Population dynamical consequences of gregariousness in a size-structured consumer-resource interaction

Many animal species live in groups. Group living may increase exploitation competition within the group, and variation among groups in intra-group competition intensity could induce life-history variability among groups. Models of physiologically structured populations generally predict single generation cycles, driven by exploitation competition within and between generations. We expect that life-history variability and habitat heterogeneity induced by group living may affect such competition-driven population dynamics. In this study, we vary the gregariousness (the tendency to aggregate in groups) of a size-structured consumer population in a spatially explicit environment. The consumer has limited mobility, and moves according to a probabilistic movement process. We study the effects on the population dynamics, as mediated through the resource and the life-history of the consumer. We find that high gregariousness leads to large spatial resource variation, and highly variable individual life-history, resulting in highly stochastic population dynamics. At reduced gregariousness, life-history of consumers synchronizes, habitat heterogeneity is reduced, and single generation cycles appear. We expect this pattern to occur for any group living organism with limited mobility. Our results indicate that constraints set by population dynamical feedback may be an important aspect in understanding group living in nature.     

Population assessment without individual identification using camera-traps: A comparison of four methods

The use of camera traps to estimate population size when animals are not individually recognizable is gaining traction in the ecological literature, because of its applicability in population conservation and management.We estimated population size of synthetic animals with four camera trap sampling-based statistical models that do not rely on individual recognition. Using a realistic model of animal movement to generate synthetic data, we compared the random encounter model, the random encounter and staying time model, the association model and the time-to-event-model and we investigated the impact of violation of assumptions on the population size estimates.While under ideal conditions these models provide reliable population estimates, when synthetic animal movements were characterised by differences in speed (due to diverse behaviours such as locomotion, grazing and resting) none of the model provided both unbiased and precise density estimates. The random encounter model and the time-to-event-model provided precise results but tended to overestimate population size, while the random encounter and staying time model was less precise and tended to underestimate population size. Lastly, the association model was unable to provide precise results. We found that each tested model was very sensitive to the method used to estimate the range of the field-of-view of camera traps. Density estimates from both random encounter model and time-to-event-model were also very sensitive to biases in the estimate of animals’ speed. We provide guidelines on how to use these statistical models to get population size estimates that could be useful to wildlife managers and practitioners.     

The Effect of Animal Movement on Line Transect Estimates of Abundance

Line transect sampling is a distance sampling method for estimating the abundance of wild animal populations. One key assumption of this method is that all animals are detected at their initial location. Animal movement independent of the transect and observer can thus cause substantial bias. We present an analytic expression for this bias when detection within the transect is certain (strip transect sampling) and use simulation to quantify bias when detection falls off with distance from the line (line transect sampling). We also explore the non-linear relationship between bias, detection, and animal movement by varying detectability and movement type. We consider animals that move in randomly orientated straight lines, which provides an upper bound on bias, and animals that are constrained to a home range of random radius. We find that bias is reduced when animal movement is constrained, and bias is considerably smaller in line transect sampling than strip transect sampling provided that mean animal speed is less than observer speed. By contrast, when mean animal speed exceeds observer speed the bias in line transect sampling becomes comparable with, and may exceed, that of strip transect sampling. Bias from independent animal movement is reduced by the observer searching further perpendicular to the transect, searching a shorter distance ahead and by ignoring animals that may overtake the observer from behind. However, when animals move in response to the observer, the standard practice of searching further ahead should continue as the bias from responsive movement is often greater than that from independent movement.     

