The evolutionary and ecological roots of human social organization

Social organization among human foragers is characterized by a three-generational system of resource provisioning within families, long-term pair-bonding between men and women, high levels of cooperation between kin and non-kin, and relatively egalitarian social relationships. In this paper, we suggest that these core features of human sociality result from the learning- and skill-intensive human foraging niche, which is distinguished by a late age-peak in caloric production, high complementarity between male and female inputs to offspring viability, high gains to cooperation in production and risk-reduction, and a lack of economically defensible resources. We present an explanatory framework for understanding variation in social organization across human societies, highlighting the interactive effects of four key ecological and economic variables: (i) the role of skill in resource production; (ii) the degree of complementarity in male and female inputs into production; (iii) economies of scale in cooperative production and competition; and (iv) the economic defensibility of physical inputs into production. Finally, we apply this framework to understanding variation in social and political organization across foraging, horticulturalist, pastoralist and agriculturalist societies.     

Environmental heterogeneity and neighbourhood interference modulate the individual response of Juniperus thurifera tree-ring growth to climate

Individual variation of tree-ring growth response to climate and heterogeneity of the local environment are usually neglected in dendrochronological research. Even if there is evidence showing that individual responsiveness to climate may depend on intrinsic traits such as tree age, size or sex, its modulation by the local heterogeneity of extrinsic factors has been less studied. Using an extensive, strictly regular sampling scheme across a 3300 ha woodland, we assessed the individual variation of tree-ring growth responses to climate in 100 Juniperus thurifera L. trees. The climatic response was evaluated by bootstrapped correlations of both population- and individual-based tree-ring chronologies with monthly records of precipitation, cloudiness, minimum and maximum temperatures. We studied also the influence of extrinsic abiotic (elevation, slope, heat load, tree location) and biotic (competition from neighbouring trees) factors on the individual growth variation and its climatic response. At a population level, growth was controlled by February–March precipitation, April minimum temperature, and June water stress. A significant proportion of individuals did not respond to those variables, but were sensitive to others not relevant at the population level. Inter-annual growth variation was strongly modulated by competition, whereas trees under lower competition levels, in eastern and warmer areas, were the most responsive to climate. The individual climatic response was, at least partially, modulated by the local heterogeneity of extrinsic factors. By considering environmental heterogeneity and neighbourhood interference we can identify the spectrum of site-dependent climatic responses in a population, which in turn will enable more realistic predictions of tree responses to ongoing climate change.     

Temporal coherence in the limnology of a suite of lakes in Wisconsin, U.S.A.

SUMMARY. 1. Temporal coherence between pairs of lakes over 7 years was measured for thirty-seven limnological variables in seven lakes at the North Temperate Lakes Long Term Ecological Research site in Northern Wisconsin. This analysis tested. First, whether lakes more similar in exposure to the atmosphere were more temporally coherent than lakes which differed more in exposure and, second, whether temporal coherence in lakes progressively decreased from variables more directly influenced by climatic factors such as thermal and hydrologic properties, to those chemical and biological properties which may be less directly influenced by climatic factors. 2. The lakes were a heterogeneous set in respect to exposure to climatic factors as estimated by the ratio of‘lake area/mean depth’and by other morphometric features, and they also differed in their position in the landscape, fertility and fish assemblages. Limnological variables formed a progression from those expected to respond directly to climatic factors to those which would not. They ranged from water level and temperatures to chemical variables such as pH, calcium concentrations and total dissolved phosphorus to biological variables such as chlorophyll concentrations, invertebrate and fish abundances. 3. Coherence was estimated by the correlation between lake pairs for each of the different variables. The mean correlation and the percentage of strong correlations were calculated for each lake pair across all variables, and for each variable across all lake pairs, and both measures of coherence gave similar answers to the questions posed above. 4. Temporal coherence between takes was higher for lakes similar in their exposure to climatic factors; mean correlation (r?) being +0.3 to +0.7 for these lakes and <+0.3 for lakes not similar in exposure. None of the lake pairs had high coherence across all variables. 5. Temporal coherence between lakes was greater for limnological variables directly influenced by climatic factors than for variables either indirectly affected by climate or complexly influenced by other types of factors. Water-level variables had a coherence near 1, r?=0.9. All biological variables had low coherence, some near r?=0.0. Chemical variables more likely to be influenced directly by climatic factors appeared to be more coherent than those more influenced by hydrology or biology. Most silica and phosphorus variables had coherences less than r?=0.15. 6. Coherence was not as strongly related to similarity in landscape position as it was to similarity in exposure to climatic factors, and was not jelated to the proximity of the lake pairs or to their similarity in fertility. 7. A conceptual model was presented to explain how climatic signals are filtered by the lake's exposure to climatic factors and by terrestrially mediated and in-lakc processes to reduce the coherence of lake pairs owing to time lags, frequency shifts and complex interference pattems. 8. Coherence is an important property to evaluate because it influences how broadly we can extrapolate results from a lake or set of lakes (for example, to a lake district) and because manipulative whole lake experiments often use adjacent lakes as reference lakes to evaluate treatment effects. The low coherence in this set of lakes in general and of individual variables such as chlorophyll concentrations and yellow perch year-class strength are discussed. 9. We found no studies that explicitly addressed interlake coherence; one long-term study of forest production made it clear that coherence among sites in a landscape will be a function of scale.     

