Social Evolution: New Horizons

Cooperation is a widespread natural phenomenon yet current evolutionary thinking is dominated by the paradigm of selfish competition. Recent advanced in many fronts of Biology and Non-linear Physics are helping to bring cooperation to its proper place. In this contribution, the most important controversies and open research avenues in the field of social evolution are reviewed. It is argued that a novel theory of social evolution must integrate the concepts of the science of Complex Systems with those of the Darwinian tradition. Current gene-centric approaches should be reviewed and com- plemented with evidence from multilevel phenomena (group selection), the constrains given by the non-linear nature of biological dynamical systems and the emergent nature of dissipative phenomena.Comment: 16 pages 5 figures, chapter in forthcoming open access book "Frontiers in Ecology, Evolution and Complexity" CopIt-arXives 2014, Mexic

A literature review of connectedness to nature and its potential for environmental management

Understanding how people's relationships with nature form, how they influence personal values and attitudes, and what behavioural implications they may have could provide more insight into how connectedness to nature (CNT) can effectively contribute to environmental management goals. This paper undertakes a review of literature published over the past decade (2002e2011) on SCOPUS; and describes the current state of knowledge regarding CNT, assesses any efforts towards the spatial mapping of CNT for environmental management, and identifies measures of CNT defined in the broader literature. This review suggests that there is quite some overlap in the literature on CNT concepts, and that more effort needs to be made towards multi-disciplinary research which explores how CNT can be useful to environmental planning and conservation research on the field. It also further corroborates the need and relevance of applying more social and affective strategies to promote conservation behaviour. The main progress in CNT theory seems to have been made in the development of measurement tools, and it is clear that there is a strong convergent validity amongst the different measures due to their similarity, and functional associations. Further efforts towards the exploration of multi-dimensional measures is recommended since they consistently stand out as showing better results. The geographic visualisation of CNT constructs is another area of research that deserves attention since it can provide a unique point of view towards guiding participatory protected area planning and management.peer-reviewe

