Geometry for the selfish herd
Abstract
This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator.
It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others.
Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations.
The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value.
References (34)
- E.E. Austin
- G.P. Baerends et al.
Behaviour
(1950) - G.M. Breder
Bull. Am. Mus. nat. Hist
(1959) - H.R. Bullis
Ecology
(1960) - P.J. Clark et al.
Science, N.Y
(1955) - J.H. Crook
Behaviour
(1960) - F.F. Darling
- Y. Espmark
Zool. Beitr
(1968) - U.R. Evans
Trans. Faraday Soc
(1945) - J. Fisher
Macmillan's Mag., Lond
Ann. math. Statist
Am. Nat
Zool. Anz
Cited by (2626)
Evolution of delayed dispersal with group size effect and population dynamics
2024, Theoretical Population BiologyIndividuals delay natal dispersal for many reasons. There may be no place to disperse to; immediate dispersal or reproduction may be too costly; immediate dispersal may mean that the individual and their relatives miss the benefits of group living. Understanding the factors that lead to the evolution of delayed dispersal is important because delayed dispersal sets the stage for complex social groups and social behavior. Here, we study the evolution of delayed dispersal when the quality of the local environment is improved by greater numbers of individuals (, safety in numbers). We assume that individuals who delay natal dispersal also expect to delay personal reproduction. In addition, we assume that improved environmental quality benefits manifest as changes to fecundity and survival. We are interested in how do the changes in these life-history features affect delayed dispersal. We use a model that ties evolution to population dynamics. We also aim to understand the relationship between levels of delayed dispersal and the probability of establishing as an independent breeder (a population-level feature) in response to changes in life-history details. Our model emphasizes kin selection and considers a sexual organism, which allows us to study parent–offspring conflict over delayed dispersal. At evolutionary equilibrium, fecundity and survival benefits of group size or quality promote higher levels of delayed dispersal over a larger set of life histories with one exception. The exception is for benefits of increased group size or quality reaped by the individuals who delay dispersal. There, the increased benefit does not change the life histories supporting delay dispersal. Next, in contrast to previous predictions, we find that a low probability of establishing in a new location is not always associated with a higher incidence of delayed dispersal. Finally, we find that increased personal benefits of delayed dispersal exacerbate the conflict between parents and their offspring. We discuss our findings in relation to previous theoretical and empirical work, especially work related to cooperative breeding.
The effect of intraspecific cooperation in a three-species cyclic predator-prey model
2024, Applied Mathematics and ComputationThe maintenance of biological diversity has perpetually remained a central focus in the field of ecology. In the pursuit of enhanced survival rates, species have begun to explore cooperation with one another. However, the consequences of such collaboration remain largely unexplored. To delve into this matter, we introduce intraspecific cooperation within the framework of the classic rock-paper-scissors (RPS) game. In this model, the competition rate is intricately tied to interactions among individuals of the same species. A greater population of individuals from the same species tends to lead to an increased predation rate and a decreased prey rate. Through extensive simulations, we observe that (i) in the case of homogeneous intraspecific cooperation (all three species have intraspecific cooperation), increased cooperation between predators tends to increase the likelihood of species coexistence. In contrast, high levels of cooperation between prey appeared to decrease the favorability of species coexistence. Measurements of the characteristic length of spiral structures revealed that the characteristic length of spirals became longer when the intensity of prey cooperation increased. (ii) In the case of heterogeneous intraspecific cooperation (two species or only one species with intraspecific cooperation), neither an increase in the intensity of intraspecific cooperation of the predator nor an increase in the intensity of intraspecific cooperation of the prey is favorable for species coexistence. Our work underscores the critical role of intraspecific cooperation in maintaining biodiversity.
