Scramble Competition Research Articles

Drivers of a temporal change in the adult sex ratio of a Roosevelt elk (Cervus canadensis roosevelti) population

Abstract Ecological processes driving female-skewed adult sex ratios (ASRs; males:females) in populations with polygynous mating systems have been addressed theoretically, but empirical support is scarce. The theoretical framework of the female substitution hypothesis (FSH) asserts that a female-skewed ASR at carrying capacity reflects an overall fitness benefit for females and for males competitive in acquiring access to reproductive females. The FSH predicts that as population abundance increases females should acquire forage more efficiently than males, thereby leading to passive displacement of males. The result is declining ASR associated with differential habitat use by both sexes as food resources are depleted by female scramble competition. We characterized the temporal variation in ASR in a Roosevelt elk population inhabiting the Redwood National and State Parks, California, across 24 years, and determined which of two possible ecological mechanisms was the driver of a declining ASR. The first mechanism explored was that increasing female abundance associated with declining forage in the study area led to the passive displacement of males into the study periphery over time. The second mechanism explored was that a declining ASR was precipitated by a lack of males within the study area and the study periphery. Systematic population surveys from a vehicle were done to estimate abundance and ASR as well as assess male abundance in the study periphery. Forage biomass was estimated in quarter-m2 plots randomly placed in meadows inhabited by female elk. Our multiple regression analysis revealed an inverse relationship between abundance and ASR indicating density dependence. We found numerous males in the study periphery when females were abundant. Our least squares models indicated declining food resources across years when female abundance increased. Our results showed that the first, and not the second, ecological mechanism examined was responsible for a female-skewed ASR. Our findings provide empirical support for the theoretical framework of the FSH in a nonmigratory population protected from hunting.

Effects of operational sex ratio and male density on size-dependent mating in Minshan’s toads, Bufo minshanicus, on the Tibetan Plateau of China

Abstract In many animal species, an increase in the operational sex ratio (OSR), density or a combination of both should lead to more intensive competition among individuals of the more abundant sex. To test this, we examined pairing patterns of Minshan’s toad (Bufo minshanicus) from six populations between 2008 and 2015 along the eastern Tibetan Plateau in south-west China. OSRs in breeding aggregations of Minshan’s toad are normally male biased and males actively compete with each other for acquisition and retention of mates. We found evidence that deviations from random mating by size varied between populations and between years according to the magnitude of the OSR and male density. Larger males were generally more successful in pairing than smaller males when the OSR was slightly male biased and male density was high. However, the resulting size-disproportionate mating was more evident when OSR was closer to 1.99, indicating a positive correlation with the intensity of aggressive scramble competition. Thus, the intensity of male-male competition may partly explain variation in size-disproportionate mating among populations.

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.

Larger is not better: no mate preference by European common frog (Rana temporaria) males

Abstract In explosive breeding frogs, high intrasexual competition between males leads to a sexual coercion ruled mating system, where males presumably evolved preferences for specific female traits. We tested these preferences in the European Common Frog by excluding intrasexual competition. We hypothesized that all males show preferences towards larger female body size, due to higher fecundity. Our results did not show any preference considering female body size, neither in the attempt to amplex a female nor during the formation of pairs. Additionally, we witnessed a high failure rate of male mating attempts, which hints at high mating costs and offers an explanation for the lack of preferences in males. Nonetheless, we observed a non-random mating pattern in successfully formed pairs, where in the absence of size dimorphism females were on average larger than males. This indicates a different mechanism for selection which is independent from male mating preference or scramble competition.

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.

Social drivers of maturation age in female geladas.

Female reproductive maturation is a critical life-history milestone, initiating an individual's reproductive career. Studies in social mammals have often focused on how variables related to nutrition influence maturation age in females. However, parallel investigations have identified conspicuous male-mediated effects in which female maturation is sensitive to the presence and relatedness of males. Here, we evaluated whether the more "classic" socioecological variables (i.e., maternal rank, group size) predict maturation age in wild geladas-a primate species with known male-mediated effects on maturation and a grassy diet that is not expected to generate intense female competition. Females delayed maturation in the presence of their fathers and quickly matured when unrelated, dominant males arrived. Controlling for these male effects, however, higher-ranking daughters matured at earlier ages than lower-ranking daughters, suggesting an effect of within-group contest competition. However, contrary to predictions related to within-group scramble competition, females matured earliest in larger groups. We attribute this result to either: 1) a shift to "faster" development in response to the high infant mortality risk posed by larger groups; or 2) accelerated maturation triggered by brief, unobserved male visits. While earlier ages at maturation were indeed associated with earlier ages at first birth, these benefits were occasionally offset by male takeovers, which can delay successful reproduction via spontaneous abortion. In sum, rank-related effects on reproduction can still occur even when socioecological theory would predict otherwise, and males (and the risks they pose) may prompt female maturation even outside of successful takeovers.

