COVID-19 inequalities in England: a mathematical modelling study of transmission risk and clinical vulnerability by socioeconomic status.

BACKGROUND: The COVID-19 pandemic resulted in major inequalities in infection and disease burden between areas of varying socioeconomic deprivation in many countries, including England. Areas of higher deprivation tend to have a different population structure-generally younger-which can increase viral transmission due to higher contact rates in school-going children and working-age adults. Higher deprivation is also associated with a higher presence of chronic comorbidities, which were convincingly demonstrated to be risk factors for severe COVID-19 disease. These two major factors need to be combined to better understand and quantify their relative importance in the observed COVID-19 inequalities. METHODS: We used UK Census data on health status and demography stratified by decile of the Index of Multiple Deprivation (IMD), which is a measure of socioeconomic deprivation. We calculated epidemiological impact using an age-stratified COVID-19 transmission model, which incorporated different contact patterns and clinical health profiles by decile. To separate the contribution of each factor, we considered a scenario where the clinical health profile of all deciles was at the level of the least deprived. We also considered the effectiveness of school closures and vaccination of over 65-year-olds in each decile. RESULTS: In the modelled epidemics in urban areas, the most deprived decile experienced 9% more infections, 13% more clinical cases, and a 97% larger peak clinical size than the least deprived; we found similar inequalities in rural areas. Twenty-one per cent of clinical cases and 16% of deaths in England observed under the model assumptions would not occur if all deciles experienced the clinical health profile of the least deprived decile. We found that more deaths were prevented in more affluent areas during school closures and vaccination rollouts. CONCLUSIONS: This study demonstrates that both clinical and demographic factors synergise to generate health inequalities in COVID-19, that improving the clinical health profile of populations would increase health equity, and that some interventions can increase health inequalities.

Chimpanzees use social information to acquire a skill they fail to innovate.

Cumulative cultural evolution has been claimed to be a uniquely human phenomenon pivotal to the biological success of our species. One plausible condition for cumulative cultural evolution to emerge is individuals' ability to use social learning to acquire know-how that they cannot easily innovate by themselves. It has been suggested that chimpanzees may be capable of such know-how social learning, but this assertion remains largely untested. Here we show that chimpanzees use social learning to acquire a skill that they failed to independently innovate. By teaching chimpanzees how to solve a sequential task (one chimpanzee in each of the two tested groups, n = 66) and using network-based diffusion analysis, we found that 14 naive chimpanzees learned to operate a puzzle box that they failed to operate during the preceding three months of exposure to all necessary materials. In conjunction, we present evidence for the hypothesis that social learning in chimpanzees is necessary and sufficient to acquire a new, complex skill after the initial innovation.

Protecting infants against RSV disease: an impact and cost-effectiveness comparison of long-acting monoclonal antibodies and maternal vaccination.

Background: Two new products for preventing Respiratory Syncytial Virus (RSV) in young children have been licensed: a single-dose long-acting monoclonal antibody (la-mAB) and a maternal vaccine (MV). To facilitate the selection of new RSV intervention programmes for large-scale implementation, this study provides an assessment to compare the costs of potential programmes with the health benefits accrued. Methods: Using an existing dynamic transmission model, we compared maternal vaccination to la-mAB therapy against RSV in England and Wales by calculating the impact and cost-effectiveness. We calibrated a statistical model to the efficacy trial data to accurately capture their immune waning and estimated the impact of seasonal and year-round programmes for la-mAB and MV programmes. Using these impact estimates, we identified the most cost-effective programme across pricing and delivery cost assumptions. Findings: For infants under six months old in England and Wales, a year-round MV programme with 60% coverage would avert 32% (95% CrI 22-41%) of RSV hospital admissions and a year-round la-mAB programme with 90% coverage would avert 57% (95% CrI 41-69%). The MV programme has additional health benefits for pregnant women, which account for 20% of the population-level health burden averted. A seasonal la-mAB programme could be cost-effective for up to £84 for purchasing and administration (CCPA) and a seasonal MV could be cost-effective for up to £80 CCPA. Interpretation: This modelling and cost-effectiveness analysis has shown that both the long-acting monoclonal antibodies and the maternal vaccine could substantially reduce the burden of RSV disease in the infant population. Our analysis has informed JCVI's recommendations for an RSV immunisation programme to protect newborns and infants. Funding: National Institute for Health Research.

