Contest Competition Research Articles

The changing South in presidential elections

We examine presidential elections in the American South. Our empirical starting point is the 2008 election of Democrat Barack Obama. We discuss political dynamics leading up to this historic contest but focus our analysis on the last four elections concluding with the defeat of President Trump in 2020. Tapping into recent southern politics scholarship emphasizing partisan patterns that differ based on population growth (or lack thereof), we show how this factor affects presidential voting with an assemblage of data collected at the state-, county-, and individual-level. We first consider the prominence of presidential Republicanism across the 11 southern states. Next, we delve into these states’ counties to further assess two-party competition in presidential contests. Finally, drawing on Cooperative Election Study (CES) data, controlling for a host of factors, we examine whether varying population rates at the state- and county-level affect presidential preferences. The U.S. South is forever changing, and in presidential politics substantial alterations to its electorate generally favor an emerging Democratic Party.

Local population dynamics of gray wolves Canis lupus and Eurasian lynx Lynx lynx exhibit consistency with intraspecific contest competition models

AbstractIn Europe, the gray wolf and Eurasian lynx populations are recovering after various levels of persecution. The two species differ in their social structure and spatial patterns of aggregation. Using model selection, we investigated the consistency of the available time series data on local wolf and lynx sub‐populations with a number of single‐species population growth models that pertain to two types of intraspecific competition, namely, scramble (SC) and contest competition (CC), and reflect random (R) or aggregated (A) distribution of individuals. The applied models of population growth—the Ricker (SCR), Skellam (CCR), Hassell (SCA), and Beverton–Holt (CCA) models—were all parameterized in terms of intrinsic growth rate and carrying capacity with unified definitions. The projected carrying capacity was allowed to show a temporal trend, which was justified by an observed increase in prey abundance in recent decades. For both species, the models pertaining to contest competition outperformed the scramble competition models, and the Beverton–Holt model had the greatest weight. However, for the lynx, the difference of performance between the scramble and contest competition models was considerably smaller than that for the wolves. In most of the models, when it was meaningful, an optional time lag operator was added to account for a delay in individual maturity and reproduction. However, the models with a time lag had a worse fit than the models without it. This study promotes the application of population models that reflect intraspecific competition for modeling population dynamics in a single‐ or multi‐species framework.

Using social networks to explore the social flexibility of urban vervet monkeys

Increasing urbanisation reduces available habitat and increases human-wildlife interactions, presenting social and ecological challenges for many species; however, flexible generalist species, such as the vervet monkey, Chlorocebus pygerythrus, thrive under these pressures. In the urban mosaic, human-food sources represent clumped, monopolisable food that can increase contest competition. Social network analysis (SNA) is a powerful tool to monitor changes in social structure, yet it has rarely been used to study urban wildlife. Using SNA, we investigated the effect of anthropogenic food and human-wildlife interactions on social cohesion in five vervet monkey groups in urban KwaZulu-Natal, South Africa. Over six months, we conducted group scan samples every 30-min on each group and recorded all humans-vervet monkey interactions during dawn to dusk follows. We analysed the effect of foraging on natural and human-related food sources and human-vervet monkey interactions on social network metrics for grooming and aggression at group (density, clustering coefficient and distance) and individual (eigenvector centrality and degree) levels, using linear mixed models. Anthropogenic food influenced almost all social metrics. At the group level, foraging on anthropogenic food was related to increased density and cohesion in both grooming and aggression networks. At the individual level, increasing anthropogenic food affected high-ranking monkeys most: eigenvector centrality and outdegree in aggression networks increased with rank. Social network analysis can be a useful tool to document urban effects on wildlife groups, and aids our understanding of wildlife behavioural flexibility, a key tool in developing educated and effective management strategies.

