The evolution of armament strength: evidence for a constraint on the biting performance of claws of durophagous decapods

Performance data for the claws of six sympatric species of Cancer crabs confirmed a puzzling pattern reported previously for two other decapod crustaceans (stone crabs, Menippe mercenaria, and lobsters, Homarus americanus): Although biting forces increased, maximum muscle stresses (force per unit area) declined with increasing claw size. The negative allometry of muscle stress and the stress at a given claw size were fairly consistent within and among Cancer species despite significant differences in adult body size and relative claw size, but were not consistent among decapod genera. Therefore, claw height can be used as a reliable predictor of maximum biting force for the genus Cancer, but must be used with caution as a predictor of maximum biting force in wider evolutionary and biogeographical comparisons of decapods. The decline in maximum muscle stress with increasing claw size in Cancer crabs contrasts with the pattern in several other claw traits. Significantly, three traits that affect maximal biting force increased intraspecifically with increasing claw size: relative claw size, mechanical advantage, and sarcomere length of the closer muscle. Closer apodeme area and angle of pinnation of the closer muscle fibers varied isometrically with claw size. The concordant behavior of these traits suggests selection for higher biting forces in larger crabs. The contrast between the size dependence of muscle stress (negative allometry) and the remaining claw traits (isometry or positive allometry) strongly suggests that an as yet unidentified constraint impairs muscle performance in larger claws. The negative allometry of muscle stress in two distantly related taxa (stone crabs and lobsters) further suggests this constraint may be widespread in decapod crustaceans. The implications of this performance constraint for the evolution of claw size and the "arms-race" between decapod predators and their hard-shelled prey is discussed.     

A PHYLOGENETIC TEST OF ECOMORPHOLOGICAL ADAPTATION IN CANCER CRABS

We used McPeek's (1995a) method of evolutionary contrasts, and phylogenetic trees derived from maximum-parsimony, neighbor-joining, and maximum-likelihood analyses of data from the cytochrome oxidase I (COI) gene to evaluate the hypothesis that macroevolutionary changes in habitat use have driven the morphological diversification of Cancer crabs. All of our analyses suggested that habitat shifts from structurally complex substrates (e.g., the rocky intertidal zone) to more homogeneous substrates (e.g., sand or mud) have occurred independently in three Cancer lineages. Evolutionary contrasts analyses indicated that these habitat shifts were accompanied by increased morphological change toward larger body sizes. These macroevolutionary patterns support the hypothesis that the morphological diversification of Cancer crabs is strongly related to size-dependent habitat use; ancillary evidence suggests that increased predation pressure in homogeneous habitats represents the main selective agent for increased body size.     

Ecological specialization in fossil mammals explains Cope's rule

Cope's rule is the trend toward increasing body size in a lineage over geological time. The rule has been explained either as passive diffusion away from a small initial body size or as an active trend upheld by the ecological and evolutionary advantages that large body size confers. An explicit and phylogenetically informed analysis of body size evolution in Cenozoic mammals shows that body size increases significantly in most inclusive clades. This increase occurs through temporal substitution of incumbent species by larger-sized close relatives within the clades. These late-appearing species have smaller spatial and temporal ranges and are rarer than the incumbents they replace, traits that are typical of ecological specialists. Cope's rule, accordingly, appears to derive mainly from increasing ecological specialization and clade-level niche expansion rather than from active selection for larger size. However, overlain on a net trend toward average size increase, significant pulses in origination of large-sized species are concentrated in periods of global cooling. These pulses plausibly record direct selection for larger body size according to Bergmann's rule, which thus appears to be independent of but concomitant with Cope's.     

A comparative analysis of vigilance in birds

Individual variation in vigilance is known to vary with factors such as group size but the ecological determinants of vigilance among species have not been examined thus far in a systematic fashion. Earlier analyses suggested that vigilance should be lower in larger species and in species living in larger groups. These analyses were based on a small number of species and failed to take into account phylogenetic relationships among species. Here, I examined ecological determinants of vigilance in a large sample of bird species using a phylogenetic framework. I focused on vigilance in foraging groups of birds in the non-breeding season. Among species, vigilance by solitary foragers was not influenced by body mass. However, among species, asymptotic vigilance, the plateau reached by vigilance in larger groups, decreased with increasing group size in vegetarian clades but not in carnivorous clades. Asymptotic vigilance also increased with increasing body mass in vegetarian clades but not in carnivorous clades. Increasing group size may allow species to reduce vigilance in response to decreased predation risk. Increasing body mass may allow species to increase vigilance because more non-foraging time is available in larger species. Diet may modulate the effect of body mass and group size through factors such as within-group vigilance or foraging techniques.     

Rise of dinosaurs reveals major body-size transitions are driven by passive processes of trait evolution

A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252-201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian-Middle Jurassic. Maximum-likelihood analyses indicate that Cope's rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope's rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle-early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely-accepted 'rule' that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates.     

Evolutionary history and adaptive significance of respiratory structures on the legs of intertidal porcelain crabs, genus Petrolisthes

Semiterrestrial and terrestrial crabs have evolved multiple strategies for aerial respiration. An uncommon strategy for aerial respiration is seen in porcelain crabs, genus Petrolisthes, where decalcified areas on the meral segments of the walking legs are used as respiratory structures. Here, the evolutionary history and adaptive significance of these structures in porcelain crabs is examined. Interspecific variation in leg membrane size is from 0% to 60% of the surface area of the meral segment. Leg membrane relative size is positively correlated with body size across species but not within one species, Petrolisthes cinctipes. Phylogenetic analyses suggest that leg membranes are ancestral to one of two eastern Pacific Petrolisthes clades. Comparative analyses using phylogenetic independent contrasts indicate a relationship between leg membrane relative size and body size that is phylogenetically independent. In large-bodied intertidal species, whole-animal lactate accumulation during aerial incubation is 200%-300% higher when the leg membranes are obscured, indicating that the leg membranes are functional respiratory structures in these species. Thus, it is possible that leg membranes have facilitated the evolution of larger body sizes by providing additional respiratory surfaces to accommodate the associated higher metabolic demands.     

Tempo and mode of mandibular shape and size evolution reveal mixed support for incumbency effects in two clades of island‐endemic rodents (Muridae: Murinae)*

Existing radiations in a spatially limited system such as an oceanic island may limit the ecological opportunity experienced by later colonists, resulting in lower macroevolutionary rates for secondary radiations. Additionally, potential colonists may be competitively excluded by these incumbent (resident) species, unless they are biologically distinct (biotic filtering). The extant phenotypic diversity of secondary colonists may thus be impacted by lower rates of phenotypic evolution, exclusion from certain phenotypes, and transitions to new morphotypes to escape competition from incumbent lineages. We used geometric morphometric methods to test whether the rates and patterns of mandibular evolution of the Luzon “old endemic” rodent clades, Phloeomyini and Chrotomyini, are consistent with these predictions. Each clade occupied nearly completely separate shape space and partially separate size space. We detected limited support for decelerating and clade‐specific evolutionary rates for both shape and size, with strong evidence for a shift in evolutionary mode within Chrotomyini. Our results suggest that decelerating phenotypic evolutionary rates are not a necessary result of incumbency interactions; rather, incumbency effects may be more likely to determine which clades can become established in the system. Nonincumbent clades that pass a biotic filter can potentially exhibit relatively unfettered evolution.     

Scaling of species diversity and body mass in mammals: Cope’s rule and the evolutionary cost of large size

Abstract

Equations are constructed describing the inverse correlation of species diversity and body mass in extant and Cenozoic mammals. Cope’s rule, the tendency for many mammal clades to increase in body size through time, through phyletic change in single lineages or turnover within species groups, is interpreted as a probability function reducing diversity potential as a tradeoff for ecological/evolutionary gains. The inverse rule predicts that large species in clades will be less diverse than smaller species and, unless origination rates remain high among smaller clade members, clades conforming to Cope’s rule will decline in diversity, moving towards extinction. This proposition is evaluated in the Cenozoic histories of five North American mammal clades; cotton rats, felids, canids, hyaenodontids, and equids. Diversity potential of different size classes within the 3.75 million year phyletic history of the muskrat, Ondatra zibethicus, is also examined. A corollary prediction of the inverse rule, that large species should have longer durations (species lifespans) than small species, is unresolved. Successful clades maintain small size or a significant number of smaller species relative to clade average size. The potential loss of unique extant large mammal species justifies the conservation effort to protect them. The similarity of scaling exponents of species diversity to mass around a slope of -1.0 suggests that species diversity is correlated with home range size, the latter related to the probability of population fragmentation.     

