Evolutionary Increase In Body Size Research Articles

Biological scaling analyses are more than statistical line fitting.

The magnitude of many biological traits relates strongly and regularly to body size. Consequently, a major goal of comparative biology is to understand and apply these 'size-scaling' relationships, traditionally quantified by using linear regression analyses based on log-transformed data. However, recently some investigators have questioned this traditional method, arguing that linear or non-linear regression based on untransformed arithmetic data may provide better statistical fits than log-linear analyses. Furthermore, they advocate the replacement of the traditional method by alternative specific methods on a case-by-case basis, based simply on best-fit criteria. Here, I argue that the use of logarithms in scaling analyses presents multiple valuable advantages, both statistical and conceptual. Most importantly, log-transformation allows biologically meaningful, properly scaled (scale-independent) comparisons of organisms of different size, whereas non-scaled (scale-dependent) analyses based on untransformed arithmetic data do not. Additionally, log-based analyses can readily reveal biologically and theoretically relevant discontinuities in scale invariance during developmental or evolutionary increases in body size that are not shown by linear or non-linear arithmetic analyses. In this way, log-transformation advances our understanding of biological scaling conceptually, not just statistically. I hope that my Commentary helps students, non-specialists and other interested readers to understand the general benefits of using log-transformed data in size-scaling analyses, and stimulates advocates of arithmetic analyses to show how they may improve our understanding of scaling conceptually, not just statistically.

Morphological divergence in giant fossil dormice.

Insular gigantism-evolutionary increases in body size from small-bodied mainland ancestors-is a conceptually significant, but poorly studied, evolutionary phenomenon. Gigantism is widespread on Mediterranean islands, particularly among fossil and extant dormice. These include an extant giant population of Eliomys quercinus on Formentera, the giant Balearic genus †Hypnomys and the exceptionally large †Leithia melitensis of Pleistocene Sicily. We quantified patterns of cranial and mandibular shape and their relationships to head size (allometry) among mainland and insular dormouse populations, asking to what extent the morphology of island giants is explained by allometry. We find that gigantism in dormice is not simply an extrapolation of the allometric trajectory of their mainland relatives. Instead, a large portion of their distinctive cranial and mandibular morphology resulted from the population- or species-specific evolutionary shape changes. Our findings suggest that body size increases in insular giant dormice were accompanied by the evolutionary divergence of feeding adaptations. This complements other evidence of ecological divergence in these taxa, which span predominantly faunivorous to herbivorous diets. Our findings suggest that insular gigantism involves context-dependent phenotypic modifications, underscoring the highly distinctive nature of island faunas.

Functional diversity of small-mammal postcrania is linked to both substrate preference and body size.

Selective pressures favor morphologies that are adapted to distinct ecologies, resulting in trait partitioning among ecomorphotypes. However, the effects of these selective pressures vary across taxa, especially because morphology is also influenced by factors such as phylogeny, body size, and functional trade-offs. In this study, we examine how these factors impact functional diversification in mammals. It has been proposed that trait partitioning among mammalian ecomorphotypes is less pronounced at small body sizes due to biomechanical, energetic, and environmental factors that favor a “generalist” body plan, whereas larger taxa exhibit more substantial functional adaptations. We title this the Divergence Hypothesis (DH) because it predicts greater morphological divergence among ecomorphotypes at larger body sizes. We test DH by using phylogenetic comparative methods to examine the postcranial skeletons of 129 species of taxonomically diverse, small-to-medium-sized (<15 kg) mammals, which we categorize as either “tree-dwellers” or “ground-dwellers.” In some analyses, the morphologies of ground-dwellers and tree-dwellers suggest greater between-group differentiation at larger sizes, providing some evidence for DH. However, this trend is neither particularly strong nor supported by all analyses. Instead, a more pronounced pattern emerges that is distinct from the predictions of DH: within-group phenotypic disparity increases with body size in both ground-dwellers and tree-dwellers, driven by morphological outliers among “medium”-sized mammals. Thus, evolutionary increases in body size are more closely linked to increases in within-locomotor-group disparity than to increases in between-group disparity. We discuss biomechanical and ecological factors that may drive these evolutionary patterns, and we emphasize the significant evolutionary influences of ecology and body size on phenotypic diversity.