Examining the influence of prey dynamics on predator–prey interactions

ABSTRACT We develop variations of a discrete-time predator–prey model to examine how intrinsic properties of the prey population may impact overall system dynamics. We focus on two properties of the prey species, namely developmental stage structure and undercompensatory (contest competition) versus overcompensatory (scramble competition) density dependence. Through analysis of these models, we examine how these different prey features affect system stability. Our results show that when prey growth is overcompensatory, the predator may have a stabilizing effect on the system dynamics with increasing predator density reversing the period doubling route to chaos observed with Ricker-type nonlinearities. Moreover, we find that stage structure in the prey does not have a destabilizing effect on the system dynamics unless the prey projection matrix is imprimitive or close to imprimitive, as may occur for semelparous species. In this case, a sufficiently large predator density may stabilize cycles that are otherwise unstable.

Contest competition and injury in adult male sub-Antarctic fur seals

AbstractWe recorded intra-sexual behaviour of adult male sub-Antarctic fur seals Arctocephalus tropicalis at Gough Island, Southern Ocean, during the 1975/76 summer breeding season. Our re-analysed data address male ‘contest competition’, which relates to the costs of intra-sexual disputes, including fights. We considered the risks/benefits of fighting through investigation of injuries (n = 353) sustained by adult males (n = 124) in fights. Injuries were predominantly on the forequarters, especially around the insertion areas of the front flippers (41%) with its sparse pelage, compared to the neck and chest areas combined (29%), an area which is well protected by thick pelage. The infliction of serious, sometimes debilitating, injuries to competitors increases a male’s access to females. Injuries predominate in injured, defeated males that gather at non-breeding sites, suggesting that injuries inflicted by dominant males were successful in excluding competing males from breeding sites.

Social stability via management of natal males in captive rhesus macaques (Macaca mulatta)

ABSTRACT Keystone individuals are expected to disproportionately contribute to group stability. For instance, rhesus macaques (Macaca mulatta) who police conflict contribute towards stability. Not all individuals’ motivations align with mechanisms of group stability. In wild systems, males typically disperse at maturity and attempt to ascend via contest competition. In a captive system, dispersal is not naturally enabled – individuals attempt to ascend in their natal groups, which can be enabled by matrilineal kin potentially destabilizing group dynamics. We relocated select high-ranking natal males from five groups and assessed group stability before and after. We quantified hierarchical metrics at the individual and group level. After removal, we found significantly higher aggression against the established hierarchy (reversals), indicative of opportunistic attempts to change the hierarchy. Mixed-sex social signaling became more hierarchical, but the strength of this effect varied. Stable structure was not uniformly reached across the groups and alpha males did not all benefit. Indiscriminate natal male removal is an unreliable solution to group instability. Careful assessment of how natal males are embedded within their group is necessary to balance individual and group welfare.

Properties and interpretation of the Skellam model—A discrete‐time contest competition population model

AbstractThe Skellam model describes discrete‐time population dynamics of a single species assuming uniform (i.e., random or Poissonian) individual distribution and intraspecific contest competition. Apart from studies on individual‐based models derived from first principles it has been rarely applied in ecological research although in specific situations it may be more appropriate than, for instance, the frequently used Ricker model, which is derived assuming scramble competition among the individuals. In this article, we offer an insight into the first principles underlying the Skellam model and provide an alternative parameterization of the model in terms of two commonly used parameters: intrinsic rate of population increase and carrying capacity. We also provide guidelines and software for fitting the Skellam model to discrete population time series data. In light of these findings, the Skellam model may be a useful alternative for a range of purposes where it has been earlier overlooked.

Finding analytical approximations for discrete, stochastic, individual-based models of ecology

Discrete time, spatially extended models play an important role in ecology, modelling population dynamics of species ranging from micro-organisms to birds. An important question is how ’bottom up’, individual-based models can be approximated by ’top down’ models of dynamics. Here, we study a class of spatially explicit individual-based models with contest competition: where species compete for space in local cells and then disperse to nearby cells. We start by describing simulations of the model, which exhibit large-scale discrete oscillations and characterize these oscillations by measuring spatial correlations. We then develop two new approximate descriptions of the resulting spatial population dynamics. The first is based on local interactions of the individuals and allows us to give a difference equation approximation of the system over small dispersal distances. The second approximates the long-range interactions of the individual-based model. These approximations capture demographic stochasticity from the individual-based model and show that dispersal stabilizes population dynamics. We calculate extinction probability for the individual-based model and show convergence between the local approximation and the non-spatial global approximation of the individual-based model as dispersal distance and population size simultaneously tend to infinity. Our results provide new approximate analytical descriptions of a complex bottom-up model and deepen understanding of spatial population dynamics.