Group-specific expressions of co-feeding tolerance in bonobos and chimpanzees preclude dichotomous species generalizations.

Bonobos are typically portrayed as more socially tolerant than chimpanzees, yet the current evidence supporting such a species-level categorization is equivocal. Here, we used validated group-level co-feeding assays to systematically test expressions of social tolerance in sixteen groups of zoo- and sanctuary-housed bonobos and chimpanzees. We found that co-feeding tolerance substantially overlaps between the species, thus precluding categorical inference at the species level. Instead, marked differences were observed between groups, with some bonobo communities exhibiting higher social tolerance than chimpanzee communities, and vice versa. Moreover, considerable intergroup variation was found within species living in the same environment, which attests to Pan's behavioral flexibility. Lastly, chimpanzees showed more tolerance in male-skewed communities, whereas bonobos responded less pronounced to sex-ratio variation. We conclude that the pervasive dichotomy between the tolerant bonobo and the belligerent chimpanzee requires quantitative nuance, and that accurate phylogenetic tracing of (human) social behavior warrants estimations of intraspecific group variation.

The potential cost-effectiveness of next generation influenza vaccines in England and Wales: A modelling analysis.

Next generation influenza vaccines are in development and have the potential for widespread health and economic benefits. Determining the potential health and economic impact for these vaccines is needed to drive investment in bringing these vaccines to the market, and to inform which groups public health policies on influenza vaccination should target. We used a mathematical modelling approach to estimate the epidemiological impact and cost-effectiveness of next generation influenza vaccines in England and Wales. We used data from an existing fitted model, and evaluated new vaccines with different characteristics ranging from improved vaccines with increased efficacy duration and breadth of protection, to universal vaccines, defined in line with the World Health Organisation (WHO) Preferred Product Characteristics (PPC). We calculated the cost effectiveness of new vaccines in comparison to the current seasonal vaccination programme. We calculated and compared the Incremental Cost-Effectiveness Ratio and Incremental Net Monetary Benefit for each new vaccine type. All analysis was conducted in R. We show that next generation influenza vaccines may result in a 21% to 77% reduction in influenza infections, dependent on vaccine characteristics. Our economic modelling shows that using any of these next generation vaccines at 2019 coverage levels would be highly cost-effective at a willingness to pay threshold of £20,000 for a range of vaccine prices. The vaccine threshold price for the best next generation vaccines in £-2019 is £230 (95%CrI £192 - £269) per dose, but even minimally-improved influenza vaccines could be priced at £18 (95%CrI £16 - £21) per dose and still remain cost-effective. This evaluation demonstrates the promise of next generation influenza vaccines for impact on influenza epidemics, and likely cost-effectiveness profiles. We have provided evidence towards a full value of vaccines assessment which bolsters the investment case for development and roll-out of next-generation influenza vaccines.

Scoring epidemiological forecasts on transformed scales.

Forecast evaluation is essential for the development of predictive epidemic models and can inform their use for public health decision-making. Common scores to evaluate epidemiological forecasts are the Continuous Ranked Probability Score (CRPS) and the Weighted Interval Score (WIS), which can be seen as measures of the absolute distance between the forecast distribution and the observation. However, applying these scores directly to predicted and observed incidence counts may not be the most appropriate due to the exponential nature of epidemic processes and the varying magnitudes of observed values across space and time. In this paper, we argue that transforming counts before applying scores such as the CRPS or WIS can effectively mitigate these difficulties and yield epidemiologically meaningful and easily interpretable results. Using the CRPS on log-transformed values as an example, we list three attractive properties: Firstly, it can be interpreted as a probabilistic version of a relative error. Secondly, it reflects how well models predicted the time-varying epidemic growth rate. And lastly, using arguments on variance-stabilizing transformations, it can be shown that under the assumption of a quadratic mean-variance relationship, the logarithmic transformation leads to expected CRPS values which are independent of the order of magnitude of the predicted quantity. Applying a transformation of log(x + 1) to data and forecasts from the European COVID-19 Forecast Hub, we find that it changes model rankings regardless of stratification by forecast date, location or target types. Situations in which models missed the beginning of upward swings are more strongly emphasised while failing to predict a downturn following a peak is less severely penalised when scoring transformed forecasts as opposed to untransformed ones. We conclude that appropriate transformations, of which the natural logarithm is only one particularly attractive option, should be considered when assessing the performance of different models in the context of infectious disease incidence.

Food sharing with friends and acquaintances: A study in preschool boys and girls.