动物生殖抑制研究进展：理论模型、方式和机制

生殖抑制指原本具有生育能力的动物个体因特定外界环境或生理条件而减少或丧失生殖能力的现象,有时是受环境变化的主动调控,更多的是出现在其他个体影响下的被动抑制,极端情况发生在社会性动物的永久性抑制,即永久无法生殖或无法生殖成熟。研究发现非社会性动物也有生殖的推迟及可恢复性的生殖抑制,生殖抑制影响着动物种群数量动态、维持和进化。随着多学科的发展,生殖抑制机理研究取得了诸多进展。从阐述生殖抑制的概念出发,解析生殖抑制的形态、激素和分子生理特征,总结了生殖抑制的原因、作用,终述现有的理论模型以及不同物种方面的最新进展,并就生殖抑制在生物资源管理方面的应用价值进行了展望,旨在丰富生殖抑制的理论,扩展应用实践,为后续的生物资源管理提供理论参考。     

The effects of density, spatial pattern, and competitive symmetry on size variation in simulated plant populations

Patterns of size inequality in crowded plant populations are often taken to be indicative of the degree of size asymmetry of competition, but recent research suggests that some of the patterns attributed to size-asymmetric competition could be due to spatial structure. To investigate the theoretical relationships between plant density, spatial pattern, and competitive size asymmetry in determining size variation in crowded plant populations, we developed a spatially explicit, individual-based plant competition model based on overlapping zones of influence. The zone of influence of each plant is modeled as a circle, growing in two dimensions, and is allometrically related to plant biomass. The area of the circle represents resources potentially available to the plant, and plants compete for resources in areas in which they overlap. The size asymmetry of competition is reflected in the rules for dividing up the overlapping areas. Theoretical plant populations were grown in random and in perfectly uniform spatial patterns at four densities under size-asymmetric and size-symmetric competition. Both spatial pattern and size asymmetry contributed to size variation, but their relative importance varied greatly over density and over time. Early in stand development, spatial pattern was more important than the symmetry of competition in determining the degree of size variation within the population, but after plants grew and competition intensified, the size asymmetry of competition became a much more important source of size variation. Size variability was slightly higher at higher densities when competition was symmetric and plants were distributed nonuniformly in space. In a uniform spatial pattern, size variation increased with density only when competition was size asymmetric. Our results suggest that when competition is size asymmetric and intense, it will be more important in generating size variation than is local variation in density. Our results and the available data are consistent with the hypothesis that high levels of size inequality commonly observed within crowded plant populations are largely due to size-asymmetric competition, not to variation in local density.     

Effect of immersion at low tide on distribution and movement of the mud snail, Ilyanassa obsoleta (Say), in the upper Bay of Fundy, eastern Canada

Habitat heterogeneity often affects movement behaviours of animals, and consequently their spatial distribution. We evaluated the effect of immersion at low tide on the distribution, fine-scale movement patterns and daily movement patterns of the mud snail Ilyanassa obsoleta on a mudflat in the upper Bay of Fundy, Canada. Mud snails migrate onto intertidal mudflats in the summer, and our field survey showed that their density was higher inside tide pools relative to adjacent areas that are exposed at low tide. Using time-lapse videography, we evaluated the effect of snail size, snail density, and immersion at low tide on fine-scale movement patterns of I. obsoleta. Time until snails stopped moving and burrowed was unaffected by snail size, but snails at low and high densities burrowed somewhat faster than those at intermediate densities. Snail size and snail density had no detectable effect on displacement speed or linearity of displacement. Immersion affected snail movement: snails within tide pools delayed burrowing and traveled in more convoluted paths compared to those on exposed mud. Snails increased their turning angles within tide pools, which is probably the mechanism by which aggregations are formed. We also performed a mark-recapture experiment to compare daily movement patterns of snails released inside and outside tide pools. Snails released in tide pools moved shorter distances, but did not orient themselves differently than snails released outside tide pools. Both groups exhibited significant directionality, moving against the mean water current direction over 24 h. In sum, immersion at low tide affected the behaviour and spatial distribution of snails, resulting in snail aggregations within tide pools. These snail aggregations, in turn, may be a major factor influencing spatial dynamics on mudflats, including causing changes in distribution patterns of the burrowing amphipod Corophium volutator, a dominant inhabitant and key species in the food web of mudflats.     

