Optimum body size: effects of food quality and temperature,when reproductive growth rate is restricted,with examples from aphids

Most models of optimum energy partitioning predict variability in adult size, although not always explicitly. Increase in size is usually attributed to an increase in the growth rate or decline in mortality. The model presented shows that this may not always be the case. Even when mortality is kept constant in organisms with overlapping generations, a constraint on the maximum reproductive growth rate may lead, when the rate of overall growth increases, to either an increase or a decline in the optimum adult body size. It is shown that adult size could be a consequence of the differential responses of life history traits to changes in temperature and food quality. This is clearly advantageous for short lived organisms, like aphids, each generation of which only experience a very small part of the great seasonal range in conditions. This hypothesis complements Iwasa's (1991) explanation of the phenotypic plasticity observed in long lived organisms. The predictions are illustrated with empirical data from aphids. The model presented, which has been verified against a very large data set, indicates that for aphids the adult weight observed at a particular combination of temperature and food quality is that at which the population growth rate, r_m, is maximized. We conclude that predictions about adult size from models based on the partitioning of energy are more likely to apply to organisms that scramble for resources, i.e., “r” selected species. The size of organisms that contest for resources is more likely to be determined by competitive status and avoidance of natural enemies.     

温度对水螅种群增长和个体大小的影响

应用群体累积培养法,以枝角类为食物,在10℃、20℃和30℃三种温度下,研究了温度对水螅(Hydra sp．)种群密度、种群增长率和个体大小的影响。结果表明温度对水螅种群密度和个体体积由极显著影响。在培养初期,30℃下的水螅种群密度最大;而在培养后期,20℃下的水螅种群密度则显著大于30℃下的值;10℃下的值则始终最小。在同一温度下,种群增长率均与时间呈曲线相关,10℃、20℃、30℃的回归方程分别为：Y=0．000433X^2-0．00262X 0．00332、Y=-0．003367X^2 0．068335X-0．066489、Y=-0．018469X^2 0．188952X-0．030933。在研究范围内,30℃的水螅个体最小。     

The relationship between body size and population size in bromeliad tank faunas

Relationships between body size and abundance in collections of animals from the tanks of 73 bromeliads belonging to five species were analysed. Unlike data in previously published studies on this relationship, these collections of species are not taxonomically restricted and represent complete communities over the macroscopic range of organisms. There is no overall tendency for there to be a positive or negative relationship between population abundance and body size of morphospecies. We can find no evidence that body size-abundance patterns are triangular in complete communities. However, there is weak evidence that the relationship in the aquatic subsets of those communities may have some underlying triangularity, with medium-sized species having the largest populations.     

The relationship between genome size, development rate, and body size in copepods

Freshwater cyclopoid copepods exhibit at least a fivefold range in somatic genome size and a mechanism, chromatin diminution, which could account for much of this interspecific variation. These attributes suggest that copepods are well suited to studies of genome size evolution. We tested the nucleotypic hypothesis of genome size evolution, which poses that variation in genome size is adaptive due to the bulk effects of both coding and noncoding DNA on cell size and division rates, and their correlates. We found a significant inverse correlation between genome size and developmental (growth) rate in five freshwater cyclopoid species at three temperatures. That is, species with smaller genomes developed faster. Species with smaller genomes had significantly smaller bodies at 22 °C, but not at cooler and warmer temperatures. Species with smaller genomes developed faster at all three temperatures, but had smaller bodies only at 22 °C. We propose a model of life history evolution that adds genome size and cell cycle dynamics to the suite of characters on which selection may act to mold life histories and to influence the distribution of traits among different habitats.     

