Latitude, elevation and body mass variation in Andean passerine birds

• 1 Relationships between body mass and latitude, and body mass and elevation are examined in the assemblage of Andean passerine birds.
• 2 Across species, body mass is positively correlated with the mid‐point of the species elevational distribution, but there is no significant relationship between body mass and latitudinal range mid‐point.
• 3 When the assemblage is separated into Andean endemic and non‐endemic species, the former group shows a significant positive relationship between body mass and elevation, and the latter a significant positive relationship between body mass and latitude (‘Bergmann’s rule’). Andean endemic species exhibit Bergmann’s rule once elevation is controlled for using multiple regression.
• 4 These relationships are not a consequence of the phylogenetic non‐independence of species. All the effects shown are very weak, with latitude and elevation explaining only a few per cent of the variation in body mass. Relationships are strongest when phylogenetically controlled analyses are performed just within genera.
• 5 The implications of these results for the mechanistic understanding of Bergmann’s rule are discussed.

     

Energy and the tempo of evolution in amphibians

Aim The evolutionary speed hypothesis (ESH) attempts to explain global patterns of species richness on the basis that rates of molecular evolution and speciation in warmer climates have led to a greater accumulation of taxa at lower latitudes. A substantial alternative hypothesis to the ESH is the tropical conservatism hypothesis (TCH). However, recent tests of the TCH, using amphibians as the model taxon, have relied on the assumption that rates of molecular evolution are stable across latitudes and elevations. Here, we test for the first time for systematic variation in rates of molecular evolution across latitude and elevation among amphibians. Location The dataset is geographically diverse with samples from all continents except Antarctica and also from many of the earth's major tropical–warm temperate archipelagos. Methods We tested for substitution rate heterogeneity across climatically varying habitats with the mitochondrial RNA genes 12S and 16S. Thus, we report here on our findings for amphibians – a taxon whose phylogenetic and trophic contexts are remote from those previously tested – using genes that have also not been examined before. The study utilized paired contrasts of sister species (188 species across 18 families, including both caudates and anurans) that are spatially separated in either latitudinal or elevational dimensions. Results We found substantially faster substitution rates for species living in warmer habitats (P= 0.001–0.002) at both lower latitudes (P < 0.02) and lower elevations (P < 0.01). Main conclusions The consistency of these results with the previous studies that used quite different organisms – and in this instance also using different genes – suggests that this is a ubiquitous pattern in nature consistent with the predictions of the ESH. Recent tests of the TCH that, in estimating diversification rates, have relied on the assumption that DNA evolution occurs at a constant rate across latitudes and elevations, require reconsideration in light of the findings presented here. Our results indicate that greater caution is required when estimating dates of divergence using DNA sequence data.     

The estimation and evolution of hominin body mass

Body mass is a critical variable in many hominin evolutionary studies, with implications for reconstructing relative brain size, diet, locomotion, subsistence strategy, and social organization. We review methods that have been proposed for estimating body mass from true and trace fossils, consider their applicability in different contexts, and the appropriateness of different modern reference samples. Recently developed techniques based on a wider range of modern populations hold promise for providing more accurate estimates in earlier hominins, although uncertainties remain, particularly in non-Homo taxa. When these methods are applied to almost 300 Late Miocene through Late Pleistocene specimens, the resulting body mass estimates fall within a 25–60 kg range for early non-Homo taxa, increase in early Homo to about 50–90 kg, then remain constant until the Terminal Pleistocene, when they decline.     

Of mice and mammoths: evaluations of causal explanations for body size evolution in insular mammals