The role of group size and environmental factors on survival in a cooperatively breeding tropical passerine

1. Variation in survival, a major determinant of fitness, may be caused by individual or environmental characteristics. Furthermore, interactions between individuals may influence survival through the negative feedback effects of density dependence. Compared to species in temperate regions, we have little knowledge about population processes and variation in fitness in tropical bird species. 2. To investigate whether variation in survival could be explained by population size or climatic variables we used capture-recapture models in conjunction with a long-term data set from an island population of the territorial, cooperatively breeding Seychelles warbler (Acrocephalus sechellensis). The lack of migration out of the study population means that our results are not confounded by dispersal. 3. Annual survival was high, both for adults (84%) and juveniles (61%), and did not differ between the sexes. Although there was significant variation in survival between years, this variation could not be explained by overall population size or weather variables. 4. For territorial species, resource competition will work mainly on a local scale. The size of a territory and number of individuals living in it will therefore be a more appropriate measure of density than overall population density. Consequently, both an index of territory quality per individual (food availability) and local density, measured as group size, were included as individual covariates in our analyses. 5. Local density had a negative effect on survival; birds living in larger groups had lower survival probabilities than those living in small groups. Food availability did not affect survival. 6. Our study shows that, in a territorial species, although density-dependent effects might not be detectable at the population level they can be detected at the individual territory level - the scale at which individuals compete. These results will help to provide a better understanding of the small-scale processes involved in the dynamics of a population in general, but in particular in tropical species living in relatively stable environments.     

Effects of body size and resource availability on dispersal in a native and a non-native estuarine snail

I manipulated snail densities of two coexisting species of salt marsh snail, Cerithidea californica Haldeman (native) and Batillaria attramentaria Sowerby (non-indigenous) to investigate how resource levels set by intraspecific competition may influence dispersal rates. I used two distinct size classes of the snails (mature and immature) to determine if the effects of competition on dispersal differed between developmental stages of a consumer. Dispersal attempts were measured within enclosure pens by counting snails climbing the sides of the enclosure. The influence of snail density per se and resource levels (which were set by snail densities) on dispersal rates were separated by comparing responses of snails to density before and after resources became depleted. For large snails of both species, dispersal increased as resource levels decreased, supporting the hypothesis that competition influences dispersal rates. Small snails of both species, in contrast, always dispersed at relatively higher rates than larger individuals, but were not influenced by variation in resource levels. This result corroborates other studies that have shown reduced competition in these species at smaller size, and suggests that another mechanism, such as genetically coded behavior to disperse when young, influences their behavior. Previous experiments demonstrated Batillaria's superior resource conversion efficiency; therefore, I had hypothesized that for any given resource level, Cerithidea would disperse more, because it was more affected by resource availability. Adult Batillaria, however, responded more sensitively to resource levels (i.e., dispersed more at any given resource level) than Cerithidea. This counter-intuitive result illustrates the potential importance of genetic limitations on behavioral responses available to a species. Constraints on behavioral responses may have been accentuated since Batillaria is a non-indigenous species whose evolved behavioral responses are not necessarily adapted to its present, non-native environment.     

