THE MICRO AND MACRO IN BODY SIZE EVOLUTION

The diversity of body sizes of organisms has traditionally been explained in terms of microevolutionary processes: natural selection owing to differential fitness of individual organisms, or to macroevolutionary processes: species selection owing to the differential proliferation of phylogenetic lineages. Data for terrestrial mammals and birds indicate that even on a logarithmic scale frequency distributions of body mass among species are significantly skewed towards larger sizes. We used simulation models to evaluate the extent to which macro- and microevolutionary processes are sufficient to explain these distributions. Simulations of a purely cladogenetic process with no bias in extinction or speciation rates for different body sizes did not produce skewed log body mass distributions. Simulations that included size-biased extinction rates, especially those that incorporated anagenetic size change within species between speciation and extinction events, regularly produced skewed distributions. We conclude that although cladogenetic processes probably play a significant role in body size evolution, there must also be a significant anagenetic component. The regular variation in the form of mammalian body size distributions among different-sized islands and continents suggests that environmental conditions, operating through both macro- and microevolutionary processes, determine to a large extent the diversification of body sizes within faunas. Macroevolution is not decoupled from microevolution.     

Body Size Is a Significant Predictor of Congruency in Species Richness Patterns: A Meta-Analysis of Aquatic Studies

Biodiversity losses over the next century are predicted to result in alterations of ecosystem functions that are on par with other major drivers of global change. Given the seriousness of this issue, there is a need to effectively monitor global biodiversity. Because performing biodiversity censuses of all taxonomic groups is prohibitively costly, indicator groups have been studied to estimate the biodiversity of different taxonomic groups. Quantifying cross-taxon congruence is a method of evaluating the assumption that the diversity of one taxonomic group can be used to predict the diversity of another. To improve the predictive ability of cross-taxon congruence in aquatic ecosystems, we evaluated whether body size, measured as the ratio of average body length between organismal groups, is a significant predictor of their cross-taxon biodiversity congruence. To test this hypothesis, we searched the published literature and screened for studies that used species richness correlations as their metric of cross-taxon congruence. We extracted 96 correlation coefficients from 16 studies, which encompassed 784 inland water bodies. With these correlation coefficients, we conducted a categorical meta-analysis, grouping data based on the body size ratio of organisms. Our results showed that cross-taxon congruence is variable among sites and between different groups (r values ranging between −0.53 to 0.88). In addition, our quantitative meta-analysis demonstrated that organisms most similar in body size showed stronger species richness correlations than organisms which differed increasingly in size (r_adj ² = 0.94, p = 0.02). We propose that future studies applying biodiversity indicators in aquatic ecosystems consider functional traits such as body size, so as to increase their success at predicting the biodiversity of taxonomic groups where cost-effective conservation tools are needed.     

A Multi-Clade Test Supports the Intermediate Dispersal Model of Biogeography

Background

Biogeography models typically focus on explaining patterns through island properties, such as size, complexity, age, and isolation. Such models explain variation in the richness of island biotas. Properties of the organisms themselves, such as their size, age, and dispersal abilities, in turn may explain which organisms come to occupy, and diversify across island archipelagos. Here, we restate and test the intermediate dispersal model (IDM) predicting peak diversity in clades of relatively intermediate dispersers.

Methodology

We test the model through a review of terrestrial and freshwater organisms in the western Indian Ocean examining the correlation among species richness and three potential explanatory variables: dispersal ability quantified as the number of estimated dispersal events, average body size for animals, and clade age.

Conclusions

Our study supports the IDM with dispersal ability being the best predictor of regional diversity among the explored variables. We find a weaker relationship between diversity and clade age, but not body size. Principally, we find that richness strongly and positively correlates with dispersal ability in poor to good dispersers while a prior study found a strong decrease in richness with increased dispersal ability among excellent dispersers. Both studies therefore support the intermediate dispersal model, especially when considered together. We note that many additional variables not here considered are at play. For example, some taxa may lose dispersal ability subsequent to island colonization and some poor dispersers have reached high diversity through within island radiations. Nevertheless, our findings highlight the fundamental importance of dispersal ability in explaining patterns of biodiversity generation across islands.     