Movement across scales: red fox spatial ecology

Rødrevens forflytningsmønster på forskjellige romlige skalaer Menneskelig påvirkning på naturlige habitater kan medføre reduserte bevegelsesmuligheter for noen dyrearter, eller fasilitere typiske generalister som rødrev Vulpes vulpes. Vår forståelse av rødrev i landskaper dominert av menneskelige aktiviteter er begrenset av manglende kunnskap om hvordan revene beveger seg i disse områdene. I denne avhandlingen undersøkte jeg temporale og romlige mønstre i forflytningene til rødrever langs en gradient av menneskelig påvirkning ved bruk av GPS-telemetridata innhentet i Norge og Sverige fra 2011 til 2019. Rødrevenes leveområder var langt større enn observert i tidligere studier, og med en markant individuell variasjon som delvis kunne relateres til miljøfaktorer langs en landskapsgradient. I lavereliggende sørlige områder med relativt høy primærproduksjon og mye landbruk var revenes leveområder fire ganger mindre enn i høyereliggende nordlige barskogområder. I gjennomsnitt var 43% av revenes posisjoner innenfor klustere som dekket bare 1% av leveområdene. Dette indikerte at rødrevenes kognitive kartlegging innebar repetitive bevegelser som bidro til avgrensing av leveområdene. Noen rever viste også en betydelig evne til å forflytte seg lange avstander under spredning, mellom populasjoner og landskaper, og potensielt over nasjonale grenser. Allikevel viste revene en genetisk struktur på liten romlig skala som var knyttet til sosiale mekanismer heller enn forflytningsevne og spredningskapasitet. Parvise distanser mellom nært beslektede hunndyr (gjennomsnitt = 6.3 km) var signifikant kortere enn avstandene mellom beslektede hanner (37.8 km). Dette understreker at sosiale forhold (som slektskap) spiller en viktig rolle i rødrevenes romlige organisasjon. Mine resultater viser at forståelsen av rødrevens forflytninger forutsetter god innsikt i hvilken temporal og romlig skala bevegelsene foregår. Å klassifisere unike og variable atferdstrekk hos en svært fleksibel art som rødrev er veldig vanskelig, og dette understreker artens økologiske plastisitet. Alt i alt har denne avhandlingen gitt ny innsikt i hvordan rødrevens forflytningsmønster er påvirket av sosial struktur og miljøfaktorer på forskjellige romlige skalaer. Dette har betydning for framtidig forskning og forvaltning, samt for modellering av revenes demografi og sykdomsspredning. Informasjonen gir økt innsikt i rødrevens områdebruk og spredning i rurale områder, og åpner for nye muligheter for forskning på denne artens påvirkning på økosystemer.Movement across scales: red fox spatial ecology The impact of human activities is altering natural habitats, reducing the ability of some animals to move, while facilitating other, generalist species, such as the red fox Vulpes vulpes. Our understanding of red foxes in rural and human modified landscapes is constrained by a lack of knowledge about how foxes use these landscapes. In this thesis I investigated the spatiotemporal movement patterns of red foxes along a landscape gradient of human influence using individual based GPS telemetry data from red foxes collected in Norway and Sweden between 2011- 2019. Herein, I identified much larger home ranges than previously recorded for red foxes and a high degree of individual variation, partially explained through environmental factors along a landscape gradient. At lower elevations, where productivity and the amount of available agricultural land increased, red foxes had home ranges approximately four times smaller than the home ranges of foxes in the northern boreal vegetation areas. I also identified cognitive mapping as a feature of red fox space use, linked to recursive movements within home ranges and contributing to bounded space use. On average, 43% of a red fox’s positions were found in defined clusters that covered a proportional area of only 1% of their home range. I highlighted the red fox’s ability to traverse between populations, across landscapes, and potentially across international boundaries, by identifying six long-distance dispersal events, representing some of the longest dispersal distances recorded for red foxes. However, I also showed that fine scale familial structuring in red foxes occurred by social mechanisms not linked to their movement ability or dispersal capacity. I found significant differences in pairwise geographic distances between highly related same sex pairs with the average distance between related males, 37.8 km, being six times farther than that of related females, averaging 6.3 km. This highlights how social dynamics (e.g. kin clustering and female philopatry) play a role in the spatial organization of red foxes. Finally, I showed that recognizing red fox behaviors is dependent on not only identifying their associated movement patterns, but also understanding the temporal and spatial scales at which their movements occur. Identifying the unique and variable behaviors of a highly flexible species such as the red fox is difficult and shows the ecological plasticity of the species. Together, this information represents new observations that greatly expand our knowledge of red fox space use and dispersal in rural landscapes and opens the door for future research into the broader ecosystem consequences of such movements. Overall, this thesis increases our understanding of red fox movement behaviors and their interactions with social and environmental factors at multiple spatial scales, with implications for future research, management and demographic and disease modeling.publishedVersio

Context‐dependent dispersal determines relatedness and genetic structure in a patchy amphibian population

Dispersal is a central process in ecology and evolution with far reaching consequences for the dynamics and genetics of spatially structured populations (SSPs). Individuals can adjust their decisions to disperse according to local fitness prospects, resulting in context-dependent dispersal. By determining dispersal rate, distance, and direction, these individual-level decisions further modulate the demography, relatedness, and genetic structure of SSPs. Here, we examined how context-dependent dispersal influences the dynamics and genetics of a Great Crested Newt (Triturus cristatus) SSP. We collected capture-recapture data of 5564 individuals and genetic data of 950 individuals across a SSP in northern Germany. We added genetic data from six sites outside this SSP to assess genetic structure and gene flow at a regional level. Dispersal rates within the SSP were high but dispersal distances were short. Dispersal was context-dependent: individuals preferentially immigrated into high-quality ponds where breeding probabilities were higher. The studied SSP behaved like a patchy population, where subpopulations at each pond were demographically interdependent. High context-dependent dispersal led to weak but significant spatial genetic structure and relatedness within the SSP. At the regional level, a strong hierarchical genetic structure with very few first-generation migrants as well as low effective dispersal rates suggest the presence of independent demographic units. Overall, our study highlights the importance of habitat quality for driving context-dependent dispersal and therefore demography and genetic structure in SSPs. Limited capacity for long-distance dispersal seems to increase genetic structure within a population and leads to demographic isolation in anthropogenic landscapes.Microsatellite Genotypes: Missing values are coded "-9". Presence/Absence Data: Missing values are coded "-". Funding provided by: Deutsche ForschungsgemeinschaftCrossref Funder Registry ID: http://dx.doi.org/10.13039/501100001659Award Number: STE 1130/7-1Demographic Data (CMR and Presence/Absence Data): We surveyed 33 water bodies using mark-recapture methods for the presence, demography and reproduction of crested newts between 2012 and 2014. Newts were captured during two capture sessions (cs) per year, one early (April/May) and one late (June/July) in the breeding season. Every capture session thereby consisted of three consecutive capture events in intervals of two days. Within the context of a presence/absence analysis, all sites were surveyed for one more day in late July/early August in order to detect larvae. If a pond dried out and was therefore not surveyed during a capture session, such an event was treated as a missing observation. Newts were captured using Ortmann's funnel traps which were evenly distributed along the shoreline of a pond. The number of traps deployed per capture event varied according to pond perimeter (one trap per 10m shoreline), ranging from one to 27 traps. For individual recognition of newts during the CMR study, we used photographs of the ventral side of an individual which provides a natural marking in form of a highly variable but individually unique and stable color pattern through the time. Recaptured individuals were identified automatically by the software AmphIdent. Microsatellite Genotypes: Tissue samples were taken from seven sampling sites by puncturing the tails of captured great crested newts (Triturus cristatus) using micro haematocrit capillary tubes (Carl Roth, Ø 1.6 mm) and were then stored in 80% ethanol. Total genomic DNA was extracted using the sodium dodecyl sulfate (SDS)-proteinase K/ Phenol-Chloroform extraction method. Genomic DNA was stored in Tris-EDTA buffer (10 mM Tris-HCl, 0.1 mM EDTA, pH 8.0) and used for all subsequent reactions. Each individual sample was mugenotyped for 17 microsatellite loci. Primers were combined in three multiplex mixes (Drechsler et al., 2013). 10 µl Type-it Multiplex PCRs (Qiagen) containing 1 µl of genomic DNA were performed. The PCR profile was as follows: (1) 5 min at 95°C, (2) 30 s at 94°C, (3) 90 s at an annealing temperature of 60°C, (4) 60 s at 72°C, (5) return to step 2 for 30 times, (6) 30 min at 60°C. Obtained PCR products were diluted with 50-200 μl water depending on the strength of obtained PCR products. 1 µl of each diluted multiplex reaction was added to 20 μl of Genescan 500-LIZ size standard (Applied Biosystem) and then run on an ABI 3730 96-capillary or an ABI 3130 16-capillary automated DNA-sequencer. Allele scoring of microsatellite loci was performed using Genemarker software (SoftGenetics version 1.95)

Extended Inclusive Fitness Theory bridges Economics and Biology through a common understanding of Social Synergy

Inclusive Fitness Theory (IFT) was proposed half a century ago by W.D. Hamilton to explain the emergence and maintenance of cooperation between individuals that allows the existence of society. Contemporary evolutionary ecology identified several factors that increase inclusive fitness, in addition to kin-selection, such as assortation or homophily, and social synergies triggered by cooperation. Here we propose an Extend Inclusive Fitness Theory (EIFT) that includes in the fitness calculation all direct and indirect benefits an agent obtains by its own actions, and through interactions with kin and with genetically unrelated individuals. This formulation focuses on the sustainable cost/benefit threshold ratio of cooperation and on the probability of agents sharing mutually compatible memes or genes. This broader description of the nature of social dynamics allows to compare the evolution of cooperation among kin and non-kin, intra- and inter-specific cooperation, co-evolution, the emergence of symbioses, of social synergies, and the emergence of division of labor. EIFT promotes interdisciplinary cross fertilization of ideas by allowing to describe the role for division of labor in the emergence of social synergies, providing an integrated framework for the study of both, biological evolution of social behavior and economic market dynamics.Comment: Bioeconomics, Synergy, Complexit

An Exploration of the Relationships Among Connectedness to Nature, Quality of Life, and Mental Health

The current study examined the relationships among connectedness to nature (CTN), quality of life (QOL), and mental health (MH). Theory in biophilia and ecopsychology has emphasized the importance of the human relationship with the natural world for the health of individuals, our species, and our planet as a whole. Previous research has documented the relationship between experiences in nature and outcomes of health and well-being. However, scant research has examined the correlates of the concept of CTN. Furthermore, no research has examined the relationship between CTN and measures of well-being such as QOL or MH. In the current study, 267 undergraduate students completed a series of self-report items measuring CTN, QOL, MH, and demographic characteristics. Significant relationships between CTN and all categories of dependent variables (QOL, MH, and demographics) were found. In addition, the subcategory of CTN closely related to the desire for direct experiences in nature, NR experience was shown to have a stronger relationship to QOL and MH than overall CTN--as evidenced by more significant correlations and by serving as a better predictive model through multiple regression. QOL, MH, and demographic variables were collectively found to predict 21% of the variance in overall CTN, while those same variables were found to predict 35% of the variance in NR experience. Implications of these findings based on previous research, limitations of the current study, and future avenues of research are discussed