Density-dependent reproductive output in relation to a drastically varying food supply: getting the density measure right

When a limiting resource (e.g. food) varies drastically between years, and population density is measured in the conventional way as individuals per area, demographic processes such as productivity and survival may erroneously be considered density-independent. We tested the hypothesis that if the variation in a limiting resource is not taken into account in the density measure, this may lead to erroneous conclusions about the density-dependence of demographic variables. We studied the food-related variation in productivity of bramblings Fringilla montifringilla , an insectivorous passerine bird, using 19 years of standardised insect censusing, bird censusing and mist-netting of birds in subalpine birch forest in Swedish Lapland. The yearly variation in our measure of brambling per capita productivity (numbers of juveniles per adult trapped) was explained to 30–40% by the larvae abundance of the moth Epirrita autumnata . Taking larvae density into account, no other environmental variable (inferred predation pressure, breeding phenology, and summer temperature) was significantly related to variation in reproductive output. There was no effect of brambling population density on per capita productivity, that is, when density was measured the conventional way, productivity seemed density-independent. However, per capita productivity was significantly and negatively correlated to the food-related population density (population density divided by larval density), supporting the hypothesis that not including a limiting resource into the density measure may indeed lead to erroneous conclusions about the density-dependence of demographic variables.     

Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus (L.)

Models for sex allocation assume that increased expenditure of resources on male function decreases the resources available for female function. Under some circumstances, a negative genetic correlation between investment in stamens and investment in ovules or seeds is expected. Moreover, if fitness returns for investment in male and female function are different with respect to size, sex allocation theory predicts size‐specific gender changes. We studied sex allocation and genetic variation for investment in stamens, ovules and seeds at both the flower and the plant level in a Dutch population of the wind‐pollinated and predominantly outcrossing Plantago coronopus. Data on biomass of floral structures, stamens, ovules, seedset and seedweight were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and (genetic) correlations were estimated from the greenhouse‐grown progeny of maternal families, raised at two nutrient levels. The proportion of reproductive biomass invested in male function was high at flowering (0.86 at both nutrient levels) and much lower at fruiting (0.30 and 0.40 for the high and low nutrient treatment, respectively). Androecium and gynoecium mass exhibited moderately high levels of genetic variance, with broad‐sense heritabilities varying from 0.35 to 0.56. For seedweight no genetic variation was detected. Significant among‐family variation was also detected for the proportion of resources invested in male function at flowering, but not at fruiting. Phenotypic and broad‐sense genetic correlations between androecium and gynoecium mass were positive. Even after adjusting for plant size, as a measure of resource acquisition, maternal families that invested more biomass in the androecium also invested more in the gynoecium. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the overall lack of a negative correlation between male and female function. Larger plants had a more female‐biased allocation pattern, brought about by an increase in seedset and seedweight, whereas stamen biomass did not differ between small and large plants. These results are discussed in relation to size‐dependent sex allocation theory (SDS). Our results indicate that the studied population harboured substantial genetic variation for reproductive characters.     