Local Competition Between Foraging Relatives: Growth and Survival of Bruchid Beetle Larvae

Kin selection theory states that when resources are limited and all else is equal, individuals will direct competition away from kin. However, when competition between relatives is completely local, as is the case in granivorous insects whose larval stages spend their lives within a single seed, this can reduce or even negate the kin-selected benefits. Instead, an increase in competition may have the same detrimental effects on individuals that forage with kin as those that forage with non-kin. In a factorial experiment we assessed the effects of relatedness and competition over food on the survival and on fitness-related traits of the bruchid beetle Callosobruchus maculatus. Relatedness of competitors did not affect the survival of larvae. Larval survival substantially decreased with increasing larval density, and we found evidence that beetles maturing at a larger size were more adversely affected by competition, resulting in lower survival rates. Furthermore, females showed a reduction in their growth rate with increasing larval density, emerging smaller after the same development time. Males increased their growth rate, emerging earlier but at a similar size when food was more limited. Our results add to the growing number of studies that fail to show a relationship between relatedness and a reduction in competition between relatives in closed systems, and emphasize the importance of the scale at which competition between relatives occurs.     

Rate of gene transfer from mitochondria to nucleus: effects of cytoplasmic inheritance system and intensity of intracellular competition

Endosymbiotic theory states that mitochondria originated as bacterial intracellular symbionts, the size of the mitochondrial genome gradually reducing over a long period owing to, among other things, gene transfer from the mitochondria to the nucleus. Such gene transfer was observed in more genes in animals than in plants, implying a higher transfer rate of animals. The evolution of gene transfer may have been affected by an intensity of intracellular competition among organelle strains and the organelle inheritance system of the organism concerned. This article reveals a relationship between those factors and the gene transfer rate from organelle to nuclear genomes, using a mathematical model. Mutant mitochondria that lose a certain gene by deletion are considered to replicate more rapidly than normal ones, resulting in an advantage in intracellular competition. If the competition is intense, heteroplasmic individuals possessing both types of mitochondria change to homoplasmic individuals including mutant mitochondria only, with high probability. According to the mathematical model, it was revealed that the rate of gene transfer from mitochondria to the nucleus can be affected by three factors, the intensity of intracellular competition, the probability of paternal organelle transmission, and the effective population size. The gene transfer rate tends to increase with decreasing intracellular competition, increasing paternal organelle transmission, and decreasing effective population size. Intense intracellular competition tends to suppress gene transfer because it is likely to exclude mutant mitochondria that lose the essential gene due to the production of lethal individuals.     

Modelling space use and dispersal of mammals in real landscapes: a tool for conservation

Aim To explore the usefulness of Spatially Explicit Population Models (SEPMs), incorporating dispersal, as tools for animal conservation, as illustrated by the contrasting cases of four British mammals. Methods For each of the four species (American mink, Mustela vison, pine marten, Martes martes, dormouse, Muscardinus avellanarius and water vole, Arvicola terrestris) a spatial dynamics model was developed based on an integrated geographical information system (GIS) population model that linked space use to the incidence of the species. Each model had, first, a GIS, which stored environmental, habitat and animal population information, and secondly, an individual‐based population dynamics module, which simulated home range formation, individual life histories and dispersal within the GIS‐held landscape. Results The four models illustrated different interactions between species life‐history variables and the landscape, particularly with respect to dispersal. As water voles and dormice occupy home ranges that are small relative to blocks of their habitat, they were most effectively modelled in terms of the dynamics of local populations within habitat blocks but linked by dispersal. In contrast, because the home ranges of American mink and pine marten are large relative to blocks of habitat, they were best modelled as individuals moving through a landscape of more or less useful patches of habitat. For the water vole, the most significant predictors of population size were the carrying capacity of each habitat and the annual number of litters. For the dormouse, the likelihood of catastrophe and the upper limit to dispersal movement were the key variables determining persistence. Adult mortality and home‐range size were the only significant partial correlates of total population size for the American mink. Adult mortality was also a significant correlate of total population size in the pine marten, as were litter size and juvenile mortality. In neither the marten nor the mink was dispersal distance a significant factor in determining their persistence in the landscape. Main conclusions At a landscape scale it is difficult to measure animal distributions directly and yet conservation planning often necessitates knowledge of where, and in what numbers, animals are found, and how their distributions will be affected by interventions. SEPMs offer a useful tool for predicting this, and for refining conservation plans before irreversible decisions are taken in practice.     

