART); 2) Increasing ART coverage along with high prices of antiretroviral drugs; and 3) The funding for prevention activities among MARPs rely almost exclusively on external donors. These threats call for strengthened actions by civil society and governments to protect and advance gains against HIV in LAC.     

Influences of State and Trait Affect on Behavior,Feedback-Related Negativity,and P3b in the Ultimatum Game

The present study investigates how different emotions can alter social bargaining behavior. An important paradigm to study social bargaining is the Ultimatum Game. There, a proposer gets a pot of money and has to offer part of it to a responder. If the responder accepts, both players get the money as proposed by the proposer. If he rejects, none of the players gets anything. Rational choice models would predict that responders accept all offers above 0. However, evidence shows that responders typically reject a large proportion of all unfair offers. We analyzed participants’ behavior when they played the Ultimatum Game as responders and simultaneously collected electroencephalogram data in order to quantify the feedback-related negativity and P3b components. We induced state affect (momentarily emotions unrelated to the task) via short movie clips and measured trait affect (longer-lasting emotional dispositions) via questionnaires. State happiness led to increased acceptance rates of very unfair offers. Regarding neurophysiology, we found that unfair offers elicited larger feedback-related negativity amplitudes than fair offers. Additionally, an interaction of state and trait affect occurred: high trait negative affect (subsuming a variety of aversive mood states) led to increased feedback-related negativity amplitudes when participants were in an angry mood, but not if they currently experienced fear or happiness. We discuss that increased rumination might be responsible for this result, which might not occur, however, when people experience happiness or fear. Apart from that, we found that fair offers elicited larger P3b components than unfair offers, which might reflect increased pleasure in response to fair offers. Moreover, high trait negative affect was associated with decreased P3b amplitudes, potentially reflecting decreased motivation to engage in activities. We discuss implications of our results in the light of theories and research on depression and anxiety.     

Brain serotonin deficiency leads to social communication deficits in mice

A deficit in brain serotonin is thought to be associated with deteriorated stress coping behaviour, affective disorders and exaggerated violence. We challenged this hypothesis in mice with a brain-specific serotonin depletion caused by a tryptophan hydroxylase 2 (TPH2) deficiency. We tested TPH2-deficient (Tph2^−/–) animals in two social situations. As juveniles, Tph2^−/− mice displayed reduced social contacts, whereas ultrasonic vocalizations (USVs) were unchanged within same-sex same-genotype pairings. Interestingly, juvenile females vocalized more than males across genotypes. Sexually naive adult males were exposed to fresh male or female urine, followed by an interaction with a conspecific, and re-exposed to urine. Although Tph2^−/− mice showed normal sexual preference, they were hyper-aggressive towards their interaction partners and did not vocalize in response to sexual cues. These results highlight that central serotonin is essential for prosocial behaviour, especially USV production in adulthood, but not for sexual preference.     

Field and laboratory studies of social organization in Bushbabies (Galago senegalensis)

During the course of a one year field study of the Lesser Bushbaby (Galago senegalensis moholi), using a technique of direct observation with red light, it was discovered that family groups which slept together during the daytime split up at night, with each member spending most of the time alone. During the night individuals would come into contact with one another or with members of adjacent groups and the resulting social behaviour varied considerably. An interpretation has been made of more than 120 interactions observed in the field based on the behaviour of bushbabies introduced to one another under controlled semi-natural conditions.     

Effects of deception in social networks

Honesty plays a crucial role in any situation where organisms exchange information or resources. Dishonesty can thus be expected to have damaging effects on social coherence if agents cannot trust the information or goods they receive. However, a distinction is often drawn between prosocial lies (‘white’ lies) and antisocial lying (i.e. deception for personal gain), with the former being considered much less destructive than the latter. We use an agent-based model to show that antisocial lying causes social networks to become increasingly fragmented. Antisocial dishonesty thus places strong constraints on the size and cohesion of social communities, providing a major hurdle that organisms have to overcome (e.g. by evolving counter-deception strategies) in order to evolve large, socially cohesive communities. In contrast, white lies can prove to be beneficial in smoothing the flow of interactions and facilitating a larger, more integrated network. Our results demonstrate that these group-level effects can arise as emergent properties of interactions at the dyadic level. The balance between prosocial and antisocial lies may set constraints on the structure of social networks, and hence the shape of society as a whole.     