Signal modulation as a mechanism for handicap disposal

Signal honesty may be compromised when heightened competition provides incentive for signal exaggeration. Some degree of honesty might be maintained by intrinsic handicap costs on signalling or through imposition of extrinsic costs, such as social punishment of low quality cheaters. Thus, theory predicts a delicate balance between signal enhancement and signal reliability that varies with degree of social competition, handicap cost, and social cost. We investigated whether male sexual signals of the electric fish Brachyhypopomus gauderio would become less reliable predictors of body length when competition provides incentives for males to boost electric signal amplitude. As expected, social competition under natural field conditions and in controlled lab experiments drove males to enhance their signals. However, signal enhancement improved the reliability of the information conveyed by the signal, as revealed in the tightening of the relationship between signal amplitude and body length. Signal augmentation in male B. gauderio was independent of body length, and thus appeared not to be curtailed through punishment of low quality (small) individuals. Rather, all individuals boosted their signals under high competition, but those whose signals were farthest from the predicted value under low competition boosted signal amplitude the most. By elimination, intrinsic handicap cost of signal production, rather than extrinsic social cost, appears to be the basis for the unexpected reinforcement of electric signal honesty under social competition. Signal modulation may provide its greatest advantage to the signaller as a mechanism for handicap disposal under low competition rather than as a mechanism for exaggeration of quality under high competition.     

Influence of nerve supply on hand electromyography coherence during a three-digit task

Intermuscular coupling has been investigated to understand neural inputs to coordinate muscles in a motor performance. However, little is known on the role of nerve innervation on intermuscular coupling. The purpose of this study was to investigate how the anatomy of nerve distribution affected intermuscular coupling in the hand during static grip. Electromyographic (EMG) signals were recorded from intrinsic and extrinsic muscles while subjects performed a static grip. Coherence was computed for muscle pairs innervated by either the same or different nerves. The results did not support the hypothesis that muscles sharing the same nerve exhibit greater coupling than muscles innervated by different nerves. In general, extrinsic muscle pairs displayed higher coherence than intrinsic pairs. The results suggest that intermuscular coupling in a voluntary motor task is likely modulated in a functional manner and that different nerves might transport common neural inputs to functionally coupled muscles.     

Niche overlap between herbivorous cladocerans; the role of food quality and habitat homogeneity

In spite of resource limitation, five abundant species of herbivorous metazoan zooplankton in a humic lake exhibited extensive niche overlap both with regard to seasonal and spatial occurrence, time of reproductive maxima, juvenile release and food choice. Their coexistence could not be explained by modifying predation, environmental oscillations or recolonization.Laboratory bottle experiments indicated only weak interspecific interactions between the three tested species at low food levels, but negative interactions were induced at elevated food levels. Bosmina appeared as competitively inferior during enrichment with cultured algae, but as the superior species during starvation. At low nutrient levels, all species coexisted for several generations with low reproduction, in accordance with the lake situation. It was concluded that the observed niche overlaps would be promoted if; 1) Intraspecific competition is more important than interspecific competition. 2) All species are co-adapted to low nutrient availability, food is quantitatively in surplus, but qualitatively deficient (mainly recycled detritus). During such conditions, no species would be capable of obtaining a population increase until extinction of the other species. This situation may be typical of oligotrophic humic lakes, and of other localities with low predation pressure and high inputs of allochthonous particulate carbon.     

Climatic variability and body size variation in the muskrats (Ondatra zibethicus) of North America

Summary Patterns of geographic variation in nine morphological characters of adult muskrats (Ondatra zibethicus) are investigated by multiple regression and canonical correlation analysis. Three variables describing the annual precipitation regimen account for 16 to 33% of the variance in each of the skeletal measurements. Highly variable precipitation patterns result in low rates of germination and survivorship for preferred aquatic food plants which in turn reduces average total food availability for muskrats. In large individuals nutritive demands may exceed supply, thus selection favors smaller body size in areas of low food availability.After removing the variation attributable to three precipitation variables, the residual variation in the morphological variables is exposed to canonical correlation analysis with a set of 10 environmental and geographic variables. A canonical variate loaded for climatic seasonality accounts for 60% of the variance in a canonical variate of the morphological residuals which load as a general body size variate. It is argued that seasonality is a major factor selecting for large body size in muskrats and other organisms. During seasonal periods of resource abundance natural selection favors individuals with rapid growth to a large size, while concurrently enhancing survivorship through oncoming periods of resource shortage.     

