Interspecific Allometry Research Articles

Geometric morphometry of the Rhodniusprolixus complex (Hemiptera, Triatominae): patterns of intraspecific and interspecific allometry and their taxonomic implications.

In the subfamily Triatominae, the genus Rhodnius is one of the most studied, not only because of its epidemiological importance, but also because of the difficulty in differentiating its species. Currently, one of the strategies to control Chagas disease, besides other initiatives such as the analysis of donated blood, is focused on fighting the vector. Correctly identifying triatomines is essential for the entomoepidemiological surveillance of Chagas disease. The objective of the present work was to compare the species of the R.prolixus complex using geometric morphometry of hemelytra and heads to evaluate the patterns of intraspecific and interspecific allometry and their taxonomic implications. This method can help in the diagnosis of close species, whose morphological characteristics are insufficient for correct identification. Specimens from five different collections were used, covering the species included in the R.prolixus complex (R.barretti, R.dalessandroi, R.domesticus, R.marabaensis, R.milesi, R.montenegrensis, R.nasutus, R.neglectus, R.neivai, R.prolixus and R.robustus). Morphometric analyses indicated that the hemelytra are not structures with good resolution for separating species and, for this reason, the use of the heads proved to be more adequate for this group (thus allowing differentiation of all species of the R.prolixus complex). The results suggest that R.milesi is a variant of R.neglectus and confirms that R.prolixus and R.robustus are distinct species. Furthermore, we propose the creation of the R.neivai complex comprising R.domesticus and R.neivai.

Length-mass allometries of the larvae of aquatic dipterans: differences between taxa, morphological traits, and methods.

Body mass underpins many ecological processes at the level of individuals, populations, and communities. Often estimated in arthropods from linear morphological traits such as body length or head width, these relationships can vary even between closely related taxa. Length-mass relationships of mosquito (Diptera: Culicidae) larvae are poorly known despite the importance of this family to disease and aquatic ecology. To fill this gap, we measured ontogenetic changes in linear traits (body length, head width, and thorax width) and dry and wet masses and estimated length- and width-mass relationships in larvae of 3 culicid species inhabiting different niches: the tropical Aedes albopictus (Skuse, 1894), the temperate Culex pipiens (Linnaeus, 1758), and the snowmelt Ochlerotatus punctor (Kirby, 1837). We compared our results with published length-mass allometries of other aquatic dipteran larvae. We showed that thorax width and body length, but not head width, reliably predicted body mass for our 3 species. The length-mass allometry slopes in aquatic dipterans varied considerably between and within families but were independent of phylogeny, specimen handling, preservation techniques, and data fitting methods. Slope estimates became less precise with decreasing sample size and size range. To obtain reliable estimates of the allometric slopes, we have thus recommended using data on all larval stages for intraspecific allometries and a wide range of species for interspecific allometries. We also cautioned against the indiscriminate use of length-mass allometries obtained for other taxa or collected at lower taxonomic resolutions, e.g., when using length-mass relationships to estimate biomass production at a given site.

Activity, heart rate, and energy expenditure of a cold-climate mesocarnivore, the Canada lynx (Lynx canadensis)

The energetic consequences of body size, behaviour, and fine-scale environmental variation remain understudied, particularly among free-ranging carnivores, due to logistical and methodological challenges of studying them in the field. Here, we present novel activity, heart rate, and metabolic data on free-ranging Canada lynx (Lynx canadensis Kerr, 1792) to (i) investigate intraspecific patterns of energy expenditure, particularly how they relate to body size, environmental conditions, and activity variation, and (ii) position lynx — a cold-climate, mesocarnivore — within interspecific allometries of carnivore energetics. Lynx demonstrated limited behavioural and metabolic responses to environmental conditions, despite extreme cold and moderate snow depths during our study, but marked body size patterns with larger lynx having higher activity and lower resting heart rate than smaller lynx. Compared with similar-sized carnivores, lynx were less active and had lower heart rate, likely due to their ambush hunting style, but higher energy expenditure, likely due to their cold-climate existence and access to abundant prey. Overall, lynx were more similar to other ambush hunters than to sympatric cold-climate species and mesocarnivores. Our data provide insight into the relative importance of abiotic and biotic drivers of carnivore energetics and the ways in which predators maintain energy balance in variable environments.

Common Patterns of Skull Bone Fusion and Their Potential to Discriminate Different Ontogenetic Stages in Extant Birds

The degree of sutural closure between bones generally allows for the classification of skeleton maturity in tetrapods. In mammals, the sutural closure of skull bones was previously used as proxy to evaluate the ontogenetic stage of single individuals. However, due to temporal variation, this process can be only applied among mammalian subclades, but not for all mammals in general. In contrast, the process of sutural closures in bird skulls could be a more reliable ontogenetic proxy for this clade as adult birds commonly show a generally high degree of bone fusion. To test this, we studied the process of sutural closure in ontogenetic series of 18 extant bird species regarding the presence of an ontogenetic signal and compared the results with changes in skull size and proportions. Univariate analyses indicate that bone fusion happens faster in altricial than in precocial birds. However, the use of PCoA and multivariate regressions reveal that the skull bone fusion follows a common pattern among birds and thus can be used as proxy to identify different ontogenetic stages. In general, the process of sutural closure spreads from posterior to anterior and from ventral to dorsal. In contrast, skull measurements reflect rather interspecific allometry than ontogeny. The used of bone fusion as proxy will help to better identify and compare different stages of maturation in birds, including historical material from osteological collections.

