The observed range for temporal mean-variance scaling exponents can be explained by reproductive correlation

The mean-variance scaling relationship known as Taylor's power law has been well documented empirically over the past four decades but a general theoretical explanation for the phenomenon does not exist. Here we provide an explanation that relates empirical patterns of temporal mean-variance scaling to individual level reproductive behavior. Initially, we review the scaling behavior of population growth models to establish theoretical limits for the scaling exponent b that is in agreement with the empirically observed range (1≤b≤2). We go on to show that the degree of reproductive covariance among individuals determines the scaling exponent b. Independent reproduction results in an exponent of one, while completely correlated reproduction results in the upper limit of two. Intermediate exponents, which are common empirically, can be generated through the decay of reproductive covariance with increasing population size. Finally, we describe how the link between reproductive correlation and the scaling exponent provides a way to infer properties of individual-level reproductive behavior, such as the relative influence of demographic stochasticity, from a macroecological pattern.     

A resource budget model to explain intraspecific variation in mast reproductive dynamics

Rowan, Sorbus aucuparia L., exhibits conspicuous geographic variation in its inter-annual patterns of reproduction across southern Norway. Along the southern west coast, trees flower and produce clusters of succulent berries every other year, while in the east, trees mast every 3 years. We investigate two hypotheses for this variation: local adaptation to environmental conditions that favors different reproductive schedules at different locations or variation arising from single stereotyped reproductive strategy in the face of a geographic variation in productivity. To assess the theoretical plausibility of each of these hypotheses, we develop a refined resource budget model for mast reproduction. The refined model assumes that there is a lower bound on the amount of resource that may be allocated to reproduction, which allows the prediction of the relation between reproductive schedules and productivity. From the analysis of the model, we find that the observed geographic transition in rowan mast is caused if either productivity in the east is lower than that in the west or if plants in the eastern populations have evolved to pause reproduction until energy reserves are replenished to higher levels. The prediction by the productivity gradient hypothesis match with the empirical finding that the productivity is, in fact, likely to be lower in the east than the west, although we lack empirical data to test the likelihood of the local adaptation hypothesis. There is a need for experimental studies to clarify the validity of the local adaptation hypothesis. Akiko Satake is the recipient of the 11th Denzaburo Miyadi Award.     

Current-year shoot based approach for annual variation in the reproductive output in Siberian alder (<Emphasis Type="Italic">Alnus hirsuta</Emphasis> var. <Emphasis Type="Italic">sibirica</Emphasis>)

Siberian alder (Alnus hirsuta var. sibirica) shows annual variation in reproductive output. However, this phenomenon has not been explained by hypotheses proposed in previous studies. In this study, we constructed a matrix model of current-year shoot dynamics to estimate and compare the reproductive output of current-year shoot population with or without annual variation, and explained the annual variation of reproductive output in terms of a reproductive strategy of the current-year shoot population of Siberian alder. The current-year shoot population of Siberian alder was divided into three functional groups: reproductive, maintenance, and explorative shoot sub-populations. A transition matrix was calculated from the relationships between 1-year-old shoots and the current-year shoots on them. The dynamics of the current-year shoot population was simulated using the estimated matrix on the patterns with or without annual variation in reproductive output. The pattern with annual variation in reproductive output yielded more reproductive current-year shoots than the pattern with high reproductive output every year. The annual variation of reproductive output may well be regarded as a reproductive strategy of current-year shoots to increase lifetime fecundity.     

Semi-empirical power-law scaling of new infection rate to model epidemic dynamics with inhomogeneous mixing

The expected number of new infections per day per infectious person during an epidemic has been found to exhibit power-law scaling with respect to the susceptible fraction of the population. This is in contrast to the linear scaling assumed in traditional epidemiologic modeling. Based on simulated epidemic dynamics in synthetic populations representing Los Angeles, Chicago, and Portland, we find city-dependent scaling exponents in the range of 1.7-2.06. This scaling arises from variations in the strength, duration, and number of contacts per person. Implementation of power-law scaling of the new infection rate is quite simple for SIR, SEIR, and histogram-based epidemic models. Treatment of the effects of the social contact structure through this power-law formulation leads to significantly lower predictions of final epidemic size than the traditional linear formulation.     