Stabilization of population fluctuations due to cannibalism promotes resource polymorphism in fish

Resource polymorphism is a well-known phenomenon in many taxa, assumed to be a consequence of strong competition for resources and to be facilitated by stable environments and the presence of several profitable resources on which to specialize. In fish, resource polymorphism, in the form of planktivore-benthivore pairs, is found in a number of species. We gathered literature data on life-history characteristics and population dynamics for 15 fish species and investigated factors related to the presence of such resource polymorphism. This investigation indicated that early cannibalism and low overall population variability are typically associated with the presence of resource polymorphism. These findings match previously reported patterns of population dynamics for size-structured fish populations, whereby early cannibalism has been shown to decrease temporal variation in population dynamics and to equalize the profitability of the zooplankton and macroinvertebrate resources. Our study suggests that competition alone is not a sufficient condition for the development of resource polymorphism because overly strong competition is typically associated with increased temporal variation (environmental instability). We conclude that although resource competition is an important factor regulating the development of resource polymorphism, cannibalism may also play a fundamental role by dampening population oscillations and possibly by equalizing the profitability of different resources.     

Hierarchical models of intra-specific competition: scramble versus contest

Hierarchical structured models for scramble and contest intraspecific competition are derived. The dynamical consequences of the two modes of competition are studied under the assumption that both populations divide up the same amount of a limiting resource at equal population levels. A comparison of equilibrium levels and their resiliences is made in order to determine which mode of competition is more advantageous. It is found that the concavity of the resource uptake rate is an important determining factor. Under certain circumstances contest competition is more advantageous for a population while under other circumstances scramble competition is more advantageous.Supported by NSF grant DMS-9306271     

Intra-sexual competition alters the relationship between testosterone and ornament expression in a wild territorial bird

In a reliable signalling system, individual quality is expected to mediate the costs associated with ornamental displays, with relatively lower costs being paid by individuals of higher quality. These relative costs should depend not only on individual quality, but also on levels of intra-sexual competition. We explored the current and delayed effects that testosterone implants have on bird ornamentation in populations with contrasted population densities, as a proxy for intra-sexual competition. In a replicated experiment, we manipulated testosterone in 196 yearling male red grouse Lagopus lagopus scoticus in autumn in populations of high and low levels of intra-sexual competition. Males were assigned to one of three exogenous testosterone (T) treatments: empty implants (T0), small T implants (T1) or larger T implants (T2). We monitored subsequent changes in testosterone levels, ornament size and carotenoid-based colouration, carotenoid levels and body condition from autumn to spring. Testosterone implants increased testosterone levels, comb redness and comb size, and decreased body condition but these effects depended on levels of intra-sexual competition. Specifically, T2-implanted birds increased testosterone levels and comb size more, and reduced body condition more, in populations where intra-sexual competition was low. In the following spring, testosterone levels of T2-treated birds kept increasing in populations where intra-sexual competition was high but not in populations where intra-sexual competition was low. Our results highlight that levels of intra-sexual competition alter the relationship between testosterone levels and ornament expression, influencing their condition-dependence; they also indicate that the outcome of standard hormone manipulation conducted in free-living animals vary depending on the population context.     

Dietary niche and population dynamic feedbacks in a novel habitat

Population dynamics and resource use are often intricately connected via density‐dependent intraspecific competition. However, experimental studies of concurrent change in population and resource use dynamics are scarce. In particular, the impact of factors such as genetic diversity, which can affect both population dynamics and competition, remains unexplored. Using stable isotope analysis and periodic population censuses, we quantified both diet and population dynamics in wheat‐adapted Tribolium castaneum (flour beetle) populations provided with an additional novel resource (corn). Populations were initiated with different levels of genetic variation for traits relevant to population growth and resource use (e.g. fecundity and survival).We found that high population size decreased subsequent corn use, and high corn use in turn lowered population size. Surprisingly, we did not detect a significant effect of founding genetic variation on resource niche expansion, although genetic variation increased overall population size and stability. In contrast, dietary niche expansion decreased both population size and stability. Finally, larval and adult niche dynamics were uncorrelated, suggesting that various life stages perceive or respond differentially to intraspecific competition and resource availability. Our experiments indicate that population performance in a novel habitat depends on stage‐specific interactions between resource use, standing genetic variation, and population size.     

