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Morphological traits often covary within and among species according to simple power laws referred to as allometry. Such allometric relationships may result from common growth regulation, and this has given rise to the hypothesis that allometric exponents may have low evolvability and constrain trait evolution. We formalize hypotheses for how allometry may constrain morphological trait evolution across taxa, and test these using more than 300 empirical estimates of static (within-species) allometric relations of animal morphological traits. Although we find evidence for evolutionary changes in allometric parameters on million-year, cross-species time scales, there is limited evidence for microevolutionary changes in allometric slopes. Accordingly, we find that static allometries often predict evolutionary allometries on the subspecies level, but less so across species. Although there is a large body of work on allometry in a broad sense that includes all kinds of morphological trait–size relationships, we found relatively little information about the evolution of allometry in the narrow sense of a power relationship. Despite the many claims of microevolutionary changes of static allometries in the literature, hardly any of these apply to narrow-sense allometry, and we argue that the hypothesis of strongly constrained static allometric slopes remains viable.

Forest tree communities are largely structured by interactions between phenotypes and their environments. Functional traits have been popularized as providing key insights into plant functional tradeoffs. Similarly, tree crown—stem diameter and tree height—stem diameter allometric relationships are likely to be strongly coordinated with functional trait tradeoff axes. Specifically, species with functional traits indicative of conservative strategies (i.e., dense wood, heavy seeds) should be related to tree architectures that have lower heights and wider crowns for a given stem diameter. For example, shade‐tolerant species in tropical forests are typically characterized as having dense wood, large seeds, and relatively broad crowns at early ontogenetic stages. Here, we focus on 14 dominant dicot tree species in a tropical forest. We utilized hierarchical Bayesian models to characterize species‐specific height and crown size allometric parameters. We sampled from the posterior distributions for these parameters and correlated them with six functional traits. We also characterize the expected height and crown size for a series of reference stem diameters to quantify the relationship between traits and tree architecture across size classes. We find little interspecific variation in allometric slopes, but clear variation in allometric intercepts. Allometeric height intercepts were negatively correlated with wood density and crown size intercepts were positively related to wood density and seed mass and negatively related to leaf percent phosphorus. Thus, interspecific variation in tree architecture is generated by interspecific variation in allometric intercepts and not slopes. These intercepts could be predicted using a handful of functional traits where conservative traits were indicative of trees that are relatively short and have larger crown sizes. This demonstrates a coordination of tropical tree life histories that can be characterized simultaneously with functional traits and tree allometries. The ontogenetic trajectory of height and crown allocation of tropical trees is an essential indicator of their life histories. Here, we show that this whole plant strategy can be linked to organ‐level plant functional traits leading to insights into how phenotypes are integrated.

Sexual selection has driven the evolution of exaggerated traits among diverse animal taxa. The production of exaggerated traits can come at a cost to other traits through trade-offs when resources allocated to trait development are limited. Alternatively some traits can be selected for in parallel to support or compensate for the cost of bearing the exaggerated trait. Male giraffe weevils (Lasiorhynchus barbicornis) display an extremely elongated rostrum used as a weapon during contests for mates. Here we characterise the scaling relationship between rostrum and body size and show that males have a steep positive allometry, but that the slope is non-linear due to a relative reduction in rostrum length for the largest males, suggesting a limitation in resource allocation or a diminishing requirement for large males to invest increasingly into larger rostra. We also measured testes, wings, antennae, fore- and hind-tibia size and found no evidence of a trade-off between these traits and rostrum length when comparing phenotypic correlations. However, the relative length of wings, antennae, fore- and hind-tibia all increased with relative rostrum length suggesting these traits may be under correlational selection. Increased investment in wing and leg length is therefore likely to compensate for the costs of flying with, and wielding the exaggerated rostrum of larger male giraffe weevils. These results provide a first step in identifying the potential for trait compensation and trades-offs, but are phenotypic correlations only and should be interpreted with care in the absence of breeding experiments.

Allometry is vital for understanding the mechanisms underlying phenotypic evolution. Despite a large body of literature on allometry, studies based on fossil time series are limited for solitary organisms and nonexistent for colonial organisms. Allometric relationships have been found to be relatively constant across Recent populations of the same species, separated by space, but variable among fossil populations separated by thousands of years. How stable are allometric relationships at the module level for colonial organisms? We address this question using two extant species of the cheilostome bryozoan Microporella with fossil records spanning the Pleistocene of New Zealand. We investigate size covariation between feeding modules and three traits with separate functions (reproductive, resource uptake, and defense). We found that within-population (static) allometry can change on timescales of at least 0.1 million years. These within-population relationships do not consistently predict overintraspecific evolutionary allometry, which in turn does not predict those estimated at the genus level. Different functional traits are constrained to different extents by module size with defensive traits being the least constrained and most evolvable, compared with reproductive and resource uptake traits. Our study highlights the potential of colonial organisms in understanding the constraints and drivers of long-term phenotypic change.

[This corrects the article DOI: 10.1371/journal.pone.0151482.].

1. Promoting mixed-species forests is an important strategy for adaptation and risk reduction in the face of global change. Concurrently, a main challenge in ecology is to quantify the effects of species diversity on ecosystem functioning. In forests, the effects of individual tree species on ecosystem functions depend largely on their dimensions, which are commonly predicted using allometric equations. However, little is known about how diversity influences allometry or how to incorporate this effect into allometric equations. Ignoring the effects of interspecific interactions on allometric relationships may result in severely biased predictions. 2. This study examined the effects of tree-species diversity, competition and tree social status on crown-projection area (cpa), height (h) and live-crown length (lcl) of trees using a European-wide data set containing 17 target species and 12 939 trees. The cpa, h and lcl were predicted as functions of stem diameter at 1·3 m, tree-species diversity, tree height relative to the stand mean height (rh) and a competition index (CI) that accounted for stand density and interspecific differences in competitive ability based on species-specific wood density or shade tolerance. 3. Averaged across species, diameter had the greatest effect on cpa and lcl, followed by the competition index, while rh had the greatest effect on lcl. Tree-species diversity had the smallest effect on cpa, h and lcl. Interspecific variability in cpa, h or lcl responses to diversity, CI, or rh was sometimes related to wood density or shade tolerance. 4. Synthesis. This study shows the strong influence of stand structure and species composition on allometric relationships. These influences can be quantified using measures of competition, tree-species diversity and relative tree height so that general equations can be developed for a given species to be applied to a wide range of species compositions and stand structures. This new approach will greatly improve predictions of biomass and carbon stocks in structurally and compositionally diverse forests.

Ontogenetic allometry, how species change with size through their lives, and heterochony, a decoupling between shape, size, and age, are major contributors to biological diversity. However, macroevolutionary allometric and heterochronic trends remain poorly understood because previous studies have focused on small groups of closely related species. Here, we focus on testing hypotheses about the evolution of allometry and how allometry and heterochrony drive morphological diversification at the level of an entire species-rich and diverse clade. Pythons are a useful system due to their remarkably diverse and well-adapted phenotypes and extreme size disparity. We collected detailed phenotype data on 40 of the 44 species of python from 1191 specimens. We used a suite of analyses to test for shifts in allometric trajectories that modify morphological diversity. Heterochrony is the main driver of initial divergence within python clades, and shifts in the slopes of allometric trajectories make exploration of novel phenotypes possible later in divergence history. We found that allometric coefficients are highly evolvable and there is an association between ontogenetic allometry and ecology, suggesting that allometry is both labile and adaptive rather than a constraint on possible phenotypes.

Accounting for phylogenetic relatedness in the analysis of shape is generally deemed necessary to acknowledge the non-independence between species due to common ancestry. However, some response variation can be both attributable to a predictor variable and phylogenetically structured, an effect termed conservative trait correlation ( CTC ). For CTC s, the effects of predictors versus phylogenetic structure cannot easily be disentangled. This is problematic because standard phylogenetic tests wholly attribute CTC to phylogenetic structuring when adjusting error distributions for relatedness. By contrast, non-phylogenetic methods retain CTC as wholly predictor-related, neglecting to estimate the effects of phylogenetic structuring altogether. Characterising how CTC contributes to a shape response is therefore crucial to support the interpretation of shape analyses. We here demonstrate a strong CTC effect in the geometric morphometric shape variation among 370 crania from the speciose Australian rock-wallabies (genus Petrogale ). Specifically, significant allometry supporting mechanistically well-explained allometric patterns such as scaling of the braincase (Haller’s rule) and snout length (craniofacial evolutionary allometry) is reported by ordinary least squares ( OLS ) regression, but not phylogenetic generalised least squares ( PGLS ) analysis. We then apply two methods capable of characterising the missing variation: variation partitioning ( VARPART ), which estimates the proportion of variation shared between the predictor and phylogeny, and multi-response phylogenetic mixed models ( MR-PMM ), which identify the strength of correlation within the phylogenetic component of trait variance. Both methods show that CTC dominates the allometric shape variation in our sample. Our results therefore highlight CTC estimation as an important additional tool to clarify the relationship between morphological evolution and its predictors.

Several investigations in recent years have reported patterns of discontinuous, biphasic, loglinear variation in the metabolic allometry of aquatic animals. These putative shifts in pattern of allometry have been attributed to changes in the primary site for gas exchange from cutaneous to branchial as animals undergo ontogenetic changes in size, shape, and surface area. Because of the important implications of the earlier research with regard to both physiology and evolution, I re-examined data that purportedly support claims of discontinuous, biphasic allometry in oxygen consumption versus body size of American eels (Anguilla rostrata) and spiny lobsters (Sagmariasus verreauxi). I used ANCOVA to fit three different statistical models to each set of logarithmic transformations and then assessed the fits by Akaike's Information Criterion. The observations for both species were described better by a single straight line fitted to the full distribution than by a biphasic model. Eels, lobsters, and other aquatic animals undergo changes in shape and surface area as they grow, but such changes are not necessarily accompanied by changes in the pattern of metabolic allometry.

Complex allometry describes a smooth, curvilinear relationship between logarithmic transformations of a biological variable and a corresponding measure for body size when the observations are displayed on a bivariate graph with linear scaling. The curvature in such a display is commonly captured by fitting a quadratic equation to the distribution; and the quadratic term is typically interpreted, in turn, to mean that the mathematically equivalent equation for describing the arithmetic distribution is a two-parameter power equation with an exponent that changes with body size. A power equation with an exponent that is itself a function of body size is virtually uninterpretable, yet numerous attempts have been made in recent years to incorporate such an exponent into theoretical models for the evolution of form and function in both plants and animals. However, the curvature that is described by a quadratic equation fitted to logarithms usually means that an explicit, non-zero intercept is required in the power equation describing the untransformed distribution — not that the exponent in the power equation varies with body size. Misperceptions that commonly accompany reports of complex allometry can be avoided by using nonlinear regression to examine untransformed data.