Plants buffer some of the effects of a pair of cadmium-exposed zebrafish on the un-exposed majority
2024, Environmental Toxicology and PharmacologyCertain individuals have a disproportionate effect on group responses. Characteristics may include susceptibility to pollutants, such as cadmium (Cd), a potent trace metal. Here, we show how a pair of Cd-exposed individuals can impact the behavior of unexposed groups. We used behavioral assessments to characterize the extent of the effects of the Cd-exposed individuals on group boldness, cohesion, foraging, activity, and responses to plants. We found that groups with a pair of Cd-exposed fish remained closer to novel stimuli and plants than did groups with untreated (control) fish. The presence of plants reduced Cd-induced differences in shoal cohesion and delays feeding in male shoals. Shoals with Cd- and water-treated fish were equally active. The results suggest that fish acutely exposed to environmentally relevant Cd concentrations can have profound effects on the un-exposed majority. However, the presence of plants may mitigate the effects of contaminants on some aspects of social behavior.
On aims and methods of collective animal behaviour
2024, Animal BehaviourCollective animal behaviour is a subfield of behavioural ecology, making extensive use of its tools of observation, experimental manipulation and model building. However, a fundamental behavioural ecology approach, the application of optimality theory, has been comparatively neglected in collective animal behaviour. This article seeks to address this imbalance, by outlining an evolutionary theory framework for the discipline. The application of optimality theory to collective animal behaviour requires a number of questions to be addressed. First, what is the correct quantity to optimize? This can be achieved via a combination of considering the organisms' life history, alongside tools such as statistical decision theory and stochastic dynamic programming. Second, what mechanism is appropriate for optimal behaviour? This involves ensuring that models are self-consistent rather than assuming parameter values. Third, at what level of selection does optimization act? Selection acts on the individual except in very particular circumstances, yet collective animal behaviour phenomena are group level, thus introducing a risk of confusing at what level adaptive properties emerge. This article presents examples under each of the three questions, as well as discussing mismatches between theory and observation. In doing so, it is hoped that collective animal behaviour fully inherits the tools and philosophy of its parent discipline of behavioural ecology.
Group-housed cattle may engage in agonistic interactions over resources such as feed, which can negatively affect aspects of welfare. Little is known about how contextual factors such as group size influence agonistic behaviour. We explored the frequency of agonistic interactions at the feeder when cattle were housed in different-sized groups. We also explored the consistency of the directionality of agonistic interactions in dyads and of the number of agonistic interactions initiated by individuals across the group sizes. Four replicates of 50 cows each were assessed in two group-size phases. In Phase 1, cows were kept in one group of 50. In Phase 2, these same cows were divided into five groups of 10, maintaining stocking density (i.e., ratio of animals to lying stalls and feed bunk spaces). We measured agonistic replacements (i.e., interactions that result in one cow leaving the feed bin and another taking her place) at an electronic feeder using a validated algorithm. We used these data from Phase 1 to calculate individual Elo-ratings (a type of dominance score). Cows were then categorised into five dominance categories based upon these ratings. To ensure a consistent Elo-rating distribution between phases, two cows from each dominance category were randomly assigned to each small group of 10 cows. The mean ± SE number of replacements per cow was similar regardless of whether the cows were housed in groups of 50 (34.1 ± 2.4) or 10 (31.1 ± 4.5), although the groups of 10 were more variable. Further, 81.6 ± 7.7% (mean ± SD) of dyads had the same directionality across group sizes (i.e., the same individual won the majority of interactions in the dyad) and individuals were moderately consistent in the number of replacements they initiated (intraclass correlation coefficient = 0.62 ± 0.11; mean ± SD). These results indicate that the relationship between group size and agonistic behaviour is complex; we discuss these challenges and suggest new avenues for further research.
Grounding social timing: A commentary on “The evolution of social timing” by Verga et al. (2023)
2024, Physics of Life ReviewsWe are excited about Verga et al.'s [22], [22] exhortation to look beyond humans to understand the purpose, scope, and evolution of social timing. We argue that the field should expand even further. We first point out the enabling role of the spatial environment, which constrains social interaction and in which social interaction is embedded. Second, we argue that a full appreciation of the emergence of social timing must include a focus on physical prerequisites of interactive systems, exemplified by studies of dissipative structures more broadly. By situating interacting systems—whether biological or not—within their shared dynamic environment, we can more clearly and more fully understand social timing.