Effects of ectoparasite infestation during pregnancy on physiological stress and reproductive output in a rodent-flea system

Biotic and abiotic stressors impose various fitness costs on individuals across a variety of taxa. In vertebrates, these stressors typically trigger complex neuroendocrine responses that stimulate glucocorticoid (GC) secretion from the adrenal cortex. Short-term elevation of GCs can be adaptive as it shifts energy toward physiological processes that cope with acute stressors; however, chronic increases in GC levels could have detrimental effects on fitness. Parasitism can be considered an important biotic stressor in nature and a possible cause of reproductive failure that could substantially affect an individual’s fitness. Thus, we aimed to test the effects of parasitism and maternal stress, as measured by GCs, during pregnancy and the relationship between these variables and measures of reproductive output using a rodent-flea system. Female Egyptian spiny mice (Acomys cahirinus) were randomly assigned to flea (Parapulex chephrenis) infested or uninfested treatments before and during pregnancy. The offspring of these females were flea-free. Feces were collected at five time points during the experiment to determine maternal fecal glucocorticoid metabolite (FGCM) concentrations. Overall, infested females had lower FGCM levels during gestation but higher FGCM levels post-parturition and larger mass changes than uninfested females. Additionally, models related to pup quality and quantity often included some measure of maternal investment or body condition moderating relationships between infestation and stress. This suggests that flea parasitism or high GC levels alone might not significantly impact host reproduction but rather females can experience different effects depending on their level of investment, which could be limited by body condition and/or the number of pups present in a litter.     

Space-irrelevant scaling law for fish school sizes

Universal scaling in the power-law size distribution of pelagic fish schools is established. The power-law exponent of size distributions is extracted through the data collapse. The distribution depends on the school size only through the ratio of the size to the expected size of the schools an arbitrary individual engages in. This expected size is linear in the ratio of the spatial population density of fish to the breakup rate of school. By means of extensive numerical simulations, it is verified that the law is completely independent of the dimension of the space in which the fish move. Besides the scaling analysis on school size distributions, the integrity of schools over extended periods of time is discussed.     

Population density, phenotype and reproductive output in the grasshopper Chorthippus brunneus

ABSTRACT. 1. The effects of population density on the reproductive output of individual female grasshoppers ( Chorthippus brunneus Thunberg) and the relationships between reproductive output and component elements of each female's phenotype, were investigated.
2. Reproductive output was primarily determined by the rate of egg-pod production. Increased density led to significant reductions in the rate of egg-pod production and reproductive output.
3. Exoskeleton size (reflecting nymphal experience) was positively correlated with clutch size at both high and low densities, but condition (reflecting adult experience) showed no such correlation. At high density, exoskeleton size was more strongly correlated with the rate of egg-pod production and reproductive output than condition. It is concluded that the nymphal aspects of size are more important than the adult aspects.
4. At low density, females with small exoskeletons compensated for their smaller clutches by producing egg-pods at a faster rate. Thus, there was no overall relationship between reproductive output and any of the phenotypic characteristics.
5. The effects of competition were weakly asymmetric in high density populations. The weakness of the asymmetry suggests scramble-like interactions for resources. It is concluded that not only are the effects of competition influenced by individual differences, but also that competition may reveal differences that would not otherwise be apparent.     

Maternal body-volume as a constraint on reproductive output in lizards: evidence from the evolution of viviparity

A few species of squamate reptiles contain both oviparous (egg-laying) and viviparous (live-bearing) populations, and thus offer exceptional opportunities to test adaptationist hypotheses on the determinants of reproductive output. We focus on the hypothesis that maternal body-volume constrains reproductive output in squamate reptiles. If females are “full” of eggs, what happens when viviparity evolves within a lineage? Eggs increase in volume and mass during development, primarily due to the uptake of water, so how can they be accommodated within the mother's abdomen? We predict that the resultant increase in relative clutch mass (RCM) will be lessened by (1) a decrease in reproductive output (by reducing the number or size of offspring), and/or (2) an increase in maternal body-volume (via modifications of size or shape of adult females). Our comparisons of conspecific oviparous and viviparous lizards (Lerista bougainvillii) confirm that live-bearers carry heavier clutches (in both absolute and relative terms) and show the predicted shifts in body size and shape of reproductive females. However, offspring size and number were unaffected by the evolution of viviparity, and the shifts in maternal morphology were too small to fully offset the increase in clutch mass. Thus, RCMs increased by 50%, indicating that viviparous females produced clutches which more completely filled the space available in the abdominal cavity. We conclude that maternal body-volume does play a role in determining reproductive output, but that the observed clutch masses may be optimized, rather than maximized, with respect to the abdominal space available.     