Body mass evolution and diversification within horses (family Equidae)

Horses (family Equidae) are a classic example of adaptive radiation, exhibiting a nearly 60‐fold increase in maximum body mass and a peak taxonomic diversity of nearly 100 species across four continents. Such patterns are commonly attributed to niche competition, in which increased taxonomic diversity drives increased size disparity. However, neutral processes, such as macroevolutionary ‘diffusion’, can produce similar increases in disparity without increased diversity. Using a comprehensive database of Equidae species size estimates and a common mathematical framework, we measure the contributions of diversity‐driven and diffusion‐driven mechanisms for increased disparity during the Equidae radiation. We find that more than 90% of changes in size disparity are attributable to diffusion alone. These results clarify the role of species competition in body size evolution, indicate that morphological disparity and species diversity may be only weakly coupled in general, and demonstrate that large species may evolve from neutral macroevolutionary diffusion processes alone.     

Are most species small? Not within species-level phylogenies.

The robust macro-ecological observation that there are more small-bodied species implies that small-bodied organisms have experienced elevated net rates of diversification. We investigate the role of body size in creating non-random differences in rates of cladogenesis using a set of 38 species-level phylogenies drawn from a range of animal groups. We use independent contrasts to explore the relationship between body size and species richness within individual phylogenies and across related sets of phylogenies. We also carry out a meta-analysis looking for associations between body size and species richness across the taxa. We find little evidence for increased cladogenesis among small-bodied organisms within taxa, and no evidence for any consistent differences between taxa. We explore possible explanations for the inconsistency of our findings with macro-ecological patterns.     

Habitat disturbance selects against both small and large species across varying climates

Global extinction drivers, including habitat disturbance and climate change, are thought to affect larger species more than smaller species. However, it is unclear if such drivers interact to affect assemblage body size distributions. We asked how these two key global change drivers differentially affect the interspecific size distributions of ants, one of the most abundant and ubiquitous animal groups on earth. We also asked whether there is evidence of synergistic interactions and whether effects are related to species’ trophic roles. We generated a global dataset on ant body size from 333 local ant assemblages collected by the authors across a broad range of climates and in disturbed and undisturbed habitats. We used head length (range: 0.22–4.55 mm) as a surrogate of body size and classified species to trophic groups. We used generalized linear models to test whether body size distributions changed with climate and disturbance, independent of species richness. Our analysis yielded three key results: 1) climate and disturbance showed independent associations with body size; 2) assemblages included more small species in warmer climates and fewer large species in wet climates; and 3) both the largest and smallest species were absent from disturbed ecosystems, with predators most affected in both cases. Our results indicate that temperature, precipitation and disturbance have differing effects on the body size distributions of local communities, with no evidence of synergistic interactions. Further, both large and small predators may be vulnerable to global change, particularly through habitat disturbance.     

Abundance, diversity and body size: patterns from a grassland arthropod community

1. The empirical relationships among body size, species richness and number of individuals may give insight into the factors controlling species diversity and the relative abundances of species. To determine these relationships, we sampled the arthropods of grasslands and savannahs at Cedar Creek, MN using sweep nets (90 525 individuals of 1225 species) and pitfall traps (12 721 individuals of 92 species). Specimens were identified, enumerated and measured to determine body size.
2. Both overall and within abundant taxonomic orders, species richness and numbers of individuals peaked at body sizes intermediate for each group. Evolution could create unimodal diversity patterns by random diversification around an ancestral body size or from size-dependent fitness differences. Local processes such as competition or predation could also create unimodal diversity distributions.
3. The average body size of a species depended significantly on its taxonomic order, but on contemporary trophic role only within the context of taxonomic order.
4. Species richness ( S _i) within size classes was related to the number of individuals ( I _i) as S _i =  I i^0·5. This relationship held across a 100 000-fold range of body sizes. Within size classes, abundance distributions of size classes were all similar power functions. A general rule of resource division, together with similar minimum population sizes, is sufficient to generate the relationship between species richness and number of individuals.
5. Smaller bodied species had slightly shallower abundance distributions and may, in general, persist at lower densities than larger species.
6. Our results suggest there may be fewer undescribed small arthropod species than previously thought and that most undescribed species will be smaller than arthropods.     