Introduction: The current study examined whether preschoolers in a (semi-)natural situation shared more food with friends or acquaintances, and whether this was different between boys and girls, older and younger children, and for preferred and non- preferred food. In order to do so, we replicated and extended the classical work of Birch and Billman in a Dutch sample. Methods: Participants included 91 children aged between 3 to 6 years (52.7% boys, 93.4% Western European) from a middle- to upper-middle-class neighborhood in the Netherlands. Results: The results revealed that children shared more non-preferred than preferred food with others. Girls gave more non-preferred food to acquaintances than to friends, whereas boys gave more to friends than to acquaintances. No effect of relationship was found for preferred food. Older children shared more food than younger children. Compared to acquaintances, friends made more active attempts to get food. Moreover, children who were not shared with were just as likely to share food as children who were shared with. Discussion: Overall, only a small degree of agreement with the original study was found: Some significant findings could not be replicated, and some unconfirmed hypotheses of the original study were supported. The results underscore both the need for replications and studying the effect of social-contextual factors in natural settings.

Potential health and economic impact of paediatric vaccination using next-generation influenza vaccines in Kenya: a modelling study.

BACKGROUND: Influenza is a major year-round cause of respiratory illness in Kenya, particularly in children under 5. Current influenza vaccines result in short-term, strain-specific immunity and were found in a previous study not to be cost-effective in Kenya. However, next-generation vaccines are in development that may have a greater impact and cost-effectiveness profile. METHODS: We expanded a model previously used to evaluate the cost-effectiveness of seasonal influenza vaccines in Kenya to include next-generation vaccines by allowing for enhanced vaccine characteristics and multi-annual immunity. We specifically examined vaccinating children under 5 years of age with improved vaccines, evaluating vaccines with combinations of increased vaccine effectiveness, cross-protection between strains (breadth) and duration of immunity. We evaluated cost-effectiveness using incremental cost-effectiveness ratios (ICERs) and incremental net monetary benefits (INMBs) for a range of values for the willingness-to-pay (WTP) per DALY averted. Finally, we estimated threshold per-dose vaccine prices at which vaccination becomes cost-effective. RESULTS: Next-generation vaccines can be cost-effective, dependent on the vaccine characteristics and assumed WTP thresholds. Universal vaccines (assumed to provide long-term and broad immunity) are most cost-effective in Kenya across three of four WTP thresholds evaluated, with the lowest median value of ICER per DALY averted ($263, 95% Credible Interval (CrI): $ - 1698, $1061) and the highest median INMBs. At a WTP of $623, universal vaccines are cost-effective at or below a median price of $5.16 per dose (95% CrI: $0.94, $18.57). We also show that the assumed mechanism underlying infection-derived immunity strongly impacts vaccine outcomes. CONCLUSIONS: This evaluation provides evidence for country-level decision makers about future next-generation vaccine introduction, as well as global research funders about the potential market for these vaccines. Next-generation vaccines may offer a cost-effective intervention to reduce influenza burden in low-income countries with year-round seasonality like Kenya.

Chimpanzee communities differ in their inter- and intrasexual social relationships.

Male and female human social bonding strategies are culturally shaped, in addition to being genetically rooted. Investigating nonhuman primate bonding strategies across sex groups allows researchers to assess whether, as with humans, they are shaped by the social environment or whether they are genetically predisposed. Studies of wild chimpanzees show that in some communities males have strong bonds with other males, whereas in others, females form particularly strong intrasex bonds, potentially indicative of cultural differences across populations. However, excluding genetic or ecological explanations when comparing different wild populations is difficult. Here, we applied social network analysis to examine male and female social bonds in two neighbouring semiwild chimpanzee groups of comparable ecological conditions and subspecies compositions, but that differ in demographic makeup. Results showed differences in bonding strategies across the two groups. While female-female party co-residence patterns were significantly stronger in Group 1 (which had an even distribution of males and females) than in Group 2 (which had a higher proportion of females than males), there were no such differences for male-male or male-female associations. Conversely, female-female grooming bonds were stronger in Group 2 than in Group 1. We also found that, in line with captive studies but contrasting research with wild chimpanzees, maternal kinship strongly predicted proximity and grooming patterns across the groups. Our findings suggest that, as with humans, male and female chimpanzee social bonds are influenced by the specific social group they live in, rather than predisposed sex-based bonding strategies.

Biased cultural transmission of a social custom in chimpanzees.

Cultural transmission studies in animals have predominantly focused on identifying between-group variation in tool-use techniques, while immaterial cultures remain understudied despite their potential for highlighting similarities between human and animal culture. Here, using long-term data from two chimpanzee communities, we tested whether one of chimpanzees' most enigmatic social customs-the grooming handclasp-is culturally transmitted by investigating the influence of well-documented human transmission biases on their variational preferences. After identifying differences in style preferences between the communities, we show that older and dominant individuals exert more influence over their partners' handclasp styles. Mothers were equally likely to influence their offspring's preferences as nonkin, indicating that styles are transmitted both vertically and obliquely. Last, individuals gradually converged on the group style, suggesting that conformity guides chimpanzees' handclasp preferences. Our findings show that chimpanzees' social lives are influenced by cultural transmission biases that hitherto were thought to be uniquely human.

Does tolerance allow bonobos to outperform chimpanzees on a cooperative task? A conceptual replication of Hare et al., 2007.

Across various taxa, social tolerance is thought to facilitate cooperation, and many species are treated as having species-specific patterns of social tolerance. Yet studies that assess wild and captive bonobos and chimpanzees result in contrasting findings. By replicating a cornerstone experimental study on tolerance and cooperation in bonobos and chimpanzees (Hare et al. 2007 Cur. Biol. 17, 619-623 (doi:10.1016/j.cub.2007.02.040)), we aim to further our understanding of current discrepant findings. We tested bonobos and chimpanzees housed at the same facility in a co-feeding and cooperation task. Food was placed on dishes located on both ends or in the middle of a platform. In the co-feeding task, the tray was simply made available to the ape duos, whereas in the cooperation task the apes had to simultaneously pull at both ends of a rope attached to the platform to retrieve the food. By contrast to the published findings, bonobos and chimpanzees co-fed to a similar degree, indicating a similar level of tolerance. However, bonobos cooperated more than chimpanzees when the food was monopolizable, which replicates the original study. Our findings call into question the interpretation that at the species level bonobos cooperate to a higher degree because they are inherently more tolerant.

Chimpanzees communicate to coordinate a cultural practice.

Human culture thrives by virtue of communication, yet whether communication plays an influential role in the cultural lives of other animals remains understudied. Here, we investigated whether chimpanzees use communication to engage in a cultural practice by analysing grooming handclasp (GHC) interactions-a socio-cultural behaviour requiring interindividual coordination for successful execution. Previous accounts attributed GHC initiations to behavioural shaping, whereby the initiator physically moulds the partner's arm into the desired GHC posture. Using frame-by-frame analysis and matched-control methodology, we find that chimpanzees do not only shape their partner's posture (22%), but also use gestural communication to initiate GHC (44%), which requires an active and synchronized response from the partner. Moreover, in a third (34%) of the GHC initiations, the requisite coordination was achieved by seemingly effortless synchrony. Lastly, using a longitudinal approach, we find that for GHC initiations, communication occurs more frequently than shaping in experienced dyads and less in mother-offspring dyads. These findings are consistent with ontogenetic ritualization, thereby reflecting first documentation of chimpanzees communicating to coordinate a cultural practice. We conclude that chimpanzees show interactional flexibility in the socio-cultural domain, opening the possibility that the interplay between communication and culture is rooted in our deep evolutionary history.

Human judgement forecasting of COVID-19 in the UK.

Background: In the past, two studies found ensembles of human judgement forecasts of COVID-19 to show predictive performance comparable to ensembles of computational models, at least when predicting case incidences. We present a follow-up to a study conducted in Germany and Poland and investigate a novel joint approach to combine human judgement and epidemiological modelling. Methods: From May 24th to August 16th 2021, we elicited weekly one to four week ahead forecasts of cases and deaths from COVID-19 in the UK from a crowd of human forecasters. A median ensemble of all forecasts was submitted to the European Forecast Hub. Participants could use two distinct interfaces: in one, forecasters submitted a predictive distribution directly, in the other forecasters instead submitted a forecast of the effective reproduction number R t. This was then used to forecast cases and deaths using simulation methods from the EpiNow2 R package. Forecasts were scored using the weighted interval score on the original forecasts, as well as after applying the natural logarithm to both forecasts and observations. Results: The ensemble of human forecasters overall performed comparably to the official European Forecast Hub ensemble on both cases and deaths, although results were sensitive to changes in details of the evaluation. R t forecasts performed comparably to direct forecasts on cases, but worse on deaths. Self-identified "experts" tended to be better calibrated than "non-experts" for cases, but not for deaths. Conclusions: Human judgement forecasts and computational models can produce forecasts of similar quality for infectious disease such as COVID-19. The results of forecast evaluations can change depending on what metrics are chosen and judgement on what does or doesn't constitute a "good" forecast is dependent on the forecast consumer. Combinations of human and computational forecasts hold potential but present real-world challenges that need to be solved.