Land‐use alters the form of larval density dependence to increase extinction risk in a grassland amphibian

AbstractAn understanding of interactions between anthropogenic stressors and intrinsic population drivers is needed to fully understand wildlife population declines. Density dependence is a key aspect of population regulation for many species, especially for species that have high reproductive potential, such as amphibians. However, patterns of density dependence have been characterized for only a few species and little work has evaluated how density‐dependent interactions may be altered by anthropogenic stressors. We combined the results of a mesocosm experiment with demographic population modeling to investigate how the conversion of native prairie to agricultural grasslands dominated by Tall Fescue grass (Lolium arundinacea) affected larval density dependence and adult population size of an imperiled amphibian, Lithobates areolatus (Crawfish Frog). Overall, density dependence was overcompensatory, suggesting that L. areolatus exhibits scramble competition as larvae. Both vegetation treatments had low survival at high densities, but more individuals survived to metamorphosis at moderate densities in Fescue treatments compared to Prairie treatments. We evaluated the implications of our experimental results using a stochastic density‐dependent matrix population model to project long‐term population dynamics. Simulated populations breeding in Fescue‐dominated wetlands had a more variable population size and up to 400% higher probability of quasi‐extinction within 200 years, compared to populations breeding in ponds with prairie vegetation. Without varying density in experimental treatments and using mathematical models to project emergent population dynamics, our mesocosm experiment results would have suggested a slightly positive effect of Fescue grass on amphibian development and survival. Vegetation changes surrounding breeding wetlands might play an important role in the decline of amphibian populations persisting in low‐intensity agricultural areas.

Multi-modal locomotor costs favor smaller males in a sexually dimorphic leaf-mimicking insect

BackgroundIn most arthropods, adult females are larger than males, and male competition is a race to quickly locate and mate with scattered females (scramble competition polygyny). Variation in body size among males may confer advantages that depend on context. Smaller males may be favored due to more efficient locomotion leading to higher mobility during mate searching. Alternatively, larger males may benefit from increased speed and higher survivorship. While the relationship between male body size and mobility has been investigated in several systems, how different aspects of male body morphology specifically affect their locomotor performance in different contexts is often unclear.ResultsUsing a combination of empirical measures of flight performance and modelling of body aerodynamics, we show that large body size impairs flight performance in male leaf insects (Phyllium philippinicum), a species where relatively small and skinny males fly through the canopy in search of large sedentary females. Smaller males were more agile in the air and ascended more rapidly during flight. Our models further predicted that variation in body shape would affect body lift and drag but suggested that flight costs may not explain the evolution of strong sexual dimorphism in body shape in this species. Finally, empirical measurements of substrate adhesion and subsequent modelling of landing impact forces suggested that smaller males had a lower risk of detaching from the substrates on which they walk and land.ConclusionsBy showing that male body size impairs their flight and substrate adhesion performance, we provide support to the hypothesis that smaller scrambling males benefit from an increased locomotor performance and shed light on the evolution of sexual dimorphism in scramble competition mating systems.

Size variation, allometry and mating success in Aotearoa|New Zealand kelp flies (Coelopidae)

The need to respond quickly to the presence of an ephemeral resource required for breeding is often a feature of scramble competition mating systems. Scramble competition mating systems can feature extreme levels of sexual conflict and coercive mating by males. As a result, sexual selection can act on various traits used by males to overcome female resistance behaviours. Selection on these traits may result in significant intra and intersexual size variation and sexual dimorphism. Additionally, traits that influence mating success in males often show positive static allometry. Kelp flies (Coelopidae) are a small family of Diptera which specialise on wrack (beach cast marine macroalgae), a highly ephemeral resource. The mating system of these flies involves high levels of sexual conflict, with females rejecting all male mating attempts. In this study we describe intra and intersexual size variation and static allometry of traits in two of Aotearoa|New Zealand’s species, Coelopella curvipes and Chaetocoelopa littoralis. In addition, we investigate the mating behaviour of C. littoralis under ecologically relevant mating conditions. We found high levels of variation in both species with significant evidence of sexual dimorphism across all traits measured in C. littoralis, and in mid tibia length in C. curvipes. Furthermore, mid tibia length in both species exhibits positive static allometry and is disproportionally larger in larger males, suggesting that this trait in particular may be under strong sexual selection. We found that larger male C. littoralis which attempt to mate are significantly more likely to mate successfully demonstrating a large-size advantage in this species similar to findings across the Coelopidae. However, we only found a non-significant trend towards a mating advantage for males with longer mid-tibiae. We discuss these findings with reference to the population dynamics and ecology of these species.