Effect of ration and body size on the energy budget of juvenile white sturgeon

Growth and energy budget were measured for three sizes (2.4, 11.1 and 22.5 g) of juvenile white sturgeon Acipenser transmontanus held at 18.5° C and fed tubificid worms at different levels ranging from starvation to ad libitum . For each size-class, specific growth rate increased linearly with increasing ration, and conversion efficiency was highest at the maximum ration. Growth rate decreased with increasing fish size at the maximum ration, but increased with size at each restricted ration. Conversion efficiency increased with increasing ration for each size-class and was usually highest at the maximum ration. Faecal production accounted for 3.2–5.2% of food energy. The proportion of food energy lost in nitrogenous excretion decreased with increasing ration. With increases in ration, the allocation of metabolizable energy to metabolism decreased, while that to growth increased. Fish size had no significant effect on the allocation of metabolizable energy to metabolism or growth. At the maximum ration, on average 64.9% of metabolizable energy was spent on metabolism, and 35.1% on growth.     

Natural enemy ravine revisited: the importance of sample size for determining population growth

Abstract. 1. The population growth of three aphid species, Metopolophium dirhodum (Walker), Rhopalosiphum padi (L.), and Sitobion avenae (F.), on winter wheat, was analysed by regression. The calculations were based on censuses of aphids made in 268 plots at 3- or 7-day intervals for 10 years on leaves and 6 years on ears. The calculations were made separately for each plot each year, then repeated on the pooled data from all plots monitored in a year.
2. At the level of individual plots, no population growth was detected at very low densities. At high densities, the populations grew exponentially and the growth rates did not decrease with increasing aphid density.
3. Significant growth was always detected in the pooled data. These growth rates decreased significantly at the highest densities. Field estimates of the intrinsic rate of increase derived from these data ranged from 0.010 to 0.026 for M. dirhodum , 0.0071–0.011 for R. padi , and 0.00078–0.0061 and 0.0015–0.13 for S. avenae , on leaves and ears respectively .
4. The apparent lack of growth in the individual plots at low densities is attributable to small sample size. It is concluded that the natural enemy ravine in the population dynamics of cereal aphids, identified by Southwood and Comins (1976), is a consequence of low population densities at which population increase is undetectable unless very large samples are taken.     

The scaling of body size and mass in a host-parasitoid association: influence of host species and stage

We describe the allometry of body mass and body size as measured by hind-tibia length in males of Monoctonus paulensis (Ashmead) (Hymenoptera: Braconidae, Aphidiinae), a solitary parasitoid of aphids. To assess the influence of host quality on allometric relationships, we reared parasitoids on second and fourth nymphal instars of four different aphid species, Acyrthosiphon pisum (Harris), Macrosiphum creelii Davis, Myzus persicae (Sulzer) and Sitobion avenae (F.), under controlled conditions in the laboratory. Dry mass was positively correlated with hind-tibia length, and could be predicted from it, in unparasitized aphids, in aphid mummies containing parasitoid pupae, and in the parasitoid. The reduced-major-axis scaling exponents for the regression of dry mass on hind-tibia length were species-specific in aphids, reflecting differences in volume and shape between species. In mummified aphids, the stage at death influenced the size/mass relationship. In males of M. paulensis, the allometric exponent varied between parasitoids developing in different kinds of host. Individuals developing in pea aphid were absolutely larger in dry mass as well as proportionately larger relative to their hind-tibia length. We discuss the allometry of body size and body mass in relation to parasitoid fitness.     