Aim We investigated the hypothesis that the insular body size of mammals results from selective forces whose influence varies with characteristics of the focal islands and the focal species, and with interactions among species (ecological displacement and release). Location Islands world‐wide. Methods We assembled data on the geographic characteristics (area, isolation, maximum elevation, latitude) and climate (annual averages and seasonality of temperature and precipitation) of islands, and on the ecological and morphological characteristics of focal species (number of mammalian competitors and predators, diet, body size of mainland reference populations) that were most relevant to our hypothesis (385 insular populations from 98 species of extant, non‐volant mammals across 248 islands). We used regression tree analyses to examine the hypothesized contextual importance of these factors in explaining variation in the insular body size of mammals. Results The results of regression tree analyses were consistent with predictions based on hypotheses of ecological release (more pronounced changes in body size on islands lacking mammalian competitors or predators), immigrant selection (more pronounced gigantism in small species inhabiting more isolated islands), thermoregulation and endurance during periods of climatic or environmental stress (more pronounced gigantism of small mammals on islands of higher latitudes or on those with colder and more seasonal climates), and resource subsidies (larger body size for mammals that utilize aquatic prey). The results, however, were not consistent with a prediction based on resource limitation and island area; that is, the insular body size of large mammals was not positively correlated with island area. Main conclusions These results support the hypothesis that the body size evolution of insular mammals is influenced by a combination of selective forces whose relative importance and nature of influence are contextual. While there may exist a theoretical optimal body size for mammals in general, the optimum for a particular insular population varies in a predictable manner with characteristics of the islands and the species, and with interactions among species. This study did, however, produce some unanticipated results that merit further study – patterns associated with Bergmann’s rule are amplified on islands, and the body size of small mammals appears to peak at intermediate and not maximum values of latitude and island isolation.     

A statistical test of unbiased evolution of body size in birds

Abstract.— Of the approximately 9500 bird species, the vast majority is small-bodied. That is a general feature of evolutionary lineages, also observed for instance in mammals and plants. The avian interspecific body size distribution is right-skewed even on a logarithmic scale. That has previously been interpreted as evidence that body size evolution has been biased. However, a procedure to test for unbiased evolution from the shape of body size distributions was lacking. In the present paper unbiased body size evolution is defined precisely, and a statistical test is developed based on Monte Carlo simulation of unbiased evolution. Application of the test to birds suggests that it is highly unlikely that avian body size evolution has been unbiased as defined. Several possible explanations for this result are discussed. A plausible explanation is that the general model of unbiased evolution assumes that population size and generation time do not affect the evolutionary variability of body size; that is, that micro- and macroevolution are decoupled, which theory suggests is not likely to be the case.     

Disentangling the effects of spring anomalies in climate and net primary production on body size of temperate songbirds

Body size is implicated in individual fitness and population dynamics. Mounting interest is being given to the effects of environmental change on body size, but the underlying mechanisms are poorly understood. We tested whether body size and body condition are related to ambient temperature (heat maintenance hypothesis), or/and explained by variations in primary production (food availability hypothesis) during the period of body growth in songbirds. We also explored whether annual population‐level variations of mean body size are due to changes of juvenile growth and/or size‐dependent mortality during the first year. For 41 species, from 257 sites across France, we tested for relationships between wing length (n = 107 193) or body condition (n = 82 022) and local anomalies in temperature, precipitation and net primary production (NDVI) during the breeding period, for juveniles and adults separately. Juvenile body size was best explained by primary production: wings were longer in years with locally high NDVI, but not shorter in years with low NDVI. Temperature showed a slightly positive effect. Body condition and adult wing length did not covary with any of the other tested variables. We found no evidence of climate‐driven size‐dependent mortality for the breeding season. In our temperate system, local climatic anomalies explained little of the body size variation. A large part of wing length variance was site‐specific, suggesting that avian size was more dependent on local drivers than global ones. Net primary production influenced juvenile size the most through effects on body growth. We suggest that, during the breeding season in temperate systems, thermoregulatory mechanisms are less involved in juvenile growth than food assimilation.     

Widespread rapid reductions in body size of adult salamanders in response to climate change

Reduction in body size is a major response to climate change, yet evidence in globally imperiled amphibians is lacking. Shifts in average population body size could indicate either plasticity in the growth response to changing climates through changes in allocation and energetics, or through selection for decreased size where energy is limiting. We compared historic and contemporary size measurements in 15 Plethodon species from 102 populations (9450 individuals) and found that six species exhibited significant reductions in body size over 55 years. Biophysical models, accounting for actual changes in moisture and air temperature over that period, showed a 7.1–7.9% increase in metabolic expenditure at three latitudes but showed no change in annual duration of activity. Reduced size was greatest at southern latitudes in regions experiencing the greatest drying and warming. Our results are consistent with a plastic response of body size to climate change through reductions in body size as mediated through increased metabolism. These rapid reductions in body size over the past few decades have significance for the susceptibility of amphibians to environmental change, and relevance for whether adaptation can keep pace with climate change in the future.     