Spatial competition in a global disturbance minimum; the seabed under an Antarctic ice shelf

The marine habitat beneath Antarctica's ice shelves spans ∼1.6 million km2, and life in this vast and extreme environment is among Earth's least accessible, least disturbed and least known, yet likely to be impacted by climate-forced warming and environmental change. Although competition among biota is a fundamental structuring force of ecological communities, hence ecosystem functions and services, nothing was known of competition for resources under ice shelves, until this study. Boreholes drilled through a ∼ 200 m thick ice shelf enabled collections of novel sub-ice-shelf seabed sediment which contained fragments of biogenic substrata rich in encrusting (lithophilic) macrobenthos, principally bryozoans – a globally-ubiquitous phylum sensitive to environmental change. Analysis of sub-glacial biogenic substrata, by stereo microscopy, provided first evidence of spatial contest competition, enabling generation of a new range of competition measures for the sub-ice-shelf benthic space. Measures were compared with those of global open-water datasets traversing polar, temperate and tropical latitudes (and encompassing both hemispheres). Spatial competition in sub-ice-shelf samples was found to be higher in intensity and severity than all other global means. The likelihood of sub-ice-shelf competition being intraspecific was three times lower than for open-sea polar continental shelf areas, and competition complexity, in terms of the number of different types of competitor pairings, was two-fold higher. As posited for an enduring disturbance minimum, a specific bryozoan clade was especially competitively dominant in sub-ice-shelf samples compared with both contemporary and fossil assemblage records. Overall, spatial competition under an Antarctic ice shelf, as characterised by bryozoan interactions, was strikingly different from that of open-sea polar continental shelf sites, and more closely resembled tropical and temperate latitudes. This study represents the first analysis of sub-ice-shelf macrobenthic spatial competition and provides a new ecological baseline for exploring, monitoring and comparing ecosystem response to environmental change in a warming world.

Stability results for a hierarchical size-structured population model with distributed delay

In this paper we investigate a structured population model with distributed delay. Our model incorporates two different types of nonlinearities. Specifically we assume that individual growth and mortality are affected by scramble competition, while fertility is affected by contest competition. In particular, we assume that there is a hierarchical structure in the population, which affects mating success. The dynamical behavior of the model is analysed via linearisation by means of semigroup and spectral methods. In particular, we introduce a reproduction function and use it to derive linear stability criteria for our model. Further we present numerical simulations to underpin the stability results we obtained.

Alternative reproductive tactics of unflanged and flanged male orangutans revisited.

In many slowly developing mammal species, males reach sexual maturity well before they develop secondary sexual characteristics. Sexually mature male orangutans have exceptionally long periods of developmental arrest. The two male morphs have been associated with behavioral alternative reproductive tactics, but this interpretation is based on cross-sectional analyses predominantly of Northwest Sumatran populations. Here we present the first longitudinal analyses of behavioral changes of 10 adult males that have been observed in both unflanged and flanged morph. We also analyzed long-term behavioral data on an additional 143 individually identified males from two study sites, Suaq (Sumatra, Pongo abelii) and Tuanan (Borneo, Pongo pygmaeus wurmbii), to assess male mating tactics cross-sectionally in relation to population, male morph (unflanged and flanged), and other socio-ecological factors. Both our longitudinal and cross-sectional results confirm and refine previous cross-sectional accounts of the differences in mating tactics between the unflanged and the flanged male morphs. In the unflanged morph, males exhibit higher sociability, particularly with females, and higher rates of both copulation and sexual coercion than in the flanged morph. Based on our results and those of previous studies showing that females prefer flanged males, and that flanged males have higher reproductive success, we conclude that unflanged males face a trade-off between avoiding male-male contest competition and gaining mating access to females, and thus follow a "best-of-a-bad-job" mating strategy.

Animal conflicts escalate in a warmer world

The potential for climate change to affect animal behaviour is widely recognized, yet its possible consequences on aggressiveness are still unclear. If warming and drought limit the availability of food resources, climate change may elicit an increase of intraspecific conflicts stemming from resource competition. By measuring aggressivity indices in a group-living, herbivorous mammal (the Apennine chamois Rupicapra pyrenaica ornata) in two sites differing in habitat quality, and coupling them with estimates of plant productivity, we investigated whether harsh climatic conditions accumulated during the growing season influenced agonistic contests at feeding via vegetation-mediated effects, and their interaction with the site-specific habitat quality. We focused on females, which exhibit intra-group contest competition to access nutritious food patches. Accounting for confounding variables, we found that (1) the aggression rate between foraging individuals increased with the warming accumulated over previous weeks; (2) the probability to deliver more aggressive behaviour patterns toward contestants increased with decreasing rainfall recorded in previous weeks; (3) the effects of cumulative warming and drought on aggressivity indices occurred at time windows spanning 15–30 days, matching those found on vegetation productivity; (4) the effects of unfavourable climatic conditions via vegetation growth on aggressivity were independent of the site-specific habitat quality. Simulations conducted on our model species predict a ~50 % increase in aggression rate following the warming projected over the next 60 years. Where primary productivity will be impacted by warming and drought, our findings suggest that the anticipated climate change scenarios may trigger bottom-up consequences on intraspecific animal conflicts. This study opens the doors for a better understanding of the multifactorial origin of aggression in group-living foragers, emphasising how the escalation of agonistic contests could emerge as a novel response of animal societies to ongoing global warming.

Patch choice decisions by a fission–fusion forager as a test of the ecological constraints model

Chapman et al.’s (Behav Ecol Sociobiol 36:59–70, 1995) ecological constraints model posits that the size and distribution of food patches place restrictions upon foraging group size. Larger groups incur increased travel costs for any given array of patches, and thus, to fulfil individual energetic and nutritional requirements, foragers should adjust group sizes to balance energy obtained against that spent on travelling. Support for this model comes from both comparative and species-specific studies but findings are contradictory, and the utility of the model has been questioned. This study provides a rigorous test, analysing measurements from distinct food patches and individual inter-patch movements, on an appropriately shorter temporal scale. Using data drawn from two social groups of a species characterised by a high degree of fission–fusion dynamics, the chimpanzee (Pan troglodytes), we show that larger parties foraged in larger food patches and for longer durations, and that larger parties were associated with further travel between patches. Overt contest competition over food increased with party size. We found no evidence of distinct sex differences in either party size or travel distances: the predictive power of forager sex was low compared to that of ecological variables. We propose that analysis at the patch level is more appropriate than a daily averaging approach that may smooth out the very variation being investigated. Our findings suggest that, despite certain limitations, Chapman et al.’s (Behav Ecol Sociobiol 36:59–70, 1995) model of ecological constraints remains a useful tool. Ecology does indeed constrain grouping patterns, and the impact of this is not necessarily differentiated by sex.Significance statementForaging animals face the ‘more mouths to feed’ problem: as the numbers in a group increase, the group must travel further to find enough food, using up energy. Hence, foragers should adjust numbers to minimise these costs, but tests of this idea have proved inconclusive. We investigated the foraging behaviour of chimpanzees, a species with highly flexible grouping, considering their travel between specific patches of food. We found clear support for this proposition, with larger patches of food hosting larger numbers of foragers, and such groups having to travel further to find food. Although it is often thought that female animals should respond more strongly to foraging costs, we found little evidence of sex differences. Our results show that ecology does indeed constrain grouping patterns, and that the impact is felt equally by males and females.

Human third-party observers accurately track fighting skill and vigour along their unique paths to victory

Sexual selection via male-male contest competition has shaped the evolution of agonistic displays, weaponry, and fighting styles, and is further argued to have shaped human psychological mechanisms to detect, process, and respond appropriately to cues of fighting ability. Drawing on the largest fight-specific dataset to date across the sports and biological sciences (N = 2765 fights), we examined how different indicators of fighting ability in humans reflect unique paths to victory and indicate different forms of perceived and actual resource-holding power (RHP). Overall, we discovered that: (1) both striking skill and vigour, and grappling skill and vigour, individually and collectively predict RHP; (2) different RHP indicators are distinguished by a unique path to victory (e.g., striking skill is a knockout-typical strategy, whereas grappling vigour is a submission-typical strategy); and (3) third-party observers accurately track fighting skill and vigour along their unique paths to victory. Our argument that different measures of RHP are associated with unique paths to victory, and third-party observers accurately track fighting vigour and skill along their unique paths to victory, advance our understanding not only of human contest competition, but animal contest theory more broadly.