Empirical evidence for an optimal body size in snakes

Abstract The concept of optimal size has been invoked to explain patterns in body size of terrestrial mammals. However, the generality of this phenomenon has not been tested with similarly complete data from other taxonomic groups. In this study we describe three statistical patterns of body size in snakes, all of which indicate an optimal length of 1.0 m. First, a distribution of largest body lengths of 618 snake species had a single mode at 1.0 m. Second, we found a positive relationship between the size of the largest member of an island snake assemblage and island area and a negative relationship between the size of the smallest member of an island snake assemblage and island area. Best-fit lines through these data cross at a point corresponding to 1.0 m in body length, the presumed optimal size for a one-species island. Third, mainland snake species smaller than 1.0 m become larger on islands whereas those larger than 1.0 m become smaller on islands. The observation that all three analyses converge on a common body size is concordant with patterns observed in mammals and partial analyses of four other disparate animal clades. Because snakes differ so strikingly from mammals (ectotherms, gape-limited predators, elongate body shape) the concordant patterns of these two groups provide strong evidence for the evolution of an optimal body size within independent monophyletic groups. However, snakes differ from other taxonomic groups that have been studied in exhibiting a body size distribution that is not obviously skewed in either direction. We suggest that idiosyncratic features of the natural history of ectotherms allow relatively unconstrained distributions of body size whereas physiological limitations of endotherms constrain distributions of body size to a right skew.     

Optimal diving behaviour for foraging in relation to body size

Overall, large animals dive longer and deeper than small animals; however, after the difference in body size is taken into account, smaller divers often tend to make relatively longer dives. Neither physiological nor theoretical explanations have been provided for this paradox. This paper develops an optimal foraging diving model to demonstrate the effect of body size on diving behaviour, and discusses optimal diving behaviour in relation to body size. The general features of the results are: (1) smaller divers should rely more heavily on anaerobic respiration, (2) larger divers should not always make longer dives than smaller divers, and (3) an optimal body size exists for each diving depth. These results explain the relatively greater diving ability observed in smaller divers, and suggest that if the vertical distribution of prey in the water column is patchy, there is opportunity for a population of diving animals to occupy habitat niches related to body size.     

On size and area: Patterns of mammalian body size extremes across landmasses

We describe a biogeographic pattern in which mammalian body size extremes scale with landmass area. The relationship between the largest and the smallest mammal species found on different landbridge islands, mountaintops and continents shows that the size of the largest species increases, while that of the smallest species decreases, with increase in the area of the landmass. We offer two possible explanations: (1) that the pattern is the result of sampling artefacts, which we call the ‘statistical artefact hypothesis’, or (2) that the pattern is the result of processes related to the way body size affects the number of individuals that a particular species can pack in a given area, which we call the ‘area-scaling hypothesis’. Our results point out that the pattern is not a statistical artefact resulting from random sampling, but can be explained by considering the scaling of individual space requirements and its effect on population survival on landmasses of different area. This revised version was published online in July 2006 with corrections to the Cover Date.     

区域性农田景观格局对棉蚜种群数量的生态学效应

农田景观格局的变化显著影响害虫的发生和危害,不同景观格局会对害虫的种群数量产生不同程度的影响,因而明确农田景观格局对害虫的生态学效应是控制害虫的重要前提之一。以山东省的棉花种植区为研究区域,选取14个典型的尽量临近不同土地覆盖类型的棉花生产县,通过卫星遥感影像和土地覆盖分类数据综合分析获得取样县/区的景观因子指数,并系统调查对应县/区的棉蚜种群数量。省级范围的大空间尺度下分析景观组成、景观构成和景观结构等多因子分别与棉田中苗蚜和伏蚜种群的相关性。研究结果表明棉蚜的种群数量与景观格局有密切的关系,且棉蚜发生的两个时期苗蚜和伏蚜对景观因子的响应特征并不完全一致。苗蚜的种群数量与景观总面积、耕地的分形指数、县域范围的蔓延度和县域范围的回旋半径等呈显著正相关,与Simpson多样性指标呈显著负相关;伏蚜的种群数量与斑块丰富密度、居住工业交通的蔓延度等呈显著正相关。总之,苗蚜和伏蚜对景观的蔓延度(形)响应基本上是一致的,景观的破碎化程度越小,伏蚜和苗蚜发生越重。而苗蚜和伏蚜对景观多样性(质)的响应不一致,景观多样性高的农田景观不利于苗蚜的发生,对伏蚜的影响不显著;而丰富度密度有助于伏蚜的发生,却对苗蚜没有显著影响。这一结果显示了农业害虫的不同发生时期对农田景观格局响应的复杂性。     