Correlates of body mass evolution in primates

Body mass is undoubtedly central to the overall adaptive profile of any organism. Despite this, very little is known of what forces drive evolutionary changes in body mass and, consequently, shape patterns of body mass distribution exhibited by animal radiations. The search for factors that may influence evolutionary processes in general frequently focuses on environmental parameters such as climate change or interspecific competition. With respect to body mass, there is also the suggestion that evolutionary lineages may follow an inherent trend toward increased body mass, known as Cope's rule. The present paper investigates whether overall directional trends of body mass change, or correlations between patterns of body mass evolution and environmental factors have influenced the evolution of body mass in plesiadapiforms and primates. Analyses of the global fossil record of plesiadapiforms and primates suggest that the former did indeed follow an overall trend toward increased body mass compatible with the predictions of Cope's rule. In contrast, neither primates as a whole, nor a number of individual primate radiations (Adapiformes, Omomyiformes, and Anthropoidea), show any indication of overall directional patterns of body mass change. No correlations of primate body mass change with either the latitudinal distribution of fossil species, or with estimates of global temperature trends, were found. There is evidence, however, that direct competition between omomyiforms and adapiforms (the two main primate radiations known from the Paleogene) influenced processes of body mass evolution in omomyiforms.     

Avian eggshell thickness: scaling and maximum body mass in birds

The avian eggshell represents a highly evolved structure adapted to the physiological requirements of the embryo and the potential fracturing forces it is exposed to during incubation. Given its many roles, it is not surprising that the eggshell is also central to the current hypothesis about maximum avian body mass. Eggshell thickness ( L ) and strength has historically been scaled as a function of initial egg mass (IEM). However, maximum incubator mass (IM) is likely a better indicator of the forces the shell must be selected to withstand during incubation. We compare the results of analyses of L ² (an indicator of shell strength) as a function of IEM and IM. We conclude from IM scaling that megapode and kiwi eggshells are not thin but rather are thicker than expected and in general birds with a clutch size of 1 have thicker shells, and further, that reversed sexual dimorphism in the large, particularly extinct birds may be a strategy to avoid shell breakage during incubation of the largest eggs without creating a shell so thick as to inhibit hatching.     

The tempo and mode of evolution: body sizes of island mammals

The tempo and mode of body size evolution on islands are believed to be well known. It is thought that body size evolves relatively quickly on islands toward the mammalian modal value, thus generating extreme cases of size evolution and the island rule. Here, we tested both theories in a phylogenetically explicit context, by using two different species-level mammalian phylogenetic hypotheses limited to sister clades dichotomizing into an exclusively insular and an exclusively mainland daughter nodes. Taken as a whole, mammals were found to show a largely punctuational mode of size evolution. We found that, accounting for this, and regardless of the phylogeny used, size evolution on islands is no faster than on the continents. We compared different selection regimes using a set of Ornstein-Uhlenbeck models to examine the effects of insularity of the mode of evolution. The models strongly supported clade-specific selection regimes. Under this regime, however, an evolutionary model allowing insular species to evolve differently from their mainland relatives performs worse than a model that ignores insularity as a factor. Thus, insular taxa do not experience statistically different selection from their mainland relatives.     

The evolution of large body size in orangutans: A model for hominoid divergence

Recent Miocene fossil discoveries of large hominoids resemble orangutans. Since the evolution of large body size was functionally related to a powerful masticatory system in Miocene ape radiations, a better understanding of adaptations in extant orangutans will be informative of hominoid evolution. It is suggested here, based on the behavioral ecology of extant orangutans, that foraging energetics and large body size are tied to a dietary shift that provided access to and utilization of resources not generally available to other primates.     