Phenology constrains opportunistic growth response in <Emphasis Type="Italic">Bromus tectorum</Emphasis> L.

Seasonal resource availability may act as a constraint on plant phenology and thereby influence the range of growth responses observed among populations of annual species, especially those occupying a wide range of environments. We compared a mesic and a xeric population of the non-native, annual grass, Bromus tectorum, to examine phenology in response to interspecific competition and water availability. Using a target-neighborhood approach, we assessed how phenological patterns of the two populations affected morphological and growth responses to enhanced resource availability represented by late-season soil moisture. The xeric population exhibited a highly constrained phenology and was unable to extend the growing season despite available soil resources. Because of the low phenotypic variation, allocation to reproduction was similar across resource conditions. In contrast, the mesic population flowered later and showed a more opportunistic phenology in response to late-season water availability. The mesic population was not able to maintain consistent reproductive allocation at low resource levels. The responses of the two populations to late-season water availability were not affected by the density of neighboring plants. We suggest that post-introduction selection pressure on B. tectorum in the xeric habitat has resulted in a more fixed phenology which limits opportunistic response to unpredictable, particularly late-season resource availability. Opportunistic and fixed responses represent contrasting strategies for optimizing fitness in temporally varying environments and, while both play important roles for ensuring reproductive success, these results suggest that local adaptation to temporal resource variation may reflect a balance between flexible and inflexible phenology.     

Disentangling the causes of intrainflorescence variation in floral traits and fecundity in the hermaphrodite Silene acutifolia

Inflorescence architecture directly determines variations in floral traits and fecundity. Disentangling these patterns of variation is crucial to understanding intraplant variation, which sometimes is directly attributed to competition for resources with developing fruits. The dichasial cymes of Silene acutifolia were experimentally manipulated in the field to analyze whether the declines in petal size, ovule number, fruit set, and seed/ovule ratio along the inflorescence are constrained by ontogenetic development or are phenotypically plastic in response to environmental changes. At the same time, the level of pollen deficit was measured on different positions of the dichasia. The results showed clearly that all measured variables were more influenced by architecture than by resource competition with developing fruits; the removal of central (basal) and primary lateral flowers in the dichasia did not increase either the measures of floral characters or fecundity. On the other hand, although most of the decline in fecundity was due to architectural effects, there was also a pollen limitation, dependent to some degree on inflorescence position, which was probably due to lower pollen availability in the population when secondary flowers are in the female phase.     

Extrinsic Mortality Effects on Reproductive Strategies in a Caribbean Community

Extrinsic mortality is a key influence on organisms’ life history strategies, especially on age at maturity. This historical longitudinal study of 125 women in rural Domenica examines effects of extrinsic mortality on human age at maturity and pace of reproduction. Extrinsic mortality is indicated by local population infant mortality rates during infancy and at maturity between the years 1925 and 2000. Extrinsic mortality shows effects on age at first birth and pace of reproduction among these women. Parish death records show huge historical variation in age-specific mortality rates. The infant mortality rate (IMR) in the early 1920s was low, increased dramatically beginning in 1929, and reached a maximum in the 1950s, at which point IMR declined steadily to its present low rate. The mortality rate early in life showed a quadratic association with age at first birth. Women who experienced conditions of low IMR early in life reproduced relatively late in life. Those born into moderately high levels of infant mortality tended to reproduce earlier than those born at low levels. At very high infant mortality levels early in life, women went on to delay reproduction until relatively late, possibly as a result of somatic depletion and energetic stress associated with the conditions that lead to high IMR. Population mortality rates at age of maturity also showed a quadratic association with age at first birth. The pace of reproduction, estimated as number of surviving offspring controlled for maternal age, showed a similar quadratic effect. There were complex interactions between population mortality rates in infancy and at maturity. When extrinsic mortality was high during infancy, extrinsic mortality later in life had little effect on timing of first birth. When extrinsic mortality was low to moderate in infancy, extrinsic mortality later in life had significant effects on adult reproduction. I speculate that these effects are mediated through development of personality facets associated with reproduction.     