Life's Energy and Information: Contrasting Evolution of Volume- versus Surface-Specific Rates of Energy Consumption.

As humanity struggles to find a path to resilience amidst global change vagaries, understanding organizing principles of living systems as the pillar for human existence is rapidly growing in importance. However, finding quantitative definitions for order, complexity, information and functionality of living systems remains a challenge. Here, we review and develop insights into this problem from the concept of the biotic regulation of the environment developed by Victor Gorshkov (1935–2019). Life’s extraordinary persistence—despite being a strongly non-equilibrium process—requires a quantum-classical duality: the program of life is written in molecules and thus can be copied without information loss, while life’s interaction with its non-equilibrium environment is performed by macroscopic classical objects (living individuals) that age. Life’s key energetic parameter, the volume-specific rate of energy consumption, is maintained within universal limits by most life forms. Contrary to previous suggestions, it cannot serve as a proxy for “evolutionary progress”. In contrast, ecosystem-level surface-specific energy consumption declines with growing animal body size in stable ecosystems. High consumption by big animals is associated with instability. We suggest that the evolutionary increase in body size may represent a spontaneous loss of information about environmental regulation, a manifestation of life’s algorithm ageing as a whole.

New insights on equid locomotor evolution from the lumbar region of fossil horses.

The specialization of equid limbs for cursoriality is a classic case of adaptive evolution, but the role of the axial skeleton in this famous transition is not well understood. Extant horses are extremely fast and efficient runners, which use a stiff-backed gallop with reduced bending of the lumbar region relative to other mammals. This study tests the hypothesis that stiff-backed running in horses evolved in response to evolutionary increases in body size by examining lumbar joint shape from a broad sample of fossil equids in a phylogenetic context. Lumbar joint shape scaling suggests that stability of the lumbar region does correlate with size through equid evolution. However, scaling effects were dampened in the posterior lumbar region, near the sacrum, which suggests strong selection for sagittal mobility in association with locomotor-respiratory coupling near the lumbosacral joint. I hypothesize that small-bodied fossil horses may have used a speed-dependent running gait, switching between stiff-backed and flex-backed galloping as speed increased.

Axial allometry in a neutrally buoyant environment: effects of the terrestrial-aquatic transition on vertebral scaling.

Ecological diversification into new environments presents new mechanical challenges for locomotion. An extreme example of this is the transition from a terrestrial to an aquatic lifestyle. Here, we examine the implications of life in a neutrally buoyant environment on adaptations of the axial skeleton to evolutionary increases in body size. On land, mammals must use their thoracolumbar vertebral column for body support against gravity and thus exhibit increasing stabilization of the trunk as body size increases. Conversely, in water, the role of the axial skeleton in body support is reduced, and, in aquatic mammals, the vertebral column functions primarily in locomotion. Therefore, we hypothesize that the allometric stabilization associated with increasing body size in terrestrial mammals will be minimized in secondarily aquatic mammals. We test this by comparing the scaling exponent (slope) of vertebral measures from 57 terrestrial species (23 felids, 34 bovids) to 23 semi-aquatic species (pinnipeds), using phylogenetically corrected regressions. Terrestrial taxa meet predictions of allometric stabilization, with posterior vertebral column (lumbar region) shortening, increased vertebral height compared to width, and shorter, more disc-shaped centra. In contrast, pinniped vertebral proportions (e.g. length, width, height) scale with isometry, and in some cases, centra even become more spool-shaped with increasing size, suggesting increased flexibility. Our results demonstrate that evolution of a secondarily aquatic lifestyle has modified the mechanical constraints associated with evolutionary increases in body size, relative to terrestrial taxa.