Four not six: revealing culturally common facial expressions of emotion

As a highly social species, humans generate complex facial expressions to communicate a diverse range of emotions. Since Darwin’s work, identifying amongst these complex patterns which are common across cultures and which are culture-specific has remained a central question in psychology, anthropology, philosophy, and more recently machine vision and social robotics. Classic approaches to addressing this question typically tested the cross-cultural recognition of theoretically motivated facial expressions representing six emotions, and reported universality. Yet, variable recognition accuracy across cultures suggests a narrower cross-cultural communication, supported by sets of simpler expressive patterns embedded in more complex facial expressions. We explore this hypothesis by modelling the facial expressions of over 60 emotions across two cultures, and segregating out the latent expressive patterns. Using a multi-disciplinary approach, we first map the conceptual organization of a broad spectrum of emotion words by building semantic networks in two cultures. For each emotion word in each culture, we then model and validate its corresponding dynamic facial expression, producing over 60 culturally valid facial expression models. We then apply to the pooled models a multivariate data reduction technique, revealing four latent and culturally common facial expression patterns that each communicates specific combinations of valence, arousal and dominance. We then reveal the face movements that accentuate each latent expressive pattern to create complex facial expressions. Our data questions the widely held view that six facial expression patterns are universal, instead suggesting four latent expressive patterns with direct implications for emotion communication, social psychology, cognitive neuroscience, and social robotics

The Role of Individual Cognitive and Behavioral Ontogeny in Organization and Evolution of Social Systems