Chimpanzees organize their social relationships like humans.

Human relationships are structured in a set of layers, ordered from higher (intimate relationships) to lower (acquaintances) emotional and cognitive intensity. This structure arises from the limits of our cognitive capacity and the different amounts of resources required by different relationships. However, it is unknown whether nonhuman primate species organize their affiliative relationships following the same pattern. We here show that the time chimpanzees devote to grooming other individuals is well described by the same model used for human relationships, supporting the existence of similar social signatures for both humans and chimpanzees. Furthermore, the relationship structure depends on group size as predicted by the model, the proportion of high-intensity connections being larger for smaller groups.

Comparing human and model-based forecasts of COVID-19 in Germany and Poland.

Forecasts based on epidemiological modelling have played an important role in shaping public policy throughout the COVID-19 pandemic. This modelling combines knowledge about infectious disease dynamics with the subjective opinion of the researcher who develops and refines the model and often also adjusts model outputs. Developing a forecast model is difficult, resource- and time-consuming. It is therefore worth asking what modelling is able to add beyond the subjective opinion of the researcher alone. To investigate this, we analysed different real-time forecasts of cases of and deaths from COVID-19 in Germany and Poland over a 1-4 week horizon submitted to the German and Polish Forecast Hub. We compared crowd forecasts elicited from researchers and volunteers, against a) forecasts from two semi-mechanistic models based on common epidemiological assumptions and b) the ensemble of all other models submitted to the Forecast Hub. We found crowd forecasts, despite being overconfident, to outperform all other methods across all forecast horizons when forecasting cases (weighted interval score relative to the Hub ensemble 2 weeks ahead: 0.89). Forecasts based on computational models performed comparably better when predicting deaths (rel. WIS 1.26), suggesting that epidemiological modelling and human judgement can complement each other in important ways.

Self-interest precludes prosocial juice provisioning in a free choice group experiment in bonobos.

Previous studies on prosociality in bonobos have reported contrasting results, which might partly be explained by differences in experimental contexts. In this study, we implement a free choice group experiment in which bonobos can provide fruit juice to their group members at a low cost for themselves. Four out of five bonobos passed a training phase and understood the setup and provisioned fruit juice in a total of 17 dyads. We show that even in this egalitarian group with a shallow hierarchy, the majority of pushing was done by the alpha female, who monopolized the setup and provided most juice to two adult females, her closest social partners. Nonetheless, the bonobos in this study pushed less frequently than the chimpanzees in the original juice-paradigm study, suggesting that bonobos might be less likely than chimpanzees to provide benefits to group members. Moreover, in half of the pushing acts, subjects obtained juice for themselves, suggesting that juice provisioning was partly driven by self-regarding behavior. Our study indicates that a more nuanced view on the prosocial food provisioning nature of bonobos is warranted but based on this case study, we suggest that the observed sex differences in providing food to friends corresponds with the socio-ecological sex difference in cooperative interactions in wild and zoo-housed bonobos.

Determinants of RSV epidemiology following suppression through pandemic contact restrictions.