Physiologically structured models – from versatile technique to ecological theory

A ubiquitous feature of natural communities is the variation in size that can be observed between organisms, a variation that to a substantial degree is intraspecific. Size variation within species by necessity implies that ecological interactions vary both in intensity and type over the life cycle of an individual. Physiologically structured population models (PSPMs) constitute a modelling approach especially designed to analyse these size‐dependent interactions as they explicitly link individual level processes such as consumption and growth to population dynamics. We discuss two cases where PSPMs have been used to analyse the dynamics of size‐structured populations. In the first case, a model of a size‐structured consumer population feeding on a non‐structured prey was successful in predicting both qualitative (mechanisms) and quantitative (individual growth, survival, cycle amplitude) aspects of the population dynamics of a planktivorous fish population. We conclude that single generation cycles as a result of intercohort competition is a general outcome of size‐structured consumer–resource interactions. In the second case, involving both cohort competition and cannibalism, we show that PSPMs may predict double asymptotic growth trajectories with individuals ending up as giants. These growth trajectories, which have also been observed in field data, could not be predicted from individual level information, but are emergent properties of the population feedback on individual processes. In contrast to the size‐structured consumer–resource model, the dynamics in this case cannot be reduced to simpler lumped stage‐based models, but can only be analysed within the domain of PSPMs. Parameter values used in PSPMs adhere to the individual level and are derived independently from the system at focus, whereas model predictions involve both population level processes and individual level processes under conditions of population feedback. This leads to an increased ability to test model predictions but also to a larger set of variables that is predicted at both the individual and population level. The results turn out to be relatively robust to specific model assumptions and thus render a higher degree of generality than purely individual‐based models. At the same time, PSPMs offer a much higher degree of realism, precision and testing ability than lumped stage‐based or non‐structured models. The results of our analyses so far suggest that also in more complex species configurations only a limited set of mechanisms determines the dynamics of PSPMs. We therefore conclude that there is a high potential for developing an individual‐based, size‐dependent community theory using PSPMs.     

Experimental and model analyses of the effects of competition on individual size variation in wood frog (Rana sylvatica) tadpoles

1. Size variation is a ubiquitous feature of animal populations and is predicted to strongly influence species abundance and dynamics; however, the factors that determine size variation are not well understood. 2. In a mesocosm experiment, we found that the relationship between mean and variation in wood frog (Rana sylvatica) tadpole size is qualitatively different at different levels of competition created by manipulating resource supply rates or tadpole density. At low competition, relative size variation (as measured by the coefficient of variation) decreased as a function of mean size, while at high competition, relative size variation increased. Therefore, increased competition magnified differences in individual performance as measured by growth rate. 3. A model was developed to estimate the contribution of size-dependent factors (i.e. based on size alone) and size-independent factors (i.e. resulting from persistent inherent phenotypic differences other than size that affect growth) on the empirical patterns. 4. Model analysis of the low competition treatment indicated that size-dependent factors alone can describe the relationship between mean size and size variation. To fit the data, the size scaling exponent that describes the dependence of growth rate on size was determined. The estimated value, 0-83, is in the range of that derived from physiological studies. 5. At high competition, the model analysis indicated that individual differences in foraging ability, either size-based or due to inherent phenotypic differences (size-independent factors), were much more pronounced than at low competition. The model was used to quantify the changes in size-dependent or size-independent factors that underlie the effect of competition on size-variation. In contrast to results at low competition, parameters derived from physiological studies could not be used to describe the observed relationships. 6. Our experimental and model results elucidate the role of size-dependent and size-independent factors in the development of size variation, and highlight and quantify the context dependence of individual (intrapopulation) differences in competitive abilities.