Evolutionary diversification of clades of squamate reptiles

We analysed the diversification of squamate reptiles (7488 species) based on a new molecular phylogeny, and compared the results to similar estimates for passerine birds (5712 species). The number of species in each of 36 squamate lineages showed no evidence of phylogenetic conservatism. Compared with a random speciation-extinction process with parameters estimated from the size distribution of clades, the alethinophidian snakes (2600 species) were larger than expected and 13 clades, each having fewer than 20 species, were smaller than expected, indicating rate heterogeneity. From a lineage-through-time plot, we estimated that a provisional rate of lineage extinction (0.66 per Myr) was 94% of the rate of lineage splitting (0.70 per Myr). Diversification in squamate lineages was independent of their stem age, but strongly related to the area of the region within which they occur. Tropical vs. temperate latitude exerted a marginally significant influence on species richness. In comparison with passerine birds, squamates share several clade features, including: (1) independence of species richness and age; (2) lack of phylogenetic signal with respect to clade size; (3) general absence of exceptionally large clades; (4) over-representation of small clades; (5) influence of region size on clade size; and (6) similar rates of speciation and extinction. The evidence for both groups suggests that clade size has achieved long-term equilibrium, suggesting negative feedback of species richness on the rate of diversification.     

Cope's Rule in a modular organism: Directional evolution without an overarching macroevolutionary trend

Cope's Rule describes increasing body size in evolutionary lineages through geological time. This pattern has been documented in unitary organisms but does it also apply to module size in colonial organisms? We address this question using 1169 cheilostome bryozoans ranging through the entire 150 million years of their evolutionary history. The temporal pattern evident in cheilostomes as a whole shows no overall change in zooid (module) size. However, individual subclades show size increases: within a genus, younger species often have larger zooids than older species. Analyses of (paleo)latitudinal shifts show that this pattern cannot be explained by latitudinal effects (Bergmann's Rule) coupled with younger species occupying higher latitudes than older species (an “out of the tropics” hypothesis). While it is plausible that size increase was linked to the advantages of large zooids in feeding, competition for trophic resources and living space, other proposed mechanisms for Cope's Rule in unitary organisms are either inapplicable to cheilostome zooid size or cannot be evaluated. Patterns and mechanisms in colonial organisms cannot and should not be extrapolated from the better‐studied unitary organisms. And even if macroevolution simply comprises repeated rounds of microevolution, evolutionary processes occurring within lineages are not always detectable from macroevolutionary patterns.     

Patterns of sexual size dimorphism in Chelonia

Macroevolutionary patterns of sexual size dimorphism (SSD) indicate how sexual selection, natural selection, and genetic and developmental constraints mold sex differences in body size. One putative pattern, known as Rensch's rule, posits that, among species with female‐larger SSD, the relative degree of SSD declines with species' body size, whereas, among male‐larger SSD species, relative SSD increases with size. Using a dataset of 196 chelonian species from all fourteen families, we investigated the correlation in body size evolution between male and female Chelonia and the validity of Rensch's rule for the taxon and within its major clades. We conclude that male–female correlations in body size evolution are high, although these correlations differ among chelonian families. Overall, SSD scales isometrically with body size; Rensch's rule is valid for only one family, Testudinidae (tortoises). Because macroevolutionary patterns of SSD can vary markedly among clades, even in a taxon as morphologically conservative as Testudines, one must guard against inappropriately pooling clades in comparative studies of SSD. The results of the present study also indicate that regression models that assume the x‐variable (e.g. male body size) is measured without statistical error, although frequently reported, will result in erroneous conclusions about phylogenetic trends in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 396–413.     