Vigilance and social organization in two species of primates

Animals that live in groups may accrue anti-predator benefits by virtue of their association with other individuals. While a number of factors (e.g. habitat quality, group geometry, dominance rank) have been shown to affect the relationship between group living and individual rates of visual vigilance, sociality and vigilance have not been studied specifically in terms of the two competing demands they impose upon an individual's visual time. Two captive groups each of tamarins, Saguinus labiatus, and squirrel monkeys, Saimiri sciureus, were used in three related experiments to determine whether they looked at the social or non-social environment while they engated in a simulated foraging task. The tamarins, whose social behaviour is quite pacific and cooperative relative to the more hierarchical squirrel monkeys, looked at the non-social environment when they visually interrupted their foraging. Squirrel monkeys, however, looked more often at group mates. Detection of predators is probably more likely when an individual directs its attention not to conspecifics, but to the environment. Hence, social organizations in which individuals must pay social attention in order to monitor threats and avoid aggression may reduce individual rates of vigilance for predators. Failure to consider various targets of attention may well overestimate vigilance for predators by including other sorts of attentional phenomena in the measurement of vigilance.     

Alcohol Use among Adolescent Youth: The Role of Friendship Networks and Family Factors in Multiple School Studies

To explore the co-evolution of friendship tie choice and alcohol use behavior among 1,284 adolescents from 12 small schools and 976 adolescents from one big school sampled in the National Longitudinal Study of Adolescent to Adult Health (AddHealth), we apply a Stochastic Actor-Based (SAB) approach implemented in the R-based Simulation Investigation for Empirical Network Analysis (RSiena) package. Our results indicate the salience of both peer selection and peer influence effects for friendship tie choice and adolescent drinking behavior. Concurrently, the main effect models indicate that parental monitoring and the parental home drinking environment affected adolescent alcohol use in the small school sample, and that parental home drinking environment affected adolescent drinking in the large school sample. In the small school sample, we detect an interaction between the parental home drinking environment and choosing friends that drink as they multiplicatively affect friendship tie choice. Our findings suggest that future research should investigate the synergistic effects of both peer and parental influences for adolescent friendship tie choices and drinking behavior. And given the tendency of adolescents to form ties with their friends'' friends, and the evidence of local hierarchy in these networks, popular youth who do not drink may be uniquely positioned and uniquely salient as the highest rank of the hierarchy to cause anti-drinking peer influences to diffuse down the social hierarchy to less popular youth. As such, future interventions should harness prosocial peer influences simultaneously with strategies to increase parental support and monitoring among parents to promote affiliation with prosocial peers.     

Effects of reproductive status, social rank, sex and group size on vigilance patterns in Przewalski's gazelle

Background

Quantifying vigilance and exploring the underlying mechanisms has been the subject of numerous studies. Less attention has focused on the complex interplay between contributing factors such as reproductive status, social rank, sex and group size. Reproductive status and social rank are of particular interest due to their association with mating behavior. Mating activities in rutting season may interfere with typical patterns of vigilance and possibly interact with social rank. In addition, balancing the tradeoff between vigilance and life maintenance may represent a challenge for gregarious ungulate species rutting under harsh winter conditions. We studied vigilance patterns in the endangered Przewalski''s gazelle (Procapra przewalskii) during both the rutting and non-rutting seasons to examine these issues.

Methodology/Principal Findings

Field observations were carried out with focal sampling during rutting and non-rutting season in 2008–2009. Results indicated a complex interplay between reproductive status, social rank, sex and group size in determining vigilance in this species. Vigilance decreased with group size in female but not in male gazelles. Males scanned more frequently and thus spent more time vigilant than females. Compared to non-rutting season, gazelles increased time spent scanning at the expense of bedding in rutting season. During the rutting season, territorial males spent a large proportion of time on rutting activities and were less vigilant than non-territorial males. Although territorial males may share collective risk detection with harem females, we suggest that they are probably more vulnerable to predation because they seemed reluctant to leave rut stands under threats.

Conclusions/Significance

Vigilance behavior in Przewalski''s gazelle was significantly affected by reproductive status, social rank, sex, group size and their complex interactions. These findings shed light on the mechanisms underlying vigilance patterns and the tradeoff between vigilance and other crucial activities.     