Development of allometric relations for three mangrove species in South Florida for use in the Greater Everglades Ecosystem restoration

Mathematical relations that use easily measured variables to predict difficult-to-measure variables are important to resource managers. In this paper we develop allometric relations to predict total aboveground biomass and individual components of biomass (e.g., leaves, stems, branches) for three species of mangroves for Everglades National Park, Florida, USA. The Greater Everglades Ecosystem is currently the subject of a 7.8-billion-dollar restoration program sponsored by federal, state, and local agencies. Biomass and production of mangroves are being used as a measure of restoration success. A technique for rapid determination of biomass over large areas is required. We felled 32 mangrove trees and separated each plant into leaves, stems, branches, and for Rhizophora mangle L., prop roots. Wet weights were measured in the field and subsamples returned to the laboratory for determination of wet-to-dry weight conversion factors. The diameter at breast height (DBH) and stem height were also measured. Allometric equations were developed for each species for total biomass and components of biomass. We compared our equations with those from the same, or similar, species from elsewhere in the world. Our equations explained ≥93% of the variance in total dry weight using DBH. DBH is a better predictor of dry weight than is stem height and DBH is much easier to measure. Furthermore, our results indicate that there are biogeographic differences in allometric relations between regions. For a given DBH, stems of all three species have less mass in Florida than stems from elsewhere in the world.     

Parental care, cost of reproduction and reproductive skew: a general costly young model

Understanding the mechanisms by which animals resolve conflicts of interest is the key to understanding the basis of cooperation in social species. Conflict over reproductive portioning is the critical type of conflict among cooperative breeders. The costly young model represents an important, but underappreciated, idea about how an individual's intrinsic condition and cost of reproduction should affect the resolution of conflict over the distribution of reproduction within a cooperatively breeding group. However, dominant control in various forms and fixed parental care (offspring fitness dependent solely on total brood size) are assumed in previous versions of costly young models. Here, we develop a general costly young model by relaxing the restrictive assumptions of existing models. Our results show that (1) when the complete-control assumption is relaxed, the costly young model behaves very differently from the original model, and (2) when the fixed parental care assumption is relaxed, the costly young-costly care model displays similar predictions to the tug-of-war model, although the underlying mechanisms causing these similar patterns are different. These results, we believe, help simplify the seemingly divergent predictions of different reproductive skew models and highlight the importance of studying the group members' intrinsic conditions, costs of producing young, and costs of parental care for understanding breeding conflict resolution in cooperatively breeding animals.     

Self-amplification as a source of interindividual variability: Shelter selection in cockroaches

Although group effect and collective decisions have been described in many insect species, the behavioral mechanisms involved in the process remain poorly documented at the individual level. We examined how individual behavior depends on the environmental context and we precisely characterized the behavioral rules leading to settlement of individual cockroaches in resting site. We focused on the spatial and temporal distribution of individuals in absence of conspecifics. Using isolated adult males of the cockroach Periplaneta americana, we showed that the quality of resting sites and the duration of the settlement exerted an influence on the individual decision-making: the probability of leaving a resting site decreased with the time spent under a shelter. A numerical model derived from experimental data suggested that this simple rule of self-amplification can also account for the interindividual variability.     

Indicators of habitat quality and the reproductive output of a forest songbird in small and large fragments

The distribution of individuals among habitats that vary in quality (i.e. resource availability) may affect reproductive output at a population level. I compared indicators of habitat quality including the breeding experience ratios (inexperienced:experienced birds), turnover rates, pairing success, and densities of a forest songbird, the Eastern Yellow Robin Eopsaltria australis , in two small, food-poor forest fragments, with those in two large, food-rich fragments. I then evaluated the likelihood that the breeding experience ratio affected the reproductive output of populations. Inexperienced males occurred in small fragments eight times as often as in large fragments. Male turnover rates were 1.5 times higher in the small than large fragments, and 20% (10/50) of the male population were unpaired in the small fragments compared to 0% (0/25) in the large. None of these measures differed significantly for females. Experienced birds of both sexes produced almost all of the offspring compared with inexperienced birds. Despite these findings, reproductive output did not vary with fragment size for robins. Thus, while breeding experience clearly influenced the reproductive success of individuals, there were no obvious population consequences of having disproportionately more inexperienced males in the small fragments. I conclude that while male traits may be good indicators of resource levels within fragments (specifically, food availability), they may not be adequate predictors of population performance. Thus, the reproductive output of populations must be measured directly before conclusions concerning population performance can be made. Interestingly, breeding densities were not accurate indicators of either resource levels or population performance. Densities were two times higher in the smaller fragments, and I suggest that this result reflects problems with male dispersal among fragments caused by isolation.     