Body size evolution in Mesozoic birds: little evidence for Cope’s rule

Cope's rule, the tendency towards evolutionary increases in body size, is a long-standing macroevolutionary generalization that has the potential to provide insights into directionality in evolution; however, both the definition and identification of Cope's rule are controversial and problematic. A recent study [J. Evol. Biol. 21 (2008) 618] examined body size evolution in Mesozoic birds, and claimed to have identified evidence of Cope's rule occurring as a result of among-lineage species sorting. We here reassess the results of this study, and additionally carry out novel analyses testing for within-lineage patterns in body size evolution in Mesozoic birds. We demonstrate that the nonphylogenetic methods used by this previous study cannot distinguish between among- and within-lineage processes, and that statistical support for their results and conclusions is extremely weak. Our ancestor-descendant within-lineage analyses explicitly incorporate recent phylogenetic hypotheses and find little compelling evidence for Cope's rule. Cope's rule is not supported in Mesozoic birds by the available data, and body size evolution currently provides no insights into avian survivorship through the Cretaceous-Paleogene mass extinction.

The evolution of body size in extant groups of North American freshwater fishes: speciation, size distributions, and Cope's rule.

Change in body size within an evolutionary lineage over time has been under investigation since the synthesis of Cope's rule, which suggested that there is a tendency for mammals to evolve larger body size. Data from the fossil record have subsequently been examined for several other taxonomic groups to determine whether they also displayed an evolutionary increase in body size. However, we are not aware of any species-level study that has investigated the evolution of body size within an extant continental group. Data acquired from the fossil record and data derived from the evolutionary relationships of extant species are not similar, with each set exhibiting both strengths and weaknesses related to inferring evolutionary patterns. Consequently, expectation that general trends exhibited in the fossil record will correspond to patterns in extant groups is not necessarily warranted. Using phylogenetic relationships of extant species, we show that five of nine families of North American freshwater fishes exhibit an evolutionary trend of decreasing body size. These trends result from the basal position of large species and the more derived position of small species within families. Such trends may be caused by the invasion of small streams and subsequent isolation and speciation. This pattern, potentially influenced by size-biased dispersal rates and the high percentage of small streams in North America, suggests a scenario that could result in the generation of the size-frequency distribution of North American freshwater fishes.

Skeletal allometry and interlimb scaling patterns in mustelid carnivorans.

To address the effects of an evolutionary increase in body size on long bone skeletal allometry, scaling patterns relating body mass, bone length, limb length, midshaft diameters, and cross-sectional properties of the humerus and femur were analyzed for four species of scansorial mustelids. Humeral and, to a lesser extent, femoral allometry is consistent with expectations of elastic similarity: bone and limb length scale with negative allometry on body mass while bone robusticity (cross-sectional parameters against bone length) scales with strong positive allometry. Differences between fore- and hindlimb scaling patterns, however, are observed, with size-dependent increases in forelimb length and humeral strength and robusticity exceeding those of the hindlimb and femur. It is hypothesized that this greater fore-than hindlimb lengthening results in postural modifications that serve to straighten the hindlimb of larger bodied scansorial mustelids relative to smaller mustelids. Straightening of hindlimb joints would more precisely align the long axis of the femur with peak (vertical) ground reaction forces, thereby accounting for the reduction in relative bending stresses acting on the femur compared to the humerus.

RESPONSES AND CORRELATED RESPONSES TO ARTIFICIAL SELECTION ON THORAX LENGTH IN DROSOPHILA MELANOGASTER.

Two sets of four replicate lines of Drosophila melanogaster were selected for large and small thorax with controls. F, progeny of crosses between the selected lines within each size category showed (a) a reduction in preadult viability in large lines relative to control and small lines when they were cultured at medium or high density in competition with a standard mutant marked competitor stock, and (b) an increase in larval development time in large lines relative to control and small lines. Natural selection for increased body size in adults may therefore be opposed by adverse effects on larval viability. The results are discussed in terms of the developmental mechanisms probably responsible for the change in body size. The preadult survival of the large and control lines was measured at three different temperatures, and there was no evidence for a significant interaction between size and temperature. The observed evolutionary increase in body size in response to reduced temperature in Drosophila must therefore involve either different genes from those subject to selection for size at a single temperature, or a fitness component other than preadult survival. There was no significant asymmetry in response to selection, and thorax length showed heterosis in crosses between the selected lines.