Coexistence under Hierarchical Resource Exploitation: The Role of the R*-Preemption Trade-Off.

AbstractResource competition theory predicts coexistence and exclusion patterns based on species' R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesized that coexistence under preemption requires an R*-preemption trade-off-that is, the species with the priority access should have a higher R* (lower "efficiency"). Thus, we developed an extension of resource competition theory to investigate partial and total preemption (in the latter, the preemptor is unaffected by species with lower preemption rank). We found that an R*-preemption trade-off is a necessary condition for coexistence in all models. Moreover, under total preemption, the trade-off alone is sufficient for coexistence. In contrast, under partial preemption, more conditions are needed, which restricts the parameter space of coexistence. Finally, we discuss the implications of our finding for seemingly distinct trade-offs, which we view as special cases of the R*-preemption trade-off. These trade-offs include the digger-grazer trade-off, the competition-colonization trade-off, and trade-offs related to light competition between trees and understories.

Sex-specific movement ecology of the shortest-lived tetrapod during the mating season

Sex-specific reproductive strategies are shaped by the distribution of potential mates in space and time. Labord’s chameleon (Furcifer labordi) from southwestern Madagascar is the shortest-lived tetrapod whose life-time mating opportunities are restricted to a few weeks. Given that these chameleons grow to sexual maturity within about three months and that all individuals die soon after breeding, their mating strategies should be adapted to these temporal constraints. The reproductive tactics of this or any other Malagasy chameleon species have not been studied, however. Radio-tracking and observations of 21 females and 18 males revealed that females exhibit high site fidelity, move small cumulative and linear distances, have low corresponding dispersal ratios and small occurrence distributions. In contrast, males moved larger distances in less predictable fashion, resulting in dispersal ratios and occurrence distributions 7–14 times larger than those of females, and males also had greater ranges of their vertical distribution. Despite synchronous hatching, males exhibited substantial inter-individual variation in body mass and snout-vent length that was significantly greater than in females, but apparently unrelated to their spatial tactics. Females mated with up to 6 individually-known mates, but frequent encounters with unmarked individuals indicate that much higher number of matings may be common, as are damaging fights among males. Thus, unlike perennial chameleons, F. labordi males do not seem to maintain and defend territories. Instead, they invest vastly more time and energy into locomotion for their body size than other species. Pronounced variation in key somatic traits may hint at the existence of alternative reproductive tactics, but its causes and consequences require further study. This first preliminary study of the mating system of a Malagasy chameleon indicates that, as in other semelparous tetrapods, accelerated life histories are tied to a mating system with intense contest and scramble competition among males.

Contests with gender bias: evidence from a culinary competition

ABSTRACT Multi-round contest competition determines many important social and economic outcomes such as employment, promotion, political office, as well as singing, debate, and culinary championships. The competitors in such a contest often belong to different groups based on race, ethnicity, and gender. We examine how favouritism in beliefs about ability affects both who wins the contest and the timing of being voted out. The results are tested using novel data from the culinary competition Chopped. We find that female contestants are more likely to be voted out early, but females who survive to the second half of the contest have a good chance to win. These results support an argument of gender bias by the judges.

Substantial but Misunderstood Human Sexual Dimorphism Results Mainly From Sexual Selection on Males and Natural Selection on Females.