Genome size as a determinant of growth and life-history traits in crustaceans

Genome size varies tremendously both within and among taxa, and strong correlations between genome size and various physiological and ecological attributes suggest that genome size is a key trait of organisms, yet the causalities remains vague. In the present study, we tested how genome size is related to key physiological and ecological properties in five large orders of crustaceans: Decapoda, Cladocera, Amphipoda, Calanoida, and Cyclopoida. These span a wide range in sizes, habitats and life-history traits. To some extent, genome size reflected phylogenetic footprints but, generally, a very wide range in genome size was found within all orders. Genome size was positively correlated with body size in Amphipoda, Cladocera, and Copepoda, but not for Decapoda in general. This could indicate that the evolution of body size occurs mainly by changing cell size for the three first orders, whereas it is more attributed to cell numbers for Decapoda. Cladocera, with direct development and a high growth rate, have minute genomes compared to copepods that possess a more complex life history, whereas, within Decapoda and Amphipoda, developmental complexity is not related to genome size. The present study suggests that, within the crustaceans, selection for a wide variety of life-history strategies has led to widely different genome sizes.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 393–399.     

Proximate causes of altitudinal differences in body size in an agamid lizard

Body size is directly linked to key life history traits such as growth, fecundity, and survivorship. Identifying the causes of body size variation is a critical task in ecological and evolutionary research. Body size variation along altitudinal gradients has received considerable attention; however, the underlying mechanisms are poorly understood. Here, we compared the growth rate and age structure of toad‐headed lizards (Phrynocephalus vlangalii) from two populations found at different elevations in the Qinghai‐Tibetan Plateau. We used mark‐recapture and skeletochronological analysis to identify the potential proximate causes of altitudinal variation in body size. Lizards from the high‐elevation site had higher growth rates and attained slightly larger adult body sizes than lizards from the low‐elevation site. However, newborns produced by high‐elevation females were smaller than those by low‐elevation females. Von Bertalanffy growth estimates predicted high‐elevation individuals would reach sexual maturity at an earlier age and have a lower mean age than low‐elevation individuals. Relatively lower mean age for the high‐elevation population was confirmed using the skeletochronological analysis. These results support the prediction that a larger adult body size of high‐elevation P. vlangalii results from higher growth rates, associated with higher resource availability.     

Heterogeneity of genetic architecture of body size traits in a free‐living population

Knowledge of the underlying genetic architecture of quantitative traits could aid in understanding how they evolve. In wild populations, it is still largely unknown whether complex traits are polygenic or influenced by few loci with major effect, due to often small sample sizes and low resolution of marker panels. Here, we examine the genetic architecture of five adult body size traits in a free‐living population of Soay sheep on St Kilda using 37 037 polymorphic SNPs. Two traits (jaw and weight) show classical signs of a polygenic trait: the proportion of variance explained by a chromosome was proportional to its length, multiple chromosomes and genomic regions explained significant amounts of phenotypic variance, but no SNPs were associated with trait variance when using GWAS. In comparison, genetic variance for leg length traits (foreleg, hindleg and metacarpal) was disproportionately explained by two SNPs on chromosomes 16 (s23172.1) and 19 (s74894.1), which each explained >10% of the additive genetic variance. After controlling for environmental differences, females heterozygous for s74894.1 produced more lambs and recruits during their lifetime than females homozygous for the common allele conferring long legs. We also demonstrate that alleles conferring shorter legs have likely entered the population through a historic admixture event with the Dunface sheep. In summary, we show that different proxies for body size can have very different genetic architecture and that dense SNP helps in understanding both the mode of selection and the evolutionary history at loci underlying quantitative traits in natural populations.     