Seed dispersal patterns in a temperate forest during a mast event: performance of alternative dispersal kernels

Seed dispersal patterns were studied in a north-western Spain temperate forest community to assess the performances of alternative dispersal kernels during two years with ecologically contrasting scenarios; a non-mast year, and a mast year of the dominant canopy species, beech Fagus sylvatica. Dispersal kernels were fitted under a Bayesian modeling framework. Both simple and mixture kernels were considered for the five more abundant tree species (Corylus avellana, Crataegus monogyna, F. sylvatica, Ilex aquifolium and Taxus baccata). Mixture kernels provided a better fit for almost all species, and the log-normal performed best for T. baccata. No relationship between dispersal syndromes and the best dispersal kernel function emerged. However, we found temporal changes in the shape of the dispersal kernels that seemed to be related to variation in relative fruit production among species and the resulting changes in the responses of dispersal vectors. This reveals a potential role for disperser-mediated indirect effects in terms of introducing temporal variation in species spread. In this sense, our results highlight the need to consider single species seed dispersal as a community process. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Nutrient limitation restricts growth and reproductive output in a tropical montane cloud forest bromeliad: findings from a long-term forest fertilization experiment

From studies in seasonal lowland tropical forests, bromeliad epiphytes appear to be limited mainly by water, and to a lesser extent by nutrient supply, especially phosphorous. Less is understood about the mineral nutrition of tropical montane cloud forest (TMCF) epiphytes, even though their highest diversity is in this habitat. Nutrient limitation is known to be a key factor restricting forest productivity in TMCF, and if epiphytes are nutritionally linked to their host trees, as has been suggested, we would expect that they are also nutrient limited. We studied the effect of a higher nutrient input on reproduction and growth of the tank bromeliad Werauhia sintenisii in experimental plots located in a TMCF in Puerto Rico, where all macro- and micronutrients had been added quarterly starting in 1989 and continuing throughout the duration of this study. We found that bromeliads growing in fertilized plots were receiving litterfall with higher concentrations of N, P, and Zn and had higher concentrations of P, Zn, Fe, Al, and Na in their vegetative body. The N:P ratios found (fertilized = 27.5 and non-fertilized = 33.8) suggest that W. sintenisii may also be phosphorous limited as are lowland epiphytes. Fertilized plants had slightly longer inflorescences, and more flowers per inflorescence, than non-fertilized plants, but their flowers produced nectar in similar concentrations and quantities. Fertilized plants produced more seeds per fruit and per plant. Frequency of flowering in two consecutive years was higher for fertilized plants than for controls, suggesting that fertilized plants overcome the cost of reproduction more readily than non-fertilized plants. These results provide evidence that TMCF epiphytic bromeliads are nutrient limited like their lowland counterparts.     

Transgenerational effects of food availability on age at maturity and reproductive output in an asexual collembolan species

Transgenerational effects of environmental conditions can have several important ecological and evolutionary implications. We conducted a fully factorial experiment manipulating food availability across three generations in the collembolan Folsomia candida, a springtail species that inhabits soil and leaf litter environments which vary in resource availability. Maternal and grandmaternal food availability influenced age at maturity and reproductive output. These effects appear to be cumulative rather than adaptive transgenerational life-history adjustments. Such cumulative effects can profoundly influence eco-evolutionary dynamics in both stable and fluctuating environments.     

Reproductive ecology of three ocypodid crabs I. The influence of activity differences on reproductive traits

Reproductive traits of three ocypodid crabs,Scopimera globosa, Ilyoplax pusillus andMacrophthalmus japonicus, were compared.S. globosa andI. pusillus, inhabiting the upper-middle intertidal zone, produced 1–2 large broods per year, whereasM. japonicus, inhabiting the lower intertidal zone, produced 4–5 small broods per year. InS. globosa andI. pusillus, ovigerous females remained in their plugged burrows without feeding until their eggs hatched. On the other hand, femaleM. japonicus fed actively on surface mud while incubating. We concluded that few large broods may be advantageous in crab species that incubate in burrows, whereas continuous small broods may be advantageous in species that feed actively while incubating.     