Weather Effects on Mobile Social Interactions: A Case Study of Mobile Phone Users in Lisbon,Portugal

The effect of weather on social interactions has been explored through the analysis of a large mobile phone use dataset. Time spent on phone calls, numbers of connected social ties, and tie strength were used as proxies for social interactions; while weather conditions were characterized in terms of temperature, relative humidity, air pressure, and wind speed. Our results are based on the analysis of a full calendar year of data for 22,696 mobile phone users (53.2 million call logs) in Lisbon, Portugal. The results suggest that different weather parameters have correlations to the level and character of social interactions. We found that although weather did not show much influence upon people''s average call duration, the likelihood of longer calls was found to increase during periods of colder weather. During periods of weather that were generally considered to be uncomfortable (i.e., very cold/warm, very low/high air pressure, and windy), people were found to be more likely to communicate with fewer social ties. Despite this tendency, we found that people are more likely to maintain their connections with those they have strong ties with much more than those of weak ties. This study sheds new light on the influence of weather conditions on social relationships and how mobile phone data can be used to investigate the influence of environmental factors on social dynamics.     

Estrogen Receptor-α in the Bed Nucleus of the Stria Terminalis Regulates Social Affiliation in Male Prairie Voles (Microtus ochrogaster)

Estrogen receptor alpha (ERα) typically masculinizes male behavior, while low levels of ERα in the medial amygdala (MeA) and the bed nucleus of the stria terminalis (BST) are associated with high levels of male prosocial behavior. In the males of the highly social prairie vole (Microtus ochrogaster), increasing ERα in the MeA inhibited the expression of spontaneous alloparental behavior and produced a preference for novel females. To test for the effects of increased ERα in the BST, a viral vector was used to enhance ERα expression in the BST of adult male prairie voles. Following treatment, adult males were tested for alloparental behavior with 1–3-day-old pups, and for heterosexual social preference and affiliation. Treatment did not affect alloparental behavior as 73% of ERα-BST males and 62.5% of control males were alloparental. Increasing ERα in the BST affected heterosexual affiliation, with ERα-BST males spending significantly less total time in side-by-side contact with females relative to time spent with control males. ERα-BST males did not show a preference for either the familiar or novel female. These findings differed significantly from those reported in ERα-MeA enhanced males, where ERα inhibited alloparental behavior and produced a preference for a novel female. The findings from this study suggest two things: first, that increased ERα in the BST decreases social affiliation and second, that altering ERα in different regions of the social neural circuit differentially impacts the expression of social behavior.     

Policing in Nonhuman Primates: Partial Interventions Serve a Prosocial Conflict Management Function in Rhesus Macaques

Studies of prosocial policing in nonhuman societies traditionally focus on impartial interventions because of an underlying assumption that partial support implies a direct benefit to the intervener, thereby negating the potential for being prosocial in maintaining social stability for the benefit of the group. However, certain types of partial interventions have significant potential to be prosocial in controlling conflict, e.g. support of non-kin subordinates. Here, we propose a policing support hypothesis that some types of agonistic support serve a prosocial policing function that maintains group stability. Using seven large captive groups of rhesus macaques, we investigated the relationship between intervention type and group-level costs and benefits (rates of trauma, severe aggression, social relocation) and individual level costs and benefits (preferential sex-dyad targeting, dominance ambiguity reduction, access to mates, and return aggression). Our results show that impartial interventions and support of subordinate non-kin represent prosocial policing as both (1) were negatively associated with group-level rates of trauma and severe aggression, respectively, (2) showed no potential to confer individual dominance benefits, (3) when performed outside the mating season, they did not increase chances of mating with the beneficiary, and (4) were low-cost for the highest-ranking interveners. We recommend expanding the definition of ‘policing’ in nonhumans to include these ‘policing support interventions’.     

Stronger social bonds do not always predict greater longevity in a gregarious primate

In group‐living species, individuals often have preferred affiliative social partners, with whom ties or bonds can confer advantages that correspond with greater fitness. For example, in adult female baboons and juvenile horses, individuals with stronger or more social ties experience greater survival. We used detailed behavioral and life history records to explore the relationship between tie quality and survival in a gregarious monkey (Cercopithecus mitis stuhlmanni), while controlling for dominance rank, group size, and life history strategy. We used Cox proportional hazards regressions to model the cumulative (multi‐year) and current (single‐year) relationships of social ties and the hazard of mortality in 83 wild adult females of known age, observed 2–8 years each (437 subject‐years) in eight social groups. The strength of bonds with close partners was associated with increased mortality risk under certain conditions: Females that had strong bonds with close partners that were inconsistent over multiple years had a higher risk of mortality than females adopting any other social strategy. Within a given year, females had a higher risk of death if they were strongly bonded with partners that changed from the previous year versus with partners that remained consistent. Dominance rank, number of adult female groupmates, and age at first reproduction did not predict the risk of death. This study demonstrates that costs and benefits of strong social bonds can be context‐dependent, relating to the consistency of social partners over time.     