Repeated evolution of vertebrate pollination syndromes in a recently diverged Andean plant clade

Although specialized interactions, including those involving plants and their pollinators, are often invoked to explain high species diversity, they are rarely explored at macroevolutionary scales. We investigate the dynamic evolution of hummingbird and bat pollination syndromes in the centropogonid clade (Lobelioideae: Campanulaceae), an Andean‐centered group of ∼550 angiosperm species. We demonstrate that flowers hypothesized to be adapted to different pollinators based on flower color fall into distinct regions of morphospace, and this is validated by morphology of species with known pollinators. This supports the existence of pollination syndromes in the centropogonids, an idea corroborated by ecological studies. We further demonstrate that hummingbird pollination is ancestral, and that bat pollination has evolved ∼13 times independently, with ∼11 reversals. This convergence is associated with correlated evolution of floral traits within selective regimes corresponding to pollination syndrome. Collectively, our results suggest that floral morphological diversity is extremely labile, likely resulting from selection imposed by pollinators. Finally, even though this clade's rapid diversification is partially attributed to their association with vertebrate pollinators, we detect no difference in diversification rates between hummingbird‐ and bat‐pollinated lineages. Our study demonstrates the utility of pollination syndromes as a proxy for ecological relationships in macroevolutionary studies of certain species‐rich clades.     

Climate‐driven body‐size trends in the ostracod fauna of the deep Indian Ocean

Abstract: Body size is a common focus of macroevolutionary, macroecological and palaeontological investigations. Here, we document body‐size evolution in 19 species‐level ostracod lineages from the deep Indian Ocean (Deep Sea Drilling Program Site 253) over the past 40 myr. Body‐size trajectories vary across taxa and time intervals, but most lineages (16/19) show net gains in body size. Because many modern ostracod taxa are larger in colder parts of their geographical range, we compared the timing and magnitude of these size changes to established Cenozoic deep‐water cooling patterns confirmed through δ¹⁸O measurements of benthic foraminifera in the samples studied. These data show a significant negative correlation between size changes and temperature changes (ostracods get larger as temperatures get colder), and that systematic size increases only occur during intervals of sustained cooling. In addition, statistical support for an explicit temperature‐tracking model exceeds that of purely directional evolution. We argue that this Cope’s Rule pattern is driven by secular changes in the environment, rather than any universal or intrinsic advantages to larger body sizes, and we note some difficulties in the attempts to link Cope’s Rule to observations made within a single generation.     

Rapid lineage accumulation in a non-adaptive radiation: phylogenetic analysis of diversification rates in eastern North American woodland salamanders (Plethodontidae: Plethodon)

Adaptive radiations have served as model systems for quantifying the build-up of species richness. Few studies have quantified the tempo of diversification in species-rich clades that contain negligible adaptive disparity, making the macroevolutionary consequences of different modes of evolutionary radiation difficult to assess. We use mitochondrial-DNA sequence data and recently developed phylogenetic methodologies to explore the tempo of diversification of eastern North American Plethodon, a species-rich clade of woodland salamanders exhibiting only limited phenotypic disparity. Lineage-through-time analysis reveals a high rate of lineage accumulation, 0.8 species per million years, occurring 11-8 million years ago in the P. glutinosus species group, followed by decreasing rates. This high rate of lineage accumulation is exceptional, comparable to the most rapid of adaptive radiations. In contrast to classic models of adaptive radiation where ecological niche divergence is linked to the origin of species, we propose that phylogenetic niche conservatism contributes to the rapid accumulation of P. glutinosus-group lineages by promoting vicariant isolation and multiplication of species across a spatially and temporally fluctuating environment. These closely related and ecologically similar lineages persist through long-periods of evolutionary time and form strong barriers to the geographic spread of their neighbours, producing a subsequent decline in lineage accumulation. Rapid diversification among lineages exhibiting long-term maintenance of their bioclimatic niche requirements is an under-appreciated phenomenon driving the build-up of species richness.     