INTRODUCTION: COVID-19 related non-pharmaceutical interventions (NPIs) led to a suppression of RSV circulation in winter 2020/21 in the UK and an off-season resurgence in Summer 2021. We explore how the parameters of RSV epidemiology shape the size and dynamics of post-suppression resurgence and what we can learn about them from the resurgence patterns observed so far. METHODS: We developed an age-structured dynamic transmission model of RSV and sampled the parameters governing RSV seasonality, infection susceptibility and post-infection immunity, retaining simulations fitting the UK's pre-pandemic epidemiology by a set of global criteria consistent with likelihood calculations. From Spring 2020 to Summer 2021 we assumed a reduced contact frequency, returning to pre-pandemic levels from Spring 2021. We simulated transmission forwards until 2023 and evaluated the impact of the sampled parameters on the projected trajectories of RSV hospitalisations and compared these to the observed resurgence. RESULTS: Simulations replicated an out-of-season resurgence of RSV in 2021. If unmitigated, paediatric RSV hospitalisation incidence in the 2021/22 season was projected to increase by 30-60% compared to pre-pandemic levels. The increase was larger if infection risk was primarily determined by immunity acquired from previous exposure rather than age-dependent factors, exceeding 90 % and 130 % in 1-2 and 2-5 year old children, respectively. Analysing the simulations replicating the observed early outbreak in 2021 in addition to pre-pandemic RSV data, we found they were characterised by weaker seasonal forcing, stronger age-dependence of infection susceptibility and higher baseline transmissibility. CONCLUSION: COVID-19 mitigation measures in the UK stopped RSV circulation in the 2020/21 season and generated immunity debt leading to an early off-season RSV epidemic in 2021. A stronger dependence of infection susceptibility on immunity from previous exposure increases the size of the resurgent season. The early onset of the RSV resurgence in 2021, its marginally increased size relative to previous seasons and its decline by January 2022 suggest a stronger dependence of infection susceptibility on age-related factors, as well as a weaker effect of seasonality and a higher baseline transmissibility. The pattern of resurgence has been complicated by contact levels still not back to pre-pandemic levels. Further fitting of RSV resurgence in multiple countries incorporating data on contact patterns will be needed to further narrow down these parameters and to better predict the pathogen's future trajectory, planning for a potential expansion of new immunisation products against RSV in the coming years.

Evidence for influenza and RSV interaction from 10 years of enhanced surveillance in Nha Trang, Vietnam, a modelling study.

Influenza and Respiratory Syncytial Virus (RSV) interact within their host posing the concern for impacts on heterologous viruses following vaccination. We aimed to estimate the population level impact of their interaction. We developed a dynamic age-stratified two-pathogen mathematical model that includes pathogen interaction through competition for infection and enhanced severity of dual infections. We used parallel tempering to fit its parameters to 11 years of enhanced hospital-based surveillance for acute respiratory illnesses (ARI) in children under 5 years old in Nha Trang, Vietnam. The data supported either a 41% (95%CrI: 36-54) reduction in susceptibility following infection and for 10.0 days (95%CrI 7.1-12.8) thereafter, or no change in susceptibility following infection. We estimate that co-infection increased the probability for an infection in <2y old children to be reported 7.2 fold (95%CrI 5.0-11.4); or 16.6 fold (95%CrI 14.5-18.4) in the moderate or low interaction scenarios. Absence of either pathogen was not to the detriment of the other. We find stronger evidence for severity enhancing than for acquisition limiting interaction. In this setting vaccination against either pathogen is unlikely to have a major detrimental effect on the burden of disease caused by the other.

Drivers of Dyadic Cofeeding Tolerance in Pan: A Composite Measure Approach.

This study aimed to construct a composite model of Dyadic Cofeeding Tolerance (DCT) in zoo-housed bonobos and chimpanzees using a validated experimental cofeeding paradigm and to investigate whether components resulting from this model differ between the two species or vary with factors such as sex, age, kinship and social bond strength. Using dimension reduction analysis on five behavioral variables from the experimental paradigm (proximity, aggression, food transfers, negative food behavior, participation), we found a two-factor model: "Tolerant Cofeeding" and "Agonistic Cofeeding". To investigate the role of social bond quality on DCT components alongside species effects, we constructed and validated a novel relationship quality model for bonobos and chimpanzees combined, resulting in two factors: Relationship Value and Incompatibility. Interestingly, bonobos and chimpanzees did not differ in DCT scores, and sex and kinship effects were identical in both species but biased by avoidance of the resource zone by male-male dyads in bonobos. Social bonds impacted DCT similarly in both species, as dyads with high Relationship Value showed more Tolerant Cofeeding, while dyads with higher Relationship Incompatibility showed more Agonistic Cofeeding. We showed that composite DCT models can be constructed that take into account both negative and positive cofeeding behavior. The resulting DCT scores were predicted by sex, kinship and social bonds in a similar fashion in both Pan species, likely reflecting their adaptability to changing socio-ecological environments. This novel operational measure to quantify cofeeding tolerance can now be applied to a wider range of species in captivity and the wild to see how variation in local socio-ecological circumstances influences fitness interdependence and cofeeding tolerance at the dyadic and group levels. This can ultimately lead to a better understanding of how local environments have shaped the evolution of tolerance in humans and other species.