The evolution of mammal body sizes: responses to Cenozoic climate change in North American mammals

Explanations for the evolution of body size in mammals have remained surprisingly elusive despite the central importance of body size in evolutionary biology. Here, we present a model which argues that the body sizes of Nearctic mammals were moulded by Cenozoic climate and vegetation changes. Following the early Eocene Climate Optimum, forests retreated and gave way to open woodland and savannah landscapes, followed later by grasslands. Many herbivores that radiated in these new landscapes underwent a switch from browsing to grazing associated with increased unguligrade cursoriality and body size, the latter driven by the energetics and constraints of cellulose digestion (fermentation). Carnivores also increased in size and digitigrade, cursorial capacity to occupy a size distribution allowing the capture of prey of the widest range of body sizes. With the emergence of larger, faster carnivores, plantigrade mammals were constrained from evolving to large body sizes and most remained smaller than 1 kg throughout the middle Cenozoic. We find no consistent support for either Cope's Rule or Bergmann's Rule in plantigrade mammals, the largest locomotor guild (n = 1186, 59% of species in the database). Some cold‐specialist plantigrade mammals, such as beavers and marmots, showed dramatic increases in body mass following the Miocene Climate Optimum which may, however, be partially explained by Bergmann's rule. This study reemphasizes the necessity of considering the evolutionary history and resultant form and function of mammalian morphotypes when attempting to understand contemporary mammalian body size distributions.     

Geographic gradients in body size: a clarification of Bergmann's rule

1997 marked the sesquicentenary of the publication by Carl Bergmann of the observation that, in general, large-bodied animal species tend to live further north than their small-bodied relatives. This has been dubbed Bergmann's rule in his honour. However, more than 150 years on, we appear to be little closer to a general understanding of the rule, or even to any consensus as to whether it exists. This is due in large part to confusion about the taxonomic level at which the rule is considered to operate, and to the conflation of pattern and mechanism. In this paper, we attempt to resolve this confusion by highlighting its sources, and by providing a definition of Bergmann's rule that is practical and useful, yet that retains the essential features of its original formulation. We conclude by briefly reviewing the mechanisms proposed to explain Bergmann's rule as we define it.     

Patterns of within-species body size variation of birds: strong evidence for Bergmann's rule

Aim The aim of this study is to test whether Bergmann's rule, a general intraspecific tendency towards larger body size in cooler areas and at higher latitudes, holds for birds throughout the world. Location This study includes information on species of birds from throughout the world. Methods I gathered data on body size variation from the literature and used two general meta‐analytical procedures to test the validity of Bergmann's rule in birds: a modified vote‐counting approach and calculation of overall effect sizes. Related species may show similar body size trends, thus I performed all analyses using nonphylogenetic and phylogenetic methods. I used tests of phylogenetic signal for each data set to decide which type of statistical analysis (nonphylogenetic or phylogenetic) was more appropriate. Results The majority of species of birds (76 of 100 species) are larger at higher latitudes, and in cooler areas (20 of 22 species). Birds show a grand mean correlation coefficient of +0.32 for body size and latitude, and ?0.81 for body size and temperature, both significant trends. Sedentary species show stronger body size trends in some, but not all, analyses. Neither males nor females consistently have stronger body size trends. Additionally, the strength of body size trends does not vary with latitude or body mass. Conclusions Bergmann's rule holds for birds throughout the world, regardless of whether temperature or latitude (as a proxy) is used. Previous studies have suggested that Bergmann's rule is stronger for sedentary than migratory species, males than females and temperate than tropical taxa. I did not find strong support for any of these as general themes for birds, although few studies of tropical taxa have been conducted. The processes responsible for Bergmann's rule remain somewhat of a black box; however, fasting endurance is probably a more important factor than the traditional hypothesis of heat conservation.     