Similarity of mammalian body size across the taxonomic hierarchy and across space and time

Although it is commonly assumed that closely related animals are similar in body size, the degree of similarity has not been examined across the taxonomic hierarchy. Moreover, little is known about the variation or consistency of body size patterns across geographic space or evolutionary time. Here, we draw from a data set of terrestrial, nonvolant mammals to quantify and compare patterns across the body size spectrum, the taxonomic hierarchy, continental space, and evolutionary time. We employ a variety of statistical techniques including "sib-sib" regression, phylogenetic autocorrelation, and nested ANOVA. We find an extremely high resemblance (heritability) of size among congeneric species for mammals over approximately 18 g; the result is consistent across the size spectrum. However, there is no significant relationship among the body sizes of congeneric species for mammals under approximately 18 g. We suspect that life-history and ecological parameters are so tightly constrained by allometry at diminutive size that animals can only adapt to novel ecological conditions by modifying body size. The overall distributions of size for each continental fauna and for the most diverse orders are quantitatively similar for North America, South America, and Africa, despite virtually no overlap in species composition. Differences in ordinal composition appear to account for quantitative differences between continents. For most mammalian orders, body size is highly conserved, although there is extensive overlap at all levels of the taxonomic hierarchy. The body size distribution for terrestrial mammals apparently was established early in the Tertiary, and it has remained remarkably constant over the past 50 Ma and across the major continents. Lineages have diversified in size to exploit environmental opportunities but only within limits set by allometric, ecological, and evolutionary constraints.     

Body Size Diversity and Frequency Distributions of Neotropical Cichlid Fishes (Cichliformes: Cichlidae: Cichlinae)

Body size is an important correlate of life history, ecology and distribution of species. Despite this, very little is known about body size evolution in fishes, particularly freshwater fishes of the Neotropics where species and body size diversity are relatively high. Phylogenetic history and body size data were used to explore body size frequency distributions in Neotropical cichlids, a broadly distributed and ecologically diverse group of fishes that is highly representative of body size diversity in Neotropical freshwater fishes. We test for divergence, phylogenetic autocorrelation and among-clade partitioning of body size space. Neotropical cichlids show low phylogenetic autocorrelation and divergence within and among taxonomic levels. Three distinct regions of body size space were identified from body size frequency distributions at various taxonomic levels corresponding to subclades of the most diverse tribe, Geophagini. These regions suggest that lineages may be evolving towards particular size optima that may be tied to specific ecological roles. The diversification of Geophagini appears to constrain the evolution of body size among other Neotropical cichlid lineages; non-Geophagini clades show lower species-richness in body size regions shared with Geophagini. Neotropical cichlid genera show less divergence and extreme body size than expected within and among tribes. Body size divergence among species may instead be present or linked to ecology at the community assembly scale.     