Phylogenetic perspectives on reef fish functional traits

Functional traits have been fundamental to the evolution and diversification of entire fish lineages on coral reefs. Yet their relationship with the processes promoting speciation, extinction and the filtering of local species pools remains unclear. We review the current literature exploring the evolution of diet, body size, water column use and geographic range size in reef‐associated fishes. Using published and new data, we mapped functional traits on to published phylogenetic trees to uncover evolutionary patterns that have led to the current functional diversity of fishes on coral reefs. When examining reconstructed patterns for diet and feeding mode, we found examples of independent transitions to planktivory across different reef fish families. Such transitions and associated morphological alterations may represent cases in which ecological opportunity for the exploitation of different resources drives speciation and adaptation. In terms of body size, reconstructions showed that both large and small sizes appear multiple times within clades of mid‐sized fishes and that extreme body sizes have arisen mostly in the last 10 million years (Myr). The reconstruction of range size revealed many cases of disparate range sizes among sister species. Such range size disparity highlights potential vicariant processes through isolation in peripheral locations. When accounting for peripheral speciation processes in sister pairs, we found a significant relationship between labrid range size and lineage age. The diversity and evolution of traits within lineages is influenced by trait–environment interactions as well as by species and trait–trait interactions, where the presence of a given trait may trigger the development of related traits or behaviours. Our effort to assess the evolution of functional diversity across reef fish clades adds to the burgeoning research focusing on the evolutionary and ecological roles of functional traits. We argue that the combination of a phylogenetic and a functional approach will improve the understanding of the mechanisms of species assembly in extraordinarily rich coral reef communities.     

Do key innovations unlock diversification? A case-study on the morphological and ecological impact of pharyngognathy in acanthomorph fishes

Key innovations may allow lineages access to new resources and facilitate the invasion of new adaptive zones, potentially influencing diversification patterns. Many studies have focused on the impact of key innovations on speciation rates, but far less is known about how they influence phenotypic rates and patterns of ecomorphological diversification. We use the repeated evolution of pharyngognathy within acanthomorph fishes, a commonly cited key innovation, as a case study to explore the predictions of key innovation theory. Specifically, we investigate whether transitions to pharyngognathy led to shifts in the rate of phenotypic evolution, as well as shifts and/or expansion in the occupation of morphological and dietary space, using a dataset of 8 morphological traits measured across 3,853 species of Acanthomorpha. Analyzing the 6 evolutionarily independent pharyngognathous clades together, we found no evidence to support pharyngognathy as a key innovation; however, comparisons between individual pharyngognathous lineages and their sister clades did reveal some consistent patterns. In morphospace, most pharyngognathous clades cluster in areas that correspond to deeper-bodied morphologies relative to their sister clades, while occupying greater areas in dietary space that reflects a more diversified diet. Additionally, both Cichlidae and Labridae exhibited higher univariate rates of phenotypic evolution compared with their closest relatives. However, few of these results were exceptional relative to our null models. Our results suggest that transitions to pharyngognathy may only be advantageous when combined with additional ecological or intrinsic factors, illustrating the importance of accounting for lineage-specific effects when testing key innovation hypotheses. Moreover, the challenges we experienced formulating informative comparisons, despite the ideal evolutionary scenario of multiple independent evolutionary origins of pharyngognathous clades, illustrates the complexities involved in quantifying the impact of key innovations. Given the issues of lineage specific effects and rate heterogeneity at macroevolutionary scales we observed, we suggest a reassessment of the expected impacts of key innovations may be warranted.     

Idiosyncratic species effects confound size-based predictions of responses to climate change

Understanding and predicting the consequences of warming for complex ecosystems and indeed individual species remains a major ecological challenge. Here, we investigated the effect of increased seawater temperatures on the metabolic and consumption rates of five distinct marine species. The experimental species reflected different trophic positions within a typical benthic East Atlantic food web, and included a herbivorous gastropod, a scavenging decapod, a predatory echinoderm, a decapod and a benthic-feeding fish. We examined the metabolism–body mass and consumption–body mass scaling for each species, and assessed changes in their consumption efficiencies. Our results indicate that body mass and temperature effects on metabolism were inconsistent across species and that some species were unable to meet metabolic demand at higher temperatures, thus highlighting the vulnerability of individual species to warming. While body size explains a large proportion of the variation in species'' physiological responses to warming, it is clear that idiosyncratic species responses, irrespective of body size, complicate predictions of population and ecosystem level response to future scenarios of climate change.     

Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors

The Anomura presents the greatest degree of morphological disparity in the decapod Crustacea, with body forms ranging from the symmetrical and asymmetrical hermit crabs to squat lobsters and king crabs. The phylogeny of the anomurans has been fraught with controversy. Recent debate has focused primarily on the phenomenon of carcinization, the evolution of crab-like form from a non-crab-like ancestor, focused chiefly on derivation of king crabs from asymmetrical hermit crabs--the "hermit to king" hypothesis. We show by phylogenetic analysis of five nuclear protein-coding gene sequences that hermit crabs have a single origin, but surprisingly, that almost all other major clades and body forms within the Anomura, are derived from within the hermit crabs. The crab-like form and squat lobster form have each evolved at least twice from separate symmetrical hermit crab ancestors. In each case, a carcinization trend can be posited via a transition series from the initial symmetrical long-tailed hermit crab form, through the intermediate squat lobster or asymmetrical hermit crab form, to the final crab-like form. Adaptation to dextral shell habitation evolved at least twice, once in an exclusively deep-water clade and once in the common ancestor of all other asymmetrical hermit crabs (from which king crabs are derived). These remarkable cases of parallelism suggest considerable phenotypic flexibility within the hermit crab ground plan, with a general tendency toward carcinization. Rather than having a separate origin from other major clades, hermit crabs have given rise to most other major anomuran body types.     

Adaptive processes drive ecomorphological convergent evolution in antwrens (Thamnophilidae)

Phylogenetic niche conservatism (PNC) and convergence are contrasting evolutionary patterns that describe phenotypic similarity across independent lineages. Assessing whether and how adaptive processes give origin to these patterns represent a fundamental step toward understanding phenotypic evolution. Phylogenetic model‐based approaches offer the opportunity not only to distinguish between PNC and convergence, but also to determine the extent that adaptive processes explain phenotypic similarity. The Myrmotherula complex in the Neotropical family Thamnophilidae is a polyphyletic group of sexually dimorphic small insectivorous forest birds that are relatively homogeneous in size and shape. Here, we integrate a comprehensive species‐level molecular phylogeny of the Myrmotherula complex with morphometric and ecological data within a comparative framework to test whether phenotypic similarity is described by a pattern of PNC or convergence, and to identify evolutionary mechanisms underlying body size and shape evolution. We show that antwrens in the Myrmotherula complex represent distantly related clades that exhibit adaptive convergent evolution in body size and divergent evolution in body shape. Phenotypic similarity in the group is primarily driven by their tendency to converge toward smaller body sizes. Differences in body size and shape across lineages are associated to ecological and behavioral factors.     

Diversity and Evolution of Body Size in Fishes

The diversity of body sizes observed among species of a clade is a combined result of microevolutionary processes (i.e. natural selection and genetic drift) that cause size changes within phylogenetic lineages, and macroevolutionary processes (i.e. speciation and extinction) that affect net rates of diversification among lineages. Here we assess trends of size diversity and evolution in fishes (non-tetrapod craniates), employing paleontological, macroecological, and phylogenetic information. Fishes are well suited to studies of size diversity and evolution, as they are highly diverse, representing more than 50% of all living vertebrate species, and many fish taxa are well represented in the fossil record from throughout the Phanerozoic. Further, the frequency distributions of sizes among fish lineages resemble those of most other animal taxa, in being right-skewed, even on a log scale. Using an approach that measures rates of size evolution (in darwins) within a formal phylogenetic framework, we interpret the shape of size distributions as a balance between the competing forces of diversification, pushing taxa away from ancestral values, and of conservation, drawing taxa closer to a central tendency. Within this context we show how non-directional mechanisms of evolution (i.e. passive diffusion processes) can produce an hitherto unperceived bias to larger size, when size is measured on the conventional log scale. These results demonstrate how the interpretation of macroecological datasets can be enriched from an historical perspective, and document the ways in which macroevolutionary and microevolutionary processes may be decoupled in the production of size diversity.