气候变化对鸟类影响的研究进展

气候变化对生物多样性的影响已成为热点问题.本文以鸟类为研究对象,根据鸟类受气候变化影响的最新研究成果,综述了气候变化对鸟类的分布、物候和种群等方面的影响.结果表明,在气候变化影响下,鸟类分布向高纬度或高海拔区移动,速度比以往加快,繁殖地和非繁殖地的分布移动变化并不相同,并且多数分布范围缩小,物候期发生复杂变化,种群数量下降明显.文章还讨论了该领域主要的预测和评估方法,以及进化适应等生物因素对气候变化预测结果的影响,除了以往单一的相关性模型外,目前应用最多的是集成模型,而未来最具发展潜力的是机理模型.进化适应方面的研究近来取得新进展,证实了生物个体积极应对气候变化影响的事实,从而对人为模型预测的准确性带来挑战.文章最后进行了总结和展望,结合国外研究经验和我国实际情况,提出一些建议:由于气候变化的影响及其研究是长期性的,从而对鸟类的历史监测数据提出很高的要求,当前我国急需建立一套长期、全面和可靠的鸟类数据监测系统;此外,人们需要综合评估现有各种预测模型的可靠性,在此基础上探索新的研究方法.     

Fossils and living taxa agree on patterns of body mass evolution: a case study with Afrotheria

Most of life is extinct, so incorporating some fossil evidence into analyses of macroevolution is typically seen as necessary to understand the diversification of life and patterns of morphological evolution. Here we test the effects of inclusion of fossils in a study of the body size evolution of afrotherian mammals, a clade that includes the elephants, sea cows and elephant shrews. We find that the inclusion of fossil tips has little impact on analyses of body mass evolution; from a small ancestral size (approx. 100 g), there is a shift in rate and an increase in mass leading to the larger-bodied Paenungulata and Tubulidentata, regardless of whether fossils are included or excluded from analyses. For Afrotheria, the inclusion of fossils and morphological character data affect phylogenetic topology, but these differences have little impact upon patterns of body mass evolution and these body mass evolutionary patterns are consistent with the fossil record. The largest differences between our analyses result from the evolutionary model, not the addition of fossils. For some clades, extant-only analyses may be reliable to reconstruct body mass evolution, but the addition of fossils and careful model selection is likely to increase confidence and accuracy of reconstructed macroevolutionary patterns.     

Bergmann's rule in the ant lion Myrmeleon immaculatus DeGeer (Neuroptera: Myrmeleontidae): geographic variation in body size and heterozygosity

Aim Geographic variation in body size and heterozygosity were surveyed for discrete populations of the ant lion, Myrmeleon immaculatus DeGeer, collected from the central and northeastern United States. Location Collection sites were located in the central and eastern United States ranging from western Oklahoma to northern New York. Methods We collected 872 M. immaculatus larvae from thirty-four collecting sites. At each site, we randomly sampled ant lion pits and collected between fifteen and fifty-two larvae in total. Larvae were preserved in 95% ETOH for morphological analysis and frozen in a ?80°C freezer for protein electrophoresis. We measured the body size of eighty-five preserved adult M. immaculatus obtained from museum collections using head width as an indicator of body size. Five enzymes [GPI (glucose phosphate isomerase), MDH (malate dehydrogenase), PEP (peptidase), DIA (diaphorase) and SOD (superoxide dismutase)] were used in the heterozygosity analyses. Results Larval and adult body size increased with latitude, but decreased with elevation. Average heterozygosity, measured at five polymorphic loci, also increased significantly with latitude. Minimum temperature variance was the best predictor of body size, whereas precipitation and maximum temperature were the best predictors of heterozygosity. Populations were genetically differentiated from one another and showed a pattern of isolation by distance, as measured by Wright's F_st values and Nei's genetic distances. Main conclusions Sampling artifacts, heat conservation, character displacement, cell-size variation, density-dependent mortality, and differential dispersal probably cannot account for latitudinal variation in ant lion body size. Our results implicate the importance of diurnal photoperiod, which varies with latitude, but not with elevation. Because photoperiod often controls growth, diapause, and metamorphosis, it may be an important determinant of latitudinal clines in body size and life history of insects.     