Functional differences in echolocation call design in an adaptive radiation of bats

All organisms have specialized systems to sense their environment. Most bat species use echolocation for navigation and foraging, but which and how ecological factors shaped echolocation call diversity remains unclear for the most diverse clades, including the adaptive radiation of neotropical leaf‐nosed bats (Phyllostomidae). This is because phyllostomids emit low‐intensity echolocation calls and many inhabit dense forests, leading to low representation in acoustic surveys. We present a field‐collected, echolocation call dataset spanning 35 species and all phyllostomid dietary guilds. We analyze these data under a phylogenetic framework to test the hypothesis that echolocation call design and parameters are specialized for the acoustic demands of different diets, and investigate the contributions of phylogeny and body size to echolocation call diversity. We further link call parameters to dietary ecology by contrasting minimum detectable prey size estimates (MDPSE) across species. We find phylogeny and body size explain a substantial proportion of echolocation call parameter diversity, but most species can be correctly assigned to taxonomic (61%) or functional (77%) dietary guilds based on call parameters. This suggests a degree of acoustic ecological specialization, albeit with interspecific similarities in call structure. Theoretical MDPSE are greatest for omnivores and smallest for insectivores. Omnivores significantly differ from other dietary guilds in MDPSE when phylogeny is not considered, but there are no differences among taxonomic dietary guilds within a phylogenetic context. Similarly, predators of non‐mobile/non‐evasive prey and predators of mobile/evasive prey differ in estimated MDPSE when phylogeny is not considered. Phyllostomid echolocation call structure may be primarily specialized for overcoming acoustic challenges of foraging in dense habitats, and then secondarily specialized for the detection of food items according to functional dietary guilds. Our results give insight into the possible ecological mechanisms shaping the diversity of sensory systems, and their reciprocal influence on resource use.     

Body size distributions of large Costa Rican dry forest moths and the underlying relationship between plant and pollinator morphology

There has been much recent interest in explaining patterns of body size variation within species assemblages. One observation is that frequency distributions of species' body size commonly exhibit a right-skew, even on a logarithmic scale. Here we examine the species' body size distributions in two assemblages of large Costa Rican moths. We find that neither adult Sphingidae or Saturniidae exhibit the classic log right-skewed pattern. Furthermore, the species' body size distributions in these two groups are markedly different, which we suggest is a result of differential selective pressures related to resource and mate acquisition. For Sphingidae, we show (1) that body size is positively correlated with tongue length, and (2) that the distribution of sphingid body sizes/tongue lengths closely matches the distribution of flower corolla tube depths in sphingid-pollinated plants. Thus, morphological fitting between plants and pollinators seems to underlie the species' body size distribution of this sphingid assemblage. We discuss the significance of these results in the context of current theory on mechanisms driving species' body size distributions. Finally, we present an evolutionary hypothesis for the diversity of body sizes seen in this sphingid assemblage related to reciprocal interactions between plants and pollinators. This hypothesis can be tested within a rigorous phylogenetic framework, although a systematic phylogenetic analysis of Neotropical Sphingidae does not currently exist.     

Do small animals have a biogeography?

It has been stated that small organisms do not have barriers for distribution and will not show biogeographic discreteness. General models for size-mediated biogeographies establish a transition region between ubiquitous dispersal and restricted biogeography at about 1–10 mm. We tested patterns of distribution versus size with water mites, a group of freshwater organisms with sizes between 300 μm and 10 mm.We compiled a list of all known water mite species for Sierra del Guadarrama (a mountain range in the centre of the Iberian Peninsula) from different authors and our own studies in the area. Recorded habitats include lotic, lentic and interstitial environments. Species body size and world distribution were drawn from our work and published specialized taxonomic literature. The null hypothesis was that distribution is size-independent. The relationship between distribution and size was approached via analysis of variance and between size and habitat via logistic regression. Contrary to expectations, there is no special relationship between water mite size and area size distribution. On the other hand, water mite size is differentially distributed among habitats, although this ecological sorting is very weak. Larger water mites are more common in lentic habitats and smaller water mites in lotic habitats. Size-dependent distribution in which small organisms tend to be cosmopolitan breaks down when the particular biology comes into play. Water mites do not fit a previously proposed size-dependent biogeographical distribution, and are in accordance with similar data published on Tardigrada, Rotifera, Gastrotricha and the like.     