Human sexual dimorphism has been widely misunderstood. A large literature has underestimated the effect of differences in body composition and the role of male contest competition for mates. It is often assumed that sexually dimorphic traits reflect a history of sexual selection, but natural selection frequently builds different phenotypes in males and females. The relatively small sex difference in stature (∼7%) and its decrease during human evolution have been widely presumed to indicate decreased male contest competition for mates. However, females likely increased in stature relative to males in order to successfully deliver large-brained neonates through a bipedally-adapted pelvis. Despite the relatively small differences in stature and body mass (∼16%), there are marked sex differences in body composition. Across multiple samples from groups with different nutrition, males typically have 36% more lean body mass, 65% more muscle mass, and 72% more arm muscle than women, yielding parallel sex differences in strength. These sex differences in muscle and strength are comparable to those seen in primates where sexual selection, arising from aggressive male mating competition, has produced high levels of dimorphism. Body fat percentage shows a reverse pattern, with females having ∼1.6 times more than males and depositing that fat in different body regions than males. We argue that these sex differences in adipose arise mainly from natural selection on women to accumulate neurodevelopmental resources.

From Habitat to Hormones: Year-around territorial behavior in rock-dwelling but not in forest and grassland lagomorphs and the role of DHEA

Year-around defense of extremely patchy habitat may require hormones that drive territorial behavior, but have no other costly physiological effects. The American pika (Ochotona princeps), but not the snowshoe hare (Lepus americanus) nor the eastern cottontail rabbit (Sylvilagus floridanus), exhibits this behavior. The former engages in contest competition against all individuals independent of sex to protect its territory in highly fragmented patches on mountain talus slopes; the latter in scramble competition in more continuous forests, grasslands, and shrublands. The hormonal basis for this difference in lagomorphs is unknown. Dehydroepiandrosterone is a prohormone produced by the zona reticularis of the adrenal cortex. It has no effect on aggressive behavior until converted in the brain to estrogen. We assessed levels (DHEA-S plus DHEA) in all species collected in the wild. In nonbreeding pikas, levels were 256 times higher than in hares and 22 times higher than in rabbits. Within species, females and males had similar levels. The proportion of the adrenal cortex devoted to the zona reticularis was significantly larger in pikas than in hares or rabbits. Our evidence is consistent with the hypothesis that dehydroepiandrosterone drives this individual-based, year-around territoriality in pikas. We propose a definitive experiment to determine this and recommend comparative studies in central Asia where there is high diversity of pika species whose behavior ranges from individual-based territoriality to colonial. Thus, we speculate that the wild American pika has the adrenal-brain nexus for all seasons and is an excellent model to understand how habitat drives the hormonal control of spacing behavior.

Think Fast!: Vervet Monkeys Assess the Risk of Being Displaced by a Dominant Competitor When Making Foraging Decisions

Foraging animals need to quickly assess the costs and benefits of different foraging decisions, including resource quantity, quality, preference, ease of access, dispersion, distance, and predation risk. Social animals also need to take social context into account and adapt foraging strategies that maximize net resource intake and minimize contest competition with conspecifics. We used an experimental approach to investigate how social context impacts wild vervet monkey (Chlorocebus pygerythrus) foraging decisions in a multi-destination pentagon array. We baited four platforms with less-preferred corn and one platform with a larger, preferred resource (half banana) that required handling time. We ran over 1,000 trials and found that when monkeys foraged alone, they usually took the path that minimized travel distance but prioritized the preferred-food platform when in competition. However, the foraging strategy chosen by low-ranking individuals depended on the handling skill of the decision maker (i.e., time it would take them to retrieve the banana), the relative rank of their audience members (i.e., who has priority-of-access to resources), and the distance audience members were from the experiment site (i.e., their travel time). When the risk of being displaced by a dominant competitor was low (because they were far away and/or because the decision-maker was skilled in retrieving the banana), low-ranking individuals chose a route that minimized travel costs. Conversely, when the risk of losing food to a dominant competitor was high, decision-makers rushed for the preferred-food platform at the onset of the trial. When the risk of displacement was moderate because a dominant audience member was at least 50 m away, low-ranking individuals partly prioritized the preferred-food platform but took the time to stop for one platform of corn on the way. This strategy increased the total amount of food obtained during the trial. These findings suggest that lower-ranking individuals, who experienced high contest competition at the foraging experiment, calculated the risk of being displaced by a dominant competitor when making foraging decisions. This experiment demonstrates that vervets go through a complex decision-making process that simultaneously considers the profitability of different foraging decisions and their social context.