Causes and consequences of variation in offspring body mass: meta‐analyses in birds and mammals

Early survival is highly variable and strongly influences observed population growth rates in most vertebrate populations. One of the major potential drivers of survival variation among juveniles is body mass. Heavy juveniles are better fed and have greater body reserves, and are thus assumed to survive better than light individuals. In spite of this, some studies have failed to detect an influence of body mass on offspring survival, questioning whether offspring body mass does indeed consistently influence juvenile survival, or whether this occurs in particular species/environments. Furthermore, the causes for variation in offspring mass are poorly understood, although maternal mass has often been reported to play a crucial role. To understand why offspring differ in body mass, and how this influences juvenile survival, we performed phylogenetically corrected meta‐analyses of both the relationship between offspring body mass and offspring survival in birds and mammals and the relationship between maternal mass and offspring mass in mammals. We found strong support for an overall positive effect of offspring body mass on survival, with a more pronounced influence in mammals than in birds. An increase of one standard deviation of body mass increased the odds of offspring survival by 71% in mammals and by 44% in birds. A cost of being too fat in birds in terms of flight performance might explain why body mass is a less reliable predictor of offspring survival in birds. We then looked for moderators explaining the among‐study differences reported in the intensity of this relationship. Surprisingly, sex did not influence the intensity of the offspring mass–survival relationship and phylogeny only accounted for a small proportion of observed variation in the intensity of that relationship. Among the potential factors that might affect the relationship between mass and survival in juveniles, only environmental conditions was influential in mammals. Offspring survival was most strongly influenced by body mass in captive populations and wild populations in the absence of predation. We also found support for the expected positive effect of maternal mass on offspring mass in mammals (r_pearson = 0.387). As body mass is a strong predictor of early survival, we expected heavier mothers to allocate more to their offspring, leading them to be heavier and so to have a higher survival. However, none of the potential factors we tested for variation in the maternal mass–offspring mass relationship had a detectable influence. Further studies should focus on linking these two relationships to determine whether a strong effect of offspring size on early survival is associated with a high correlation coefficient between maternal mass and offspring mass.     

The evolution of F1 postzygotic incompatibilities in birds

Abstract.— We analyzed the rate at which postzygotic incompatibilities accumulate in birds. Our purposes were to assess the role of intrinsic F₁ hybrid infertility and inviability in the speciation process, and to compare rates of loss of fertility and viability between the sexes. Among our sample more than half the crosses between species in the same genus produce fertile hybrids. Complete loss of F₁ hybrid fertility takes on the order of millions of years. Loss of F₁ hybrid viability occurs over longer timescales than fertility: some viable hybrids have been produced between taxa that appear to have been separated for more than 55 my. There is strong support for Haldane's rule, with very few examples where the male has lower fitness than the female. However, in contrast to Drosophila , fertility of the homogametic sex in the F₁ appears to be lost before viability of the heterogametic sex in the F₁. We conclude that the time span of loss of intrinsic hybrid fertility and viability is often, but not always, longer than the time to speciation. Premating isolation is an important mechanism maintaining reproductive isolation in birds. In addition, other factors causing postzygotic reproductive isolation such as ecological causes of hybrid unfitness, reduced mating success of hybrids, and genetic incompatibilities in the F₂s and backcrosses may often be involved in the speciation process.     

Atmospheric oxygen level and the evolution of insect body size

Insects are small relative to vertebrates, possibly owing to limitations or costs associated with their blind-ended tracheal respiratory system. The giant insects of the late Palaeozoic occurred when atmospheric PO₂ (aPO₂) was hyperoxic, supporting a role for oxygen in the evolution of insect body size. The paucity of the insect fossil record and the complex interactions between atmospheric oxygen level, organisms and their communities makes it impossible to definitively accept or reject the historical oxygen-size link, and multiple alternative hypotheses exist. However, a variety of recent empirical findings support a link between oxygen and insect size, including: (i) most insects develop smaller body sizes in hypoxia, and some develop and evolve larger sizes in hyperoxia; (ii) insects developmentally and evolutionarily reduce their proportional investment in the tracheal system when living in higher aPO₂, suggesting that there are significant costs associated with tracheal system structure and function; and (iii) larger insects invest more of their body in the tracheal system, potentially leading to greater effects of aPO₂ on larger insects. Together, these provide a wealth of plausible mechanisms by which tracheal oxygen delivery may be centrally involved in setting the relatively small size of insects and for hyperoxia-enabled Palaeozoic gigantism.