Latitudinal gradients in diversity: real patterns and random models

Mid-domain models have been argued lo provide a default explanation for the best known spatial pattern in biodiversity, namely the latitudinal gradient in species richness. These models assume no environmental gradients, but merely a random latitudinal association between the size and placement of the geographic ranges of species. A mid-domain peak in richness is generated because when the latitudinal extents of species in a given taxonomic group are bounded to north and south, perhaps by a physical constraint such as a continental edge or perhaps by a climatic constraint such as a critical temperature or precipitation threshold, then the number of ways in which ranges can be distributed changes systematically between the bounds. In addition, such models make predictions about latitudinal variation in the latitudinal extents of the distributions of species, and in beta diversity (the spatial turnover in species identities). Here we test how well five mid-domain models predict observed latitudinal patterns of species richness, latitudinal extent and beta diversity in two groups of birds, parrots and woodpeckers, across the New World. Whilst both groups exhibit clear gradients in richness and beta diversity and the general trend in species richness is acceptably predicted (but not accurately, unless substantial empirical information is assumed), the fit of these models is uniformly poor for beta diversity and latitudinal range extent. This suggests either that, at least for these data, as presently formulated mid-domain models are too simplistic, or that in practice the mid-domain effect is not significant in determining geographical variation in diversity.     

Relevance of evolutionary history for food web structure

Explaining the structure of ecosystems is one of the great challenges of ecology. Simple models for food web structure aim at disentangling the complexity of ecological interaction networks and detect the main forces that are responsible for their shape. Trophic interactions are influenced by species traits, which in turn are largely determined by evolutionary history. Closely related species are more likely to share similar traits, such as body size, feeding mode and habitat preference than distant ones. Here, we present a theoretical framework for analysing whether evolutionary history--represented by taxonomic classification--provides valuable information on food web structure. In doing so, we measure which taxonomic ranks better explain species interactions. Our analysis is based on partitioning of the species into taxonomic units. For each partition, we compute the likelihood that a probabilistic model for food web structure reproduces the data using this information. We find that taxonomic partitions produce significantly higher likelihoods than expected at random. Marginal likelihoods (Bayes factors) are used to perform model selection among taxonomic ranks. We show that food webs are best explained by the coarser taxonomic ranks (kingdom to class). Our methods provide a way to explicitly include evolutionary history in models for food web structure.     

From diversity indices to community assembly processes: a test with simulated data

Ecological theory suggests that spatial distribution of biodiversity is strongly driven by community assembly processes. Thus the study of diversity patterns combined with null model testing has become increasingly common to infer assembly processes from observed distributions of diversity indices. However, results in both empirical and simulation studies are inconsistent. The aim of our study is to determine with simulated data which facets of biodiversity, if any, may unravel the processes driving its spatial patterns, and to provide practical considerations about the combination of diversity indices that would produce significant and congruent signals when using null models. The study is based on simulated species’ assemblages that emerge under various landscape structures in a spatially explicit individual‐based model with contrasting, predefined assembly processes. We focus on four assembly processes (species‐sorting, mass effect, neutral dynamics and competition colonization trade‐off) and investigate the emerging species’ distributions with varied diversity indices (alpha, beta and gamma) measured at different spatial scales and for different diversity facets (taxonomic, functional and phylogenetic). We find that 1) the four assembly processes result in distinct spatial distributions of species under any landscape structure, 2) a broad range of diversity indices allows distinguishing between communities driven by different assembly processes, 3) null models provide congruent results only for a small fraction of diversity indices and 4) only a combination of these diversity indices allows identifying the correct assembly processes. Our study supports the inference of assembly processes from patterns of diversity only when different types of indices are combined. It highlights the need to combine phylogenetic, functional and taxonomic diversity indices at multiple spatial scales to effectively infer underlying assembly processes from diversity patterns by illustrating how combination of different indices might help disentangling the complex question of coexistence.     

Clever birds are lousy: co-variation between avian innovation and the taxonomic richness of their amblyceran lice

Lice (Insecta: Phthiraptera) are ectoparasites that reduce host life expectancy and sexual attractiveness. Their taxonomic richness varies considerably among their hosts. Previous studies have already explored some important factors shaping louse diversity. An unexplored potential correlate of louse taxonomic richness is host behavioural flexibility. In this comparative study, we examine the relationship between louse generic richness, innovative capabilities (as a proxy for behavioural flexibility), and brain size while controlling for host species diversity, phylogeny, body size and research effort. Using data for 108 avian families, we found a highly significant positive relationship between host innovative capabilities and the taxonomic richness of amblyceran lice, but a lack of a similar relationship in ischnoceran lice. Host brain size had only a marginal impact on amblyceran diversity and no correlation with ischnoceran diversity. This suggests that the effect in Amblycera is not mediated by metabolic limitations due to the energetic costs of brain size and maintenance, rather directly caused by the ecological differences between hosts with differing cognitive capabilities. We propose four alternative and mutually non-exclusive hypotheses that may explain this phenomenon.     

Some like it hot: body and weapon size affect thermoregulation in horned beetles

Body size and shape affect thermoregulatory properties of organisms, and in turn are believed to have shaped macroevolutionary patterns of morphological diversity across many taxa. However, it is less clear whether thermoregulation plays a role in shaping intraspecific morphological diversity such as sexual dimorphisms or the conditional expression of exaggerated secondary sexual traits. Here, we investigate individual thermoregulatory properties in two species of horned beetles that share similar ecologies and body size ranges, but differ substantially in degree of sexual and male dimorphism. We find that intraspecific variation in body size had an unexpectedly large effect on thermal preference behavior and the ability to passively regulate body temperature. Furthermore, we find that the presence or absence of exaggerated secondary sexual traits dramatically altered thermal preference behavior, consistent with a thermoregulatory cost of horn possession. Lastly, we show that the increase in surface area associated with the expression of enlarged horns is, by itself, insufficient to account for the radically altered thermoregulatory behavior observed in horn-bearing males, and discuss possible alternative, physiological explanations. These findings are among the first to link intra-and interspecific variation in body- and weapon size to thermal preferences within and between insect species.     

Environmental correlates of body size distributions of European springtails (Hexapoda: Collembola)

Aim Species–body size distributions (SBDs) are plots of species richness across body size classes. They have been linked to energetic constraints, speciation–extinction dynamics and to evolutionary trends. However, little is known about the spatial variation of size distributions. Here we study SBDs of European springtails (Collembola) at a continental scale and test whether minimum, average and maximum body size and the shapes of size distributions change across latitudinal and longitudinal gradients and whether SBDs of islands and mainlands differ. We also test whether the island rule and the positive body size–range size relationship of vertebrates also holds for Collembola. Location Europe. Methods We use a unique data set on the spatial distributions of 2102 species of European springtails across 52 countries and larger islands together with associated data on body size, area, climate variables, longitude and latitude. Differences in the central moments of SBDs are inferred from simultaneous spatial autoregression models. Results The SBD of the European Collembola and its largest suborder Entomobryomorpha is unimodal and symmetrical. Average, minimum and maximum body weight and the skewness of the mainland/island SBDs peaked at intermediate latitudes. We could not find simple latitudinal gradients in minimum and maximum body weight. Average and maximum body size increased with country/island area in accordance with the island rule in vertebrates, while minimum body size did not significantly differ between islands and mainlands. Finally, we found a weak but statistically significant positive correlation of range size and body size. Main conclusions We provide evidence for differences in body size distributions between islands and mainlands that are in part in line with the island rule in invertebrates. We also find evidence for an interspecific body size–range size relationship similar to that of vertebrates although the vertebrate pattern is much stronger than the springtail pattern. Our results on latitudinal gradients of maximum and average body size imply the need to account for species richness and area effects in the study of latitudinal gradients in body size. We recommend implementing sample size and area effects in the study of body size distributions on islands and mainlands.