A test of the reproductive cost hypothesis for sexual size dimorphism in Yarrow's spiny lizard Sceloporus jarrovii

1. Trade-offs between reproduction and growth are central assumptions of life-history theory, but their implications for sexual size dimorphism (SSD) are poorly understood. 2. Adult male Yarrow's spiny lizards Sceloporus jarrovii average 10% larger than adult females. In a low-altitude (1700 m) population, this SSD develops because males grow more quickly than females during the first year of life, particularly during the first female reproductive season. This study tests the hypothesis that SSD develops because female growth is constrained by energetic costs of reproduction. 3. To test for a growth cost of reproduction, I compared growth rates of free-living females that differed, either naturally or experimentally, in reproductive status. Females that naturally delayed reproduction until their second year grew more quickly than females that reproduced as yearlings, and ovariectomized yearlings grew more quickly and to larger sizes than reproductive controls. 4. To determine whether SSD develops in the absence of this inferred reproductive cost, I also studied a high-altitude (2500 m) population in which all females delay reproduction until their second year. Sex differences in growth trajectories were similar to those observed at low altitude, such that males averaged 10% larger than females even prior to female reproduction. 5. Although female growth may be constrained by reproduction, multiple lines of evidence indicate that this cost is insufficient to explain the full magnitude of SSD in S. jarrovii. First, differences in growth of reproductive and nonreproductive females are not observed until the final month of gestation, by which time SSD is already well developed. Second, the growth benefit accruing from experimental inhibition of reproduction accounts for only 32% of the natural sex difference in body size. Finally, SSD develops well in advance of female reproduction in a high-altitude population with delayed maturation.     

One size does not fit all: no evidence for an optimal body size on islands

Aim Optimal body size theories predict that large clades have a single, optimal, body size that serves as an evolutionary attractor, with the full body size spectrum of a clade resulting from interspecific competition. Because interspecific competition is believed to be reduced on islands, such theories predict that insular animals should be closer to the optimal size than mainland animals. We test the resulting prediction that insular clade members should therefore have narrower body size ranges than their mainland relatives. Location World‐wide. Methods We used body sizes and a phylogenetic tree of 4004 mammal species, including more than 200 species that went extinct since the last ice age. We tested, in a phylogenetically explicit framework, whether insular taxa converge on an optimal size and whether insular clades have narrow size ranges. Results We found no support for any of the predictions of the optimal size theory. No specific size serves as an evolutionary attractor. We did find consistent evidence that large (> 10 kg) mammals grow smaller on islands. Smaller species, however, show no consistent tendency to either dwarf or grow larger on islands. Size ranges of insular taxa are not narrower than expected by chance given the number of species in their clades, nor are they narrower than the size ranges of their mainland sister clades – despite insular clade members showing strong phylogenetic clustering. Main conclusions The concept of a single optimal body size is not supported by the data that were thought most likely to show it. We reject the notion that inclusive clades evolve towards a body‐plan‐specific optimum.     

Crop size as an index of chick provisioning in the Greater Flamingo Phoenicopterus roseus

After being fed by their parents, Greater Flamingo chicks store food in their crops, which protrude outwards. We allocated the crop profiles of chicks to four categories to assess the relationship between body mass and crop profile variation, and so determine whether crop size can be used as an accurate index of the amount of food ingested, and to determine the timing and frequency of provisioning. We registered changes in body mass and crop fullness in eight chicks captured with turgid crops and kept in captivity until constant mass was achieved. The meal mass ingested by the chicks during each parental feeding was around 18% of net chick mass and varied greatly with crop profile. Mean transition times between the four crop profile categories ranged from 6 to 14 h. Between 1998 and 2009, 34% of chicks caught for ringing in a breeding colony had empty crops. From crop profiles recorded during the handling of chicks, it was estimated that approximately one‐third of the chicks were fed in the evening and another third during the night. Our results have implications for the estimation of body condition indexes because body mass should be free of the influence of the mass of the food in the crop.