Mobility unevenness in rock–paper–scissors models

We investigate a tritrophic system whose cyclic dominance is modelled by the rock–paper–scissors game. We consider that organisms of one or two species are affected by movement limitations, which unbalances the cyclic spatial game. Performing stochastic simulations, we show that mobility unevenness controls the population dynamics. In the case of one slow species, the predominant species depends on the level of mobility restriction, with the slow species being preponderant if the mobility limitations are substantial. If two species face mobility limitations, our outcomes show that being higher dispersive does not constitute an advantage in terms of population growth. On the contrary, if organisms move with higher mobility, they expose themselves to enemies more frequently, being more vulnerable to being eliminated. Finally, our findings show that biodiversity benefits in regions where species are slowed. Biodiversity loss for high mobility organisms, common to cyclic systems, may be avoided with coexistence probability being higher for robust mobility limitations. Our results may help biologists understand the dynamics of unbalanced spatial systems where organisms’ dispersal is fundamental to biodiversity conservation.     

Children's ideas of animals' internal structures

Children's understanding about animal internal structure can be affected by several factors which are poorly understoodby teachers. We conducted a large sample study (n=702) of children aged 6–16 years (Grades 1-9) examining children's responses to animals of various size, species and dimensions (2D and 3D objects), and exploring factors which might affect the development of their knowledge. Each child made a drawing of what they thought was inside a specimen animal. We found that using 2D representations of an animal negatively affected the content of children drawings.The effect of animalsize was shown only in children's understanding of skeletons, but not organ systems. The effect of animal species showedclear significant effect of children's ideas on both organ systems and skeleton. Investigation showed that parents' education level, children's experiences with animal-rearing and age significantly affected their scores from organ system. Surprisingly, school books were most frequently cited as sources of children's knowledge rather than their own experience, or information from the internet, encyclopaedias, television or parents.     

Backward bifurcation and oscillations in a nested immuno-eco-epidemiological model

This paper introduces a novel partial differential equation immuno-eco-epidemiological model of competition in which one species is affected by a disease while another can compete with it directly and by lowering the first species' immune response to the infection, a mode of competition termed stress-induced competition. When the disease is chronic, and the within-host dynamics are rapid, we reduce the partial differential equation model (PDE) to a three-dimensional ordinary differential equation (ODE) model. The ODE model exhibits backward bifurcation and sustained oscillations caused by the stress-induced competition. Furthermore, the ODE model, although not a special case of the PDE model, is useful for detecting backward bifurcation and oscillations in the PDE model. Backward bifurcation related to stress-induced competition allows the second species to persist for values of its invasion number below one. Furthermore, stress-induced competition leads to destabilization of the coexistence equilibrium and sustained oscillations in the PDE model. We suggest that complex systems such as this one may be studied by appropriately designed simple ODE models.     