To harass or to respect: the economy of male persistence despite female refusal in a damselfly with scramble mate competition

In sexual conflict, males are often thought to gain fitness benefits from harassing females over mating. Yet when harassment itself incurs costs to males and if alternative, receptive females are available in a local population, theory predicts that when confronted with a female refusal, a male’s choice of persisting or retreating is determined in part by the likelihood of achieving a mating. We tested that prediction in the damselfly Enallagma hageni, whose males compete by intense scramble competition, resulting in widespread mating harassment toward females, which have a high level of control over mating. Using captive individuals of E. hageni in outdoor insectaries, we quantified male persistence in mating after refusals by pre- and post-oviposition focal females whose egg content we quantified after observations. We documented a novel, context-dependent head-turning refusal signal of sexual non-receptivity, most often displayed in tandem pairs by post-oviposition females that typically carried few mature eggs for males to fertilize. Male persistence was less likely to result in mating with post-oviposition females compared with pre-oviposition females carrying a clutch of mature eggs. Accordingly, males were less likely to persist following refusal signals given by post-oviposition females, supporting the theoretical prediction. Compared with a refusal signal known as wing spread, head-turning was significantly more effective in deterring harassing males. Our results suggest that despite on-going sexual conflict over mating, cooperation benefits both sexes when females use the honest signal of non-receptivity because they carry few mature eggs that males could fertilize.

Harmonic convergence coordinates swarm mating by enhancing mate detection in the malaria mosquito Anopheles gambiae

The mosquito Anopheles gambiae is a major African malaria vector, transmitting parasites responsible for significant mortality and disease burden. Although flight acoustics are essential to mosquito mating and present promising alternatives to insecticide-based vector control strategies, there is limited data on mosquito flight tones during swarming. Here, for the first time, we present detailed analyses of free-flying male and female An. gambiae flight tones and their harmonization (harmonic convergence) over a complete swarm sequence. Audio analysis of single-sex swarms showed synchronized elevation of male and female flight tones during swarming. Analysis of mixed-sex swarms revealed additional 50 Hz increases in male and female flight tones due to mating activity. Furthermore, harmonic differences between male and female swarm tones in mixed-sex swarms and in single-sex male swarms with artificial female swarm audio playback indicate that frequency differences of approximately 50 Hz or less at the male second and female third harmonics (M2:F3) are maintained both before and during mating interactions. This harmonization likely coordinates male scramble competition by maintaining ideal acoustic recognition within mating pairs while acoustically masking phonotactic responses of nearby swarming males to mating females. These findings advance our knowledge of mosquito swarm acoustics and provide vital information for reproductive control strategies.

A Comparative Analysis of Adult Sex Ratios in Polygynous and Monogamous Mammal Populations

An inverse density-dependent relationship between abundance and adult sex ratio (ASR, males:female) occurs in some populations of polygynous mammals due to life history differences between the sexes. Male fecundity and survival is dictated by attempts to obtain as many copulations as possible, whereas female fecundity and survival is dictated by resource acquisition. Therefore, females usually acquire forage before males as a result of interspecific scramble competition, particularly when forage becomes more limited at K carrying capacity. This leads to the passive displacement of males in a given area. The common belief is that most monogamous mammal populations exhibit balanced adult sex ratios. The coupling of sexually mature males and females in a population result in this pattern for this mating system. Present literature focuses on primary or secondary sex ratios in mammals or on ASR patterns within individual species. Our goal was to test if expected ASR patterns would be visible across numerous species in both mating systems. We hypothesized we would see an inverse relationship between abundance and ASR across polygynous populations, and no relationship between abundance and ASR across monogamous populations. We extracted time series population data from published literature for 43 populations of 15 different mammal species. Results from our analysis of a linear mixed-effects model were consistent with our hypothesis for polygynous populations, as we found a significant inverse relationship between abundance and ASR. However, our analysis also revealed a significant inverse relationship between abundance and ASR in monogamous populations that was not consistent with our hypothesis. Our findings provide quantitative support for a theoretical model explaining the evolutionary and ecological mechanisms driving a density-dependent relationship between abundance and ASR in polygynous mammals. An investigation into monogamous mammals is needed to assess why some species with this mating system display a density-dependent response in ASR as well.