Sexual dimorphism and dental variability in platyrrhine primates

Leutenegger and Cheverud (1982, 1985) propose a hypothesis to explain why larger primates are more sexually dimorphic in body weight and canine size. Their hypothesis states that any factor selecting for an evolutionary increase in body size will produce an increase in sexual dimorphism in any character if either heritability or phenotypic variability is greater in males than in females for that character. They cite no evidence for heritability but give some data to suggest that males are, in fact, more variable than females. We test the latter proposition more fully using measurements on the dentitions of platyrrhine primates. Male and female phenotypic variances are not significantly different in most cases. Cases of greater male phenotypic variance are not limited to sexually dimorphic species. We conclude that the hypothesis of Leutenegger and Cheverud does not explain the observed patterns of dental sexual dimorphism, at least in platyrrhines.

Respiratory gas exchange in ascidans: an almost diffusion limited animal with a cardiovascular system.

The ascidians are a class of the sub Phylum Tunicata, and part of the Phylum Chordata. Characteristically they are sessile marine organisms which have a motile larval stage. They are believed to represent an ancient stock from which other chordates and eventually the vertebrates have evolved. The geometry of the motile (larval) stage and of the ancestral animal is simple in the extreme, supporting the view that this species represents a very early and fundamental form in the evolution of animals. Berrill (1955) has discussed this aspect of the ascidians and has pointed out the evolutionary increase in body size which has occured, from an original animal of less than one mm length, to present day organisms of 10 cm or more. The appearance of a tunic being seen as an adaptation to provide mechanical support and protection for this increased body. The archaic form was a typical diffusion limited animal, whereas it is by no means so obvious that this is so for modern much larger organisms, which possess a cardiovascular system, although rather an unusual one in which the heart regularly and rythmically reverses the direction of flow of blood.KeywordsApparent Diffusion CoefficientHyaluronic AcidFundamental FormPartial Specific VolumeReversible OxygenThese keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.

A General Explanation for Insular Body Size Trends in Terrestrial Vertebrates

The insular body size trends for different vertebrate families are compared. Certain groups such as lagomorphs, bats, artiodactyls, elephants, foxes, raccoons, snakes, and teiid and lacertid lizards are habitually represented by relatively smaller forms on islands. On the other hand, cricetid rodents, iguanid lizards, tortoises, and bears often have races with larger body sizes on islands. Contrary to conventional niche theoretic concepts, in many instances knowledge of the body sizes of some of these animals' insular and mainland competitors does not help explain the difference in that species body size in the 2 places. To account for these divergent size changes I examine optimum body size models that use as the optimization criterion the net energy gained by an organism over a given time period. These models predict that increases in the mean amount of available food should lead to evolutionary increases in body size, but only if body size is not tightly constrained by additional physical or biotic factors: such additional factors might be important if a change in body size alters an animal's effectiveness in finding or handling preferred food items or increases competition with its neighbors. Next using arguments derived from simple non—age—structured 2 species predator—prey models, the availability of food for a given consumer species at equilibrium is compared in theoretical island and mainland situations. Because islands usually contain fewer competitors and the insular physical environment is often more moderate, food availability for colonists is initially expected to be high. On the other hand, as the population grows resources will become depleted. Further, the loss of many predator species on islands may allow consumer densities to increase to such an extent that at equilibrium food may become relatively more limiting for consumers on islands than on the mainland. Whether the supply to demand ratio (S:D) of consumers for their food is ultimately greater or lower will depend on the relative magnitude of these various factors. Within this framework, a necessary condition for island S:D ratios to be greater than on the mainland is that the consumers maintain individual feeding territories. For animals whose body sizes are not tightly bound within narrow limits by physical or competitive restraints, an increase in S:D should lead to an evolutionary increase in body size. Accordingly, a good association is found between the presence or absence of territoriality and the direction of the insular body size shift in a number of different vertebrate groups. Yet there are exceptions which fall into 2 categories: First, if a species' mainland predators preferentially take larger individuals, selection favoring small size may override selection based on optimizing energy input. Such may have been the case for the now extinct mainland relatives of certain giant relictual insular reptiles. Secondly, an animal's body size may be tightly constrained by physical or competitive factors. The body size of island foxes, rattlesnakes, and some lizards appears to be primarily adjusted to the competitive milieu along typical niche theoretic lines. That is, body size may be predicted quite well from knowing the size class of competitors which are absent from an island or from differences in the species' prey—size distribution between island and mainland sites.