Effects of Number of Animals Monitored on Representations of Cattle Group Movement Characteristics and Spatial Occupancy

The number of animals required to represent the collective characteristics of a group remains a concern in animal movement monitoring with GPS. Monitoring a subset of animals from a group instead of all animals can reduce costs and labor; however, incomplete data may cause information losses and inaccuracy in subsequent data analyses. In cattle studies, little work has been conducted to determine the number of cattle within a group needed to be instrumented considering subsequent analyses. Two different groups of cattle (a mixed group of 24 beef cows and heifers, and another group of 8 beef cows) were monitored with GPS collars at 4 min intervals on intensively managed pastures and corn residue fields in 2011. The effects of subset group size on cattle movement characterization and spatial occupancy analysis were evaluated by comparing the results between subset groups and the entire group for a variety of summarization parameters. As expected, more animals yield better results for all parameters. Results show the average group travel speed and daily travel distances are overestimated as subset group size decreases, while the average group radius is underestimated. Accuracy of group centroid locations and group radii are improved linearly as subset group size increases. A kernel density estimation was performed to quantify the spatial occupancy by cattle via GPS location data. Results show animals among the group had high similarity of spatial occupancy. Decisions regarding choosing an appropriate subset group size for monitoring depend on the specific use of data for subsequent analysis: a small subset group may be adequate for identifying areas visited by cattle; larger subset group size (e.g. subset group containing more than 75% of animals) is recommended to achieve better accuracy of group movement characteristics and spatial occupancy for the use of correlating cattle locations with other environmental factors.     

Can a Species Keep Pace with a Shifting Climate?

Consider a patch of favorable habitat surrounded by unfavorable habitat and assume that due to a shifting climate, the patch moves with a fixed speed in a one-dimensional universe. Let the patch be inhabited by a population of individuals that reproduce, disperse, and die. Will the population persist? How does the answer depend on the length of the patch, the speed of movement of the patch, the net population growth rate under constant conditions, and the mobility of the individuals? We will answer these questions in the context of a simple dynamic profile model that incorporates climate shift, population dynamics, and migration. The model takes the form of a growth-diffusion equation. We first consider a special case and derive an explicit condition by glueing phase portraits. Then we establish a strict qualitative dichotomy for a large class of models by way of rigorous PDE methods, in particular the maximum principle. The results show that mobility can both reduce and enhance the ability to track climate change that a narrow range can severely reduce this ability and that population range and total population size can both increase and decrease under a moving climate. It is also shown that range shift may be easier to detect at the expanding front, simply because it is considerably steeper than the retreating back.     

Separating the effects of prey size and speed on the kinematics of prey capture in the omnivorous lizard Gerrhosaurus major

Feeding behavior is known to be modulated as prey properties change. During prey capture, external prey properties, including size and mobility, are likely some of the most important components in predator–prey interactions. Whereas prey size has been demonstrated to elicit modulation of jaw movements during capture, how prey speed affects the approach and capture of prey remains unknown. We quantified the kinematics associated with movements of both the feeding and locomotor systems during prey capture in a lizard, Gerrhosaurus major, while facing prey differing in size and mobility (newborn mice, grasshoppers, and mealworms). Our data show that the feeding and locomotor systems were recruited differently in response to changes in the size or speed of the prey. The timing of jaw movements and of the positioning of the head are affected by changes in prey size—and speed, to a lesser extent. Changes in prey speed resulted in concomitant changes in the speed of strike and an early and greater elevation of the neck. External prey properties, and prey mobility in particular, are relevant in predator–prey interactions and elicit specific responses in different functional systems.     

BODY SIZE,SPERM COMPETITION,AND DETERMINANTS OF REPRODUCTIVE SUCCESS IN MALE SAVANNA BABOONS

One component of sexual selection is sperm competition. It has been reasoned that the intensity of sperm competition may be reflected in the relative testicular sizes of animals. Among males residing in multimale breeding systems, testicular size is relatively larger than among males residing in unimale mating systems. Information on whether differences in testicular size within a species can account for differences in male reproductive success is unavailable for natural populations of primates. A population of six troops of savanna baboons in Kenya was surveyed for morphometric analysis, and one of these troops was the subject of extensive behavioral observations afterwards. Testicular weights could not be obtained, but measurements of linear dimensions were transformed into volumetric estimates. Male weight accounted for 30% of the variance in testicular volume. Neither body size nor testicular volume was associated with differences in male reproductive activity. The outcome of fights over access to females could not be related to male body size, and ejaculatory patterns of males were independent of testicle size. Both sperm competition and aggressive competition intensified during the four-day optimum conception period, but fights over access to consort females were infrequent. Among savanna baboons, the probability of an ejaculation resulting in a conception is fairly low, which may account for the infrequency of injurious fights. Although testicle size influences sperm production, it does not influence either the timing of mating or the fertilizing capacity of spermatozoa, and both of these factors probably account for a substantial fraction of the variance in male baboon paternity. Sperm competition is an adjunct to agonistic competition as a mechanism affecting male baboon reproductive success. It is concluded that male reproductive success in baboons is affected more by social factors than by morphological traits associated with size.     

Effects of competitive asymmetry on a local density model of plant interference

Although competition between plants is usually asymmetric (i.e. larger plants have a disproportionate effect on smaller plants) almost all models of plant competition at the local level have assumed symmetric competition. We add a simple version of competitive asymmetry to the local density neighborhood models of plant interference and population dynamics developed by Pacala & Silander (1985, Am. Nat. 125, 385-411; 1987, Oikos 48, 217-224) by assuming that plants within a neighborhood can be put in a linear dominance hierarchy based upon their initial size. The size of a focal plant is a function of the number of dominant and the number of subordinate neighbors within its neighborhood, with subordinate neighbors having less of an effect than dominant ones. Asymmetry prevents precipitous changes in focal plant size with changes in local density, making the relationship between focal plant size and local density hyperbolic, even if the symmetric model is not hyperbolic. Thus, asymmetry makes the model conform to the law of constant final yield, irrespective of the form of the relationship between plant size and local crowding. Asymmetry also prevents population dynamic oscillations in the model in cases in which it would occur in the absence of asymmetry. The results show that asymmetry has major effects on a model of local interference in plants, and point to the importance of including it in such models.     

Interspecific competition in small rodents: from populations to individuals

The role of interspecific competition in shaping animal and plant communities has formed one of the major issues in ecology for decades. Small mammals, mainly rodents, have been among the model systems used for research on interspecific competition. Most studies within small mammal systems in the past have examined effects of competition on population attributes such as on population size, habitat use, or population dynamics. Population-level responses are the cumulative effects of individual responses, however, the influence of competition on individual life-history traits has rarely been studied. Research on life-histories may bridge gaps between population biology and effects of competition on individual behaviour. In this paper, we review recent research approaches to interspecific competition in rodents based on census data and species assemblages, that use regression analysis, time series analysis, removal and exclusion experiments, and showcase our own experimental research on the effects of interspecific competition on individual life-history traits in boreal voles.     

What if it gets crowded? Density‐dependent tortuosity in individual movements of a Neotropical mammal

Effects of density dependence on animal movements have received much attention in ecology, but it is still debated to what extent dispersal and movements in general are density dependent, and their potential contribution to population regulation processes. Here, we determine the occurrence and nature of density dependence in the movements of a Neotropical marsupial, the black‐eared opossum Didelphis aurita Wied‐Neuwied 1826. Using spool‐and‐line tracking devices, we estimated the tortuosity of fine‐scale movements of 149 individuals by their fractal dimension D. We evaluated the relative importance of population size, reproductive or climatic seasons and reproductive maturity of individuals as determinants of movement tortuosity, using a model selection approach. Population size was the most important determinant of movement tortuosity, with season (climatic seasons for females, reproductive seasons for males) and reproductive maturity as secondary but also important variables. We detected a positive density‐dependent effect on movement tortuosity, resulting in more intensive use of areas by individuals during periods of high population size. This positive association between movement tortuosity and population size is more likely to result from intraspecific competition, which forces individuals to explore their environment more intensively during high‐density periods. Therefore, despite being density dependent, movements in D. aurita apparently do not contribute to population regulation mechanisms.