Experimentally induced effects of water temperature and immersion time on reproductive output of bivalves in the Wadden Sea

Results of experiments are reported on the effects of water temperature and immersion time in winter on egg size and egg numbers in three intertidally living bivalves in the Dutch Wadden Sea, the Baltic tellin Macoma balthica, the common cockle Cerastoderma edule and the common mussel Mytilus edulis. Macoma (14–17 mm shell length) produced large eggs (diameter of 107 μm) in relatively small numbers (20 000–70 000) in early spring. Later in spring, Cerastoderma (28–33 mm shell length) produced smaller eggs (77 μm, excluding the surrounding jelly layer) in tenfold larger numbers (200 000–700 000). Mytilus (45–55 mm shell length) spawned even smaller eggs (72 μm) in high (but not easily assessed) numbers over a more extended period. In Macoma egg size was not affected by winter temperatures or immersion time. Effects of winter–spring temperatures and immersion time on egg size could be demonstrated in Cerastoderma. Smaller eggs were produced at the higher temperatures. Effects of immersion time were non-consistent: at lower water temperatures larger, but at higher temperatures smaller eggs were produced by animals kept at longer immersion times. In Mytilus, no temperature effects were observed. However, a longer immersion time resulted in larger eggs. In Macoma as well as in Cerastoderma significantly more eggs were produced at the lower temperature. Immersion time effects were most pronounced at the lower temperature, where more eggs were produced at the subtidal level than at the tidal level. At the higher water temperature differences between egg numbers produced at the two tidal levels were small. Just prior to spawning, egg numbers were strongly positively related to body mass at a certain shell length.     

破碎化次生林斑块面积及斑块隔离对大山雀繁殖成功的影响

研究了破碎化山地次生林中斑块面积及斑块隔离度对大山雀（Parus major)繁殖成功的影响,运用GPS定位系统确定了18块大,中,小3种类型的斑块及对照样点,观测了大山雀产第一枚卵时间,窝卵数,平均卵重,出雏量及雏鸟出飞量等生态指标,结果表明,斑块隔离度对大山雀繁殖成功没有影响,1999－2000年两年中,大山雀在连续分布次生林中的产卵时间平均早于各斑块中的产卵时间7．2d,各斑块间的产卵时间差异较小,连续分布次生林和较大面积斑块内的大山雀窝卵数略高于中,小面积斑块内的窝卵数,连续分布生林中的平均卵重最大,斑块面积对出雏量及雏鸟不量没有影响,中,小面积斑块内的巢损失率较高,最主要的原因是巢址竞争较激烈。     

Height and reproductive success in a cohort of british men

Two recent studies have shown a relationship between male height and number of offspring in contemporary developed-world populations. One of them argues as a result that directional selection for male tallness is both positive and unconstrained. This paper uses data from a large and socially representative national cohort of men who were born in Britain in March 1958. Taller men were less likely to be childless than shorter ones. They did not have a greater mean number of children. If anything, the pattern was the reverse, since men from higher socioeconomic groups tended to be taller and also to have smaller families. However, clear evidence was found that men who were taller than average were more likely to find a long-term partner, and also more likely to have several different long-term partners. This confirms the finding that tall men are considered more attractive and suggests that, in a noncontracepting environment, they would have more children. There is also evidence of stabilizing selection, since extremely tall men had an excess of health problems and an increased likelihood of childlessness. The conclusion is that male tallness has been selected for in recent human evolution but has been constrained by developmental factors and stabilizing selection on the extremely tall. The National Child Development Study is carried out by the researchers of the Centre for Longitudinal Studies, Institute of Education, London. The data are housed at the Data Archive of the University of Essex, from where they were obtained under license for the present study. I am grateful to Paul Preece for his assistance with data processing, and to Benjamin Campbell and two anonymous referees for invaluable advice which has, I hope, improved the quality of this paper. Daniel Nettle is lecturer in biological psychology at the Open University. With a first degree in psychology and a Ph.D. in biological anthropology, his work has been concerned with the application of evolutionary models to such topics as language (e.g., Linguistic Diversity, Oxford University Press, 1999) and individual differences (Strong Imagination, Oxford University Press, 2001).     

Gynodioecy as a possible populational strategy for increasing reproductive output

Summary Cytoplasmic male steriles occur regularly in wild populations of the annual crucifer Hirschfeldia incana Lagr.-Foss. in Israel. In these plants numbers of ovules per flower, numbers of seed per fruit, and total seed weights per plant are slightly higher than in the hermaphrodites with which they mate. Yet their frequencies in wild populations do not exceed 2–10 per cent. There are no signs of incipient dioecy. The species is self incompatible and no mechanisms to enforce outcrossing are needed. It is argued that in this and similar cases gynodioecy functions as a pollen saving measure. Precise pollen presentation in the flower renders some of the pollen redundant and facilitates its abolition in a sector of the population. It is possible that the enhanced seed fecundity of the pollenless sector stems from a greater availability of plant resources for seed production in the unisexual than in the bisexual seed parent.