Interaction between sexual communication functions leads to reproductive interference in two syntopic scorpion species

Sexual communication in mate choice scenarios involves nonmutually exclusive functions: species recognition, mate location and mate assessment. Most studies on sexual communication in mate choice have focused on female assessment of conspicuous male traits, such as visual and acoustic signals. Only a few studies have addressed male assessment of female traits, especially those that are inconspicuous to the human sensory system, such as chemical signals. Although chemical signals are important in sexual communication of many arachnids, there is a gap in our knowledge about how male scorpions use these signals to evaluate potential partners. Here we studied sexual communication functions in two scorpion species (Urophonius brachycentrus and Urophonius achalensis) with sympatric populations and synchronic reproduction, where males are under a scramble competition mating system. We addressed the volatile pheromones role in each of the sexual communication functions. We used Y-mazes to test volatile pheromones, exposing males to different female stimulus arrays in single-signal (conspecific only) and mixed-signal (conspecific and heterospecific) contexts. Males located conspecific females through volatile pheromones, and the time spent in proximity to the female was proportional to males' perception of the female's quality, reinforcing the pheromones' function in mate location and assessment in single-signal environments. However, these sexually selected functions were overridden in mixed-signal contexts, in which males could not differentiate conspecific from heterospecific females, suggesting that pheromones do not allow species recognition. This interaction between sexual communication functions can lead to reproductive interference in this sympatric zone. Male indiscriminateness mediated by scramble competition could attenuate pheromones' species recognition function during mate choice. We provide a comprehensive study of the pheromones' role for mate and species recognition in different social contexts in scorpions. We found support for nonindependent and interacting functions of sexual communication, indicating that mate choice is a complex process in scorpions.

Differential allocation in a gift-giving spider: males adjust their reproductive investment in response to female condition

BackgroundWhen males are selective, they can either reject low-quality females or adjust their reproductive investment in response to traits that indicate female quality (e.g., body size or condition). According to the differential allocation hypothesis, males increase their reproductive investment when paired with high-quality females (positive differential allocation) or increase their reproductive investment when paired with low-quality females (negative differential allocation). This hypothesis has been proposed for monogamous species with biparental care, and most empirical studies focus on birds. Here we used the polygamous spider Paratrechalea ornata, in which males offer prey wrapped in silk as nuptial gifts, to test whether males adjust their reproductive investment in gift size, pre-copulatory and copulatory courtship, and sperm transfer in response to female body condition.ResultsMales exposed to females in good body condition added more flies to the gift, stimulated these females longer with abdominal touches during pre-copulatory courtship, and had longer pedipalp insertions than males exposed to females in poor body condition. Female condition affected neither silk investment in nuptial gift wrapping nor the quantity of sperm transferred by males. Finally, females in good body condition oviposited faster after copulation and laid more eggs than females in poor body condition.ConclusionsWe provide experimental evidence that males of a gift-giving spider exhibit positive differential allocation in three key aspects of their reproductive investment: the size of the nutritious gift, duration of pre-copulatory courtship, and duration of pedipalp insertions, which is regarded as a form of copulatory courtship in spiders. This positive differential allocation is likely associated with the benefits of copulating with females in good body condition. These females are more fecund and oviposit faster after copulation than females in poor body condition, which under natural field conditions probably reduces the risk of multiple matings and thus the level of sperm competition faced by the males. As a final remark, our findings indicate that the hypothesis of differential allocation also applies to species with a scramble competition mating system, in which males heavily invest in nuptial gift construction, but not in parental care.

Males of a Neotropical spider adjust prey‐gift construction but not mate search in response to sperm competition