Exploration and explanation of the relationship between individual variation in behavior and the composition and adaptive success of social groups or populations are crucial problems in the fields of behavioral ecology, ethology, and comparative psychology. These questions have been the subject of a longstanding discussion at both the proximate and ultimate levels of inquiry. Adaptive mechanisms explaining social decision making, both in terms of affiliative and competitive partner choices, are at the center of such discussions. Inclusive fitness, kin-selection, handicap or prestige, risk seeking and risk avoiding strategies, pay-to-stay/reward principles, as well as other theories have been proposed and supported as these mechanisms in a variety of taxa; theories which may not be mutually exclusive. This dissertation focuses on the role of individual ontogeny in the organization of a series of charismatic social systems. Specifically, I review evolutionary aspects of siblicidal and brood parasitic systems, and present research on social/cognitive ontogeny and interactive behavior of a passerine avian cooperative breeder (Turdoides squamiceps), a phenotypically reversible teleost fish (Astatotilapia burtoni), and a eusocial mammal (Heterocephalus glaber). The theme adaptive behavioral response to fluctuating or uncertain environments and interactions unites these taxonomically diverse subjects. In Chapter 1 I review a potential evolutionary trajectory from siblicidal behavior to both obligate and intraspecific brood parasitism. This analysis focuses on a series of fitness inequalities that may function as the most parsimonious explanation of such an evolutionary trajectory. When resources provided by parents are limited, full siblings may be driven to siblicide. The inherent fitness cost to parents of siblicidal behavior by offspring may be offset by brood parasitism. Brood parasitism, however, carries its own costs, both in conspecific/intraspecific brood parasitism (where individuals in a population may still be related, requiring a kin-selected balance for any fitness advantage to result) and in interspecific brood parasitism, where the host may not be equipped to nourish offspring as effectively as biological parents. Chapter 1 also discusses the ultimate mechanisms for the evolution of one ontogenetic style to the others. For intraspecific brood parasitism to evolve from siblicide, the alleles shared with the parasitic parents by the surviving, parasitically laid offspring must exceed those eliminated from the population via competition between the parasitically laid offspring and host offspring. For interspecific brood parasitism to evolve from siblicide or intraspecific brood parasitism, offspring lost to siblicide or related individuals lost to intraspecific brood parasitism must be less than those lost to nutritional or behavioral mismatches which result from heterospecific parental care by foreign host taxa. Chapters 2 and 3 were developed in association with the Arabian Babbler Research Project, a 40+ year old ongoing study using a habituated and ID banded population of Arabian Babblers in Hazeva, Israel, maintained by Professor Amotz Zahavi. The habituation of these cooperatively breeding passerines to observer presence permits detailed long-term data sets and analysis of individual life histories. Arabian Babblers live in exclusive, male philopatric social groups with high reproductive skew, where rank is strongly positively correlated with age. In Chapter 2 I demonstrate the relationship between ontogenetic experience and developmental stage and neophilic behavior in young Arabian Babblers (Turdoides squamiceps). I used a series of novel/familiar stimulus presentations to identify the latency to and frequency of approach to stimuli by young individuals. Stimuli yielded multimodal comparisons, including stationary objects, moving objects, and sounds. Each had a familiar and a novel condition. I found that all birds approached novel stimuli more frequently than they did familiar stimuli, and that intrabrood rank positively predicted frequency of approach. Additionally, juveniles were more likely to approach novel stimuli, and did so earlier in the presentation trials, than fledglings. All young individuals were more likely to approach when adults were present. In Chapter 3 I use a dataset collected from 2002-2004 by members of the Arabian Babbler Research Project which details the behaviors occurring before and during the formation of allopreening dyads. Chapter 3 analyzes the role of autopreening in the formation of allopreening dyads, and presents evidence that it is a displacement behavior. Autopreening may occur before the social approach that is necessary to form an allopreening dyad. When relationships were hierarchically more certain (represented by an older actor in the dyad, or the formation of the dyad without invitation by the recipient) approach by the recipient occurred without autopreening. When recipients did autopreen, they were significantly less likely to approach the actor to form the dyad. In Chapter 4 I present data from observation of a phenotypically reversible cichlid fish Astatotilapia burtoni. Socially mediated morphological plasticity in this species is largely driven by male intrasexual competition, and it is physiologically necessary for a male to develop the territorial phenotype in order to reproduce. Traditionally, analysis of A. burtoni communities divides males between territorial (DOM) and non-territorial (SUB) individuals based on an index that subtracts the total losses of an individual in intrasexual conflict from the combined total wins and courtship behavior exhibited by that individual. There are dramatic morphological, physiological, and behavioral differences between these two categories, defined as having either positive (DOM) or negative (SUB) dominance index scores. Chapter 4 uses cluster analyses to propose a formalization of a third male phenotype, identified with individuals transitioning between prototypical DOMs and prototypical SUBs. Specifically, a novel behavior was identified; a potential risk-avoidance strategy in which individuals appear to ignore challenges, rather than engage or flee. Other unique behavioral traits of this male phenotype, such as frequency of certain pigment displays, were identified, and the individuals were shown to be those more likely to transition across the traditional DOM/SUB division point of a zero score on the dominance index than either prototypical DOMs or prototypical SUBs. In Chapter 5 I present the results of a preliminary rescue-behavior experiment using a eusocial mammal, the Naked Mole-Rat Heterocephalus glaber. The study population was individually tracked using subcutaneous RFID tags. This method permits high temporal resolution on location of individuals. The rescue scenario involved the experimental trapping of individuals at the distal end of tubes connected to the central enclosure. Cork was used to create an artificial “cave-in” that served as an obstacle for colony members. A plastic barrier prevented the escape of trapped individuals. A second permutation of the experiment used a bifurcated tube to present experimentally trapped individuals and empty space, both blocked by cork, simultaneously. Colony members were significantly quicker to excavate trapped individuals than empty space. Effort expended (defined as time spent in rescue and latency to initiation of rescue) varied by both caste and individual, with queens expending less effort than workers, and individuals initiating rescue events also being the most frequent actors in excavation. In Chapter 6 I discuss the connections among these findings, as well as their relevance to contemporary questions in research on behavioral ecology and comparative psychology. Together, these manuscripts provide a taxonomically varied perspective on a central issue in sociobiology and cognitive ethology: the illumination of the role of individual ontogenetic experience on the adaptive function of social groups. This dissertation does not attempt to represent an exhaustive investigation of this complex subject. Instead, it highlights promising avenues of investigation and demonstrates that social systems which may differ greatly in organization and evolutionary history remain strongly influenced and constructed by interindividual variation in ontogeny and experience, with particular regard to decision making