Mixing at the beginning of fattening moderates social buffering in beef bulls

Cattle establish strong preferential associations early in their life that determine the cohesiveness within a group and can diminish fear responses (i.e. social buffering). Mixing beef bulls at the beginning of fattening may lead to less cohesiveness. Moreover, mixing beef bulls induces aggressive interactions, especially when the bulls are of similar weights. This study was aimed at evaluating the cohesiveness and fear responses of 64 bulls that were either mixed versus unmixed and of homogeneous vs. heterogeneous body weights (2 × 2 factorial design, four groups of four bulls each per treatment). Social cohesiveness was assessed during food competition and a social separation test. Fear responses were observed during the social separation test and pre-slaughter handling. During food competition, unmixed bulls exchanged more non-agonistic interactions (P < 0.05) and were more tolerant with each other than mixed bulls (time spent eating: P < 0.05). Unmixed bulls displayed less fear responses during physical separation (elimination: P < 0.05; startle: P < 0.01) and were less stressed by pre-slaughter handling (blood cortisol: P < 0.05). Very few effects could be attributed to the homogeneity of weights within groups. It is concluded that mixing bulls at the beginning of fattening reduces the strength of positive social relationships and renders social buffering less effective during subsequent fattening. We recommend maintaining familiar bulls together throughout fattening in order to preserve cohesiveness within groups and for animals to benefit from social buffering during stressful situations.     

Using a Rebound Matrix to Estimate Consumption Changes from Saving and its Environmental Impact in Japan

We investigate the extent to which Japanese people can change their consumption and the corresponding environmental impact. We propose a new analytical framework with a rebound matrix that captures the monetary flow from potential savings to their respending (referred to as rebound). A questionnaire is used to derive the matrix. On average, respondents spent 3.4 million Yen annually, resulting in 12.4 tons of carbon dioxide (CO₂) emissions in their daily lives. The survey results suggest that acceptable spending reductions would correspond to a CO₂ emissions reduction of nearly 6%. However, the CO₂ emissions would increase by nearly the same amount when the respondents respend their savable money (rebound CO₂ emissions). The annual CO₂ emissions and the annually reducible CO₂ emissions both increase with the increase in annual expenditure. Consequently, the net CO₂ emissions also increase with the increase in annual expenditure. The rebound spending is approximated using the rebound matrix. Finally, it is suggested that the net CO₂ emissions can be reduced through lifestyle changes whereby spending on energy items is reduced and the resulting savings are spent on telecommunication, clothes, shoes, education, and housing.     

Thermal constraints on daily patterns of aggregation and density along an intertidal gradient in the periwinkle Echinolittorina peruviana

Variations in environmental temperature have both direct and indirect effects that affect organisms at levels ranging from intra-cellular physiological processes to ecological patterns. These variations are especially important for intertidal marine ectotherms such as littorinids since they alternate between periods of immersion in seawater, and must also experience long periods of emersion. In central Chile, Echinolittorina peruviana is one of the most conspicuous species on rocky intertidal shores, occurring at high tidal levels and in the splash zone. The species is known to resist direct exposure to the sun for long periods, although juveniles tend to be restricted to protected microhabitats. Adults show seasonal variations in abundance and vertical distribution and may form aggregations that have been shown to help reduce water loss and body temperature. In this study, we evaluate the relationship between daily thermal variations throughout the vertical distribution of this species and how these affect the patterns of density and aggregation. Our results suggest that one of the leading determinants of the spatio-temporal variation of density in E. peruviana may be operative temperature (T_O: the amount of stored heat resulting from the balance between heat fluxes into and out of the body, measured with taxidermic mounts mimicking heat transfer properties of the snail). T_O showed a strong negative relationship with density and a strong positive relationship with aggregation in the highest intertidal level monitored. The strength of these relationships decreased in importance at lower levels. While T_O alone cannot explain the abundance of E. peruviana throughout its range of distribution, our results show that it does have a strong influence that should be considered in addition to other ecological factors affecting the density of intertidal littorinids.