AbstractIn scramble competition mating systems, males do not guard females and females usually mate polyandrously, leading to high levels of sperm competition. Mathematical models predict that males facing sperm competition can either decrease or increase their reproductive effort. To test how male reproductive responses vary according to sperm competition risk, we ran two experiments using the polyandrous gift‐giving spiderParatrechalea ornata. First, we tested whether males prefer substrates containing silk of unmated females than substrates containing silk of once‐mated females, competitor males or without silk. Males spent more time on substrates containing silk of females but showed no preference for unmated females, which indicates that males may optimize mate search by prioritizing substrates containing cues of potential mating partners. The lack of preference for unmated females suggests that males cannot detect female mating status from silk or that the costs of searching preferentially for unmated females are high in the field. Second, we tested whether males adjust prey‐gift construction when facing sperm competition risk. Experimental groups (substrate containing silk of an unmated female and silk of another malevs. substrate containing only silk of an unmated female) did not differ in the frequency of gift construction, latency to gift construction and number of flies added to the gift. However, males facing sperm competition risk constructed gifts with less silk, suggesting the occurrence of a less evident form of male mate choice based on differential allocation of resources to prey‐gift. Additionally, given the trade‐off between investing in prey‐gifts quantity or quality, a lower silk investment should enable males to increase the number of gifts constructed and the number of courted females. Taken together, our results indicate that males detect female and male pheromones on substrate and adjust the construction of prey‐gifts in response to apparent sperm competition risk.

A Ricker–type predator–prey system with hunting cooperation in discrete time

A predator–prey system in discrete-time for which the predators engage in cooperative hunting and the prey population is subject to scramble intra-specific competition is developed and analyzed. We prove that the number of positive equilibrium points depends not only on the predator’s basic reproduction number but also on the intensity of predator cooperation. The model exhibits a discrete Hopf bifurcation when the unique positive equilibrium point loses its stability. Two-parameter bifurcation diagrams using MATLAB conclude that hunting cooperation can increase the likelihood of predator persistence but may destabilize the predator–prey interaction and drive the predator to extinction.

No risk to scrambling? Mating tactic does not affect the frequency of leg autotomy in a New Zealand harvestman

Intense sexual selection on males may drive the evolution of exaggerated weaponry, typically used in contests for females or reproductive sites. In some species, males have discontinuous variation in weapon morphology that is accompanied by alternative reproductive tactics (ARTs). Major males with enlarged weapons usually exhibit a mating tactic based on female or resource defence, which makes them relatively sedentary. Minor males with reduced or absent weapons may exhibit a scramble competition mating tactic, which involves increased movement in search of females. Thus, the main costs paid by majors involve the expression/maintenance of exaggerated structures and potential injuries during contests. In turn, the main costs paid by minors are likely related to increased exposure to predation during mate search, yet this cost has rarely been considered for minors. Harvestmen are an arachnid group with diverse male weaponry, including many cases of male polymorphism associated with ARTs. Species of the suborder Eupnoi use leg autotomy as a common antipredator defence. Here we investigated whether leg autotomy (a proxy for predation risk) is more frequent in minors of the weapon polymorphic harvestman Forsteropsalis pureora , which has three male morphs with drastically different cheliceral size and morphology. Leg autotomy was very common, with 54% of wild-caught individuals missing at least one leg (mostly leg II), but we found no evidence for differential predation risk between male morphs during adulthood. In a predator simulation experiment, we found no difference in the likelihood or latency to autotomize a leg between male morphs or leg types (legs I, II, III or IV). However, males already missing legs were more reluctant to autotomize an additional leg. Our results suggest that while leg autotomy is a common antipredator strategy, with no difference between male tactics, costs are compounded as additional legs are autotomized and males may strategically decide not to autotomize in future predator encounters. • New Zealand harvestmen autotomize (lose) legs to escape predators. • This defence strategy is common in wild populations. • Sensory legs are lost more often than locomotory legs. • Harvestmen that have lost legs in the past take longer to drop an additional leg. • Male reproductive strategy does not affect autotomy risk.

Dissecting the two mechanisms of scramble competition among the Virunga mountain gorillas

Two mechanisms have been proposed to explain why scramble competition can increase the travel requirements of individuals within larger groups. Firstly, individuals in larger groups may be more likely to encounter food sites where other group members have already eaten, leading to greater asynchronous “individual” travel to find fresh sites. Secondly, when food sites are aggregated into patches, larger groups may need to visit more patches to obtain the same amount of food per capita, leading to greater synchronous “group” travel between patches. If the first mechanism can be mitigated by increasing group spread, then we expect the second mechanism to be more sensitive to group size. Here, we examine the individual travel and group travel of the Virunga mountain gorillas, along with potential implications for the two mechanisms of scramble competition. Asynchronous individual travel accounted for 67% of the total travel time, and the remainder arose from group travel. Group spread increased significantly for larger groups, but not enough to prevent an increase in individual travel. Contrary to expectations, group travel decreased with size among most groups, and we found only limited evidence of patch depletion that would cause the second mechanism of scramble competition. Collectively, our results illustrate how the influence of group size can differ for individual travel versus group travel, just as it differs among species for overall travel. Studies that distinguish between the two mechanisms of scramble competition may enhance our understanding of ecological constraints upon group size, including potential differences between frugivores and folivores.Significance statementFeeding competition provides insight into how group size can influence the foraging patterns of social animals, but two key mechanisms are not typically compared. Firstly, larger groups may visit more patches to access the same amount of food per capita (group travel). Secondly, their individuals may also need to move past more spots where another member has already eaten (individual travel). Contrary to expectations, we found that group travel decreased with size for most groups of mountain gorillas, which may reflect extra travel by smaller groups to avoid larger groups. Individual travel increased with size in most groups, even though gorillas in larger groups compensated by spreading out over a broader area. The two mechanisms revealed patterns that were not apparent in our previous study of overall travel. Our approach may help to explain potential differences between folivores and frugivores.

Community-level consequences of harsh environmental constraints based on spatial patterns analysis in karst primary forest of southwest China

Karst region has a high rate of rock exposure, shallow and discontinuous soil, and a fragile ecological environment. However, these special habitats encompass a large area of forest with strong original nature-karst deciduous broad-leaved and evergreen mixed forest. So far, very limited information is available on the spatial patterns and internal maintenance mechanism of this community. To fill this gap, the spatial distributions together with spatial correlations of both the live and dead trees in the karst primary forests of the Maolan National Nature Reserve in China were explored by using univariate as well as bivariate g(r) functions. The results revealed that: (1) the habitat heterogeneity of the karst primary forest has insignificant influence at a larger scale (>10 m); (2) the decline in aggregation with the increasing size classes from saplings to adults suggests self-thinning process, proving that negative density constraints occurred in early life stages; (3) positive spatial correlation was found in live trees with similar diameters, which related to the microsite heterogeneity; (4) the aggregation of dead trees and non-attraction of dead and surviving trees indicated scramble competition; (5) mortality analysis indicated the absence of random mortality, favoring positive density-dependent effects; (6) strong initiation sprouting regeneration was one of the main self-renewal strategies, and regeneration was more likely to occur at the edge of the gap rather than under the gap. Our results support the view that the mechanisms underlying the observed spatial patterns were related to certain ecological processes and their interaction. These were the positive and negative density dependence, scramble competition, and microsite heterogeneity.

Plasticity of Behavioral and Growth Responses to Two Feeding Regimes and Implications for the Domestication of Steelhead Trout

AbstractThe effects of domestication selection on fitness are of serious concern in hatchery‐reared anadromous salmonids, which often experience selection for rapid growth and associated deviations from natural agonistic and feeding behaviors. The high rations that are used to raise age‐1 smolts and the reduced rations that are used to produce age‐2 smolts potentially result in very different behavioral tactics and social dynamics of competition in hatchery tanks. Consequently, steelhead Oncorhynchus mykiss hatchery programs with recovery or conservation goals may need to change their aquaculture practices to minimize domestication selection. This study investigated how the feeding rate (proportion of presented food items that were successfully captured) and relative competitive ability (ability to monopolize food items in small groups) of age‐1 and age‐2 steelhead corresponded to feeding location (surface or bottom) and growth rate in hatchery rearing tanks. We raised steelhead from a common broodstock to smolt at age 1 or age 2 by manipulating their rations (age‐1 fish received the high ration, age‐2 the low ration). The fish in the age‐1 treatment had a lower feeding rate than did those in the age‐2 treatment. Feeding rate was positively correlated with both body size and with growth rate in the rearing tanks at both rations. The age‐2 steelhead were more likely to be detected at a single feeding location than were the age‐1 fish, which were more frequently detected at both locations. The more competitive age‐1 fish were more frequently detected at both feeding locations, and the less competitive fish were detected at only one location. Relative competitive ability was not associated with feeding location in the age‐2 treatment, nor was it associated with growth rate in either treatment. Age‐2 steelhead that were detected only at the bottom or at both feeding locations had higher growth rates than those that were detected only at the surface, suggesting greater scramble competition near the point of food delivery under the restricted ration. Feeding location was not associated with growth rate in the age‐1 treatment. The results indicate that ration directly affects feeding rate and that ration and relative competitive ability interact to influence feeding rate. Because feeding rate was positively associated with growth rate at both ration levels in the hatchery tanks, behaviors that are associated with feeding motivation may be an important trait under selection in salmonid hatcheries.