Ontogeny, phylogeny and mechanisms of adaptive changes in evaporative water loss in geckos.

Body size dependence of metabolic rate, body surface and scale morphology complicate disentangling the contribution of these characteristics to adaptive changes in total evaporative water loss (TEWL) of reptiles. To separate adaptive changes from size-related dependence, we compared intra- and interspecific scaling of several candidate traits in eyelid geckos (Eublepharidae), a group exhibiting large variation in body size and TEWL. The intraspecific allometry of TEWL of a eublepharid species fits the geometric surface-mass relationship. However, evolutionary shifts to both higher and lower evaporation were strongly correlated with habitat aridity and cannot be explained by shifts in body size alone. The intraspecific allometry of standard metabolic rate is nearly the same as the interspecific allometry. Unlike for mammals and birds, this pattern rules out respiratory water loss as a driver of the adaptive changes in TEWL among eublepharids. Scale morphology was independent of TEWL variation as well, but the correlation between cutaneous water loss and TEWL suggests a crucial role of skin permeability in adaptation to habitat aridity. Our analyses demonstrate how powerful a comparison between intra- and interspecific allometries can be for detecting body size-dependent mechanisms of adaptive changes in ecophysiological traits correlated with body size.

Interspecific allometry for sexual shape dimorphism: Macroevolution of multivariate sexual phenotypes with application to Rensch's rule.

Allometric trends in the degree of sexual dimorphism with body size have long fascinated evolutionary biologists. Many male-biased clades display more prominent sexual dimorphism in larger taxa (Rensch's rule), with most examples documenting this pattern for body size dimorphism. Although sexual dimorphism in traits other than body size is equally functionally relevant, characterizing allometric patterns of sexual dimorphism in such traits is hampered by lack of an analytical framework that can accommodate multivariate phenotypes. In this article, we derive a multivariate equivalency for investigating trends in sexual dimorphism-relative to overall body size-across taxa and provide a generalized test to determine whether such allometric patterns correspond with Rensch's rule. For univariate linear traits such as body size, our approach yields equivalent results to those from standard procedures, but our test is also capable of detecting trends in multivariate datasets such as shape. Computer simulations reveal that the method displays appropriate statistical properties, and an empirical example in Mediterranean lizards provides the first demonstration of Rensch's rule in a multivariate phenotype (head shape). Our generalized procedure substantially extends the analytical toolkit for investigating macroevolutionary patterns of sexual dimorphism and seeking a better understanding of the processes that underliethem.

The Natural Selection of Metabolism Explains Curvature in Fossil Body Mass Evolution

The natural selection of metabolism and mass can explain inter-specific body mass allometries from prokaryotes to mammals (Witting in Theor Popul Biol 117:23–42, https://doi.org/10.1016/j.tpb.2017.08.005, 2017a), with allometric exponents that depend on the selected metabolism and the packing of home ranges in predominately one (1D), two (2D), or three (3D) spatial dimensions. The predicted exponent for total metabolism for a 2D packing of home ranges increases from 3/4 to 7/4 when the fraction of the inter-specific body mass variation that follows from the natural selection of metabolism increases from zero to one. While a 3/4 exponent is commonly observed for inter-specific comparisons in mammals, a 7/4 exponent has so far not been reported. Yet, I detect the full range of exponents for evolution over time in the fossil record. There are no fossil data for allometric correlations between metabolism and mass, but I show that the allometry $$ \left( {\dot{w} \propto w^{{\hat{\dot{w}}}} } \right) $$ for the rate of evolution $$ \left( {\dot{w} = {\text{ d }}w/{\text{ d }}t} \right) $$ in mass (w) in physical time (t) is given by the underlying set of allometries for life history parameters, including mass-specific metabolism. The $$\hat{\dot{w}}$$ exponent describes the curvature of body mass evolution in time, with predicted values including: 3/2 (2D) for within niche evolution in small horses over 54 million years. 5/4 (2D) and 9/8 (3D) for across niche evolution of maximum mass in four mammalian clades. 3/4 (2D) for fast evolution in large horses, and maximum mass in trunked and terrestrial mammals. And 1 for maximum mass across major lifeforms during 3.5 billion years of evolution along a metabolic bound. These results integrate the inter-specific allometries of existing species with a deeper understanding of their natural selection during evolutionary diversification over millions of years.

Interspecific variations in tree allometry and functional traits in subtropical plantations in southern China.

Mechanical stability against buckling and water transport resistance through xylem vary with increasing tree height. To explore interspecific allometry based on morphological and physiological traits can play a crucial role in revealing their ecological adaptation. Four architectural traits (tree height, diameter at the breast height (DBH), crown width and crown depth) and seven functional traits (specific leaf area (SLA), leaf total carbon concentration (TC), midday leaf water potential, leaf δ13C and δ18O, wood density and xylem water transport efficiency) were measured in Schima superba, Acacia auriculiformis and Eucalyptus citriodora plantations in the subtropical region of China. The mechanical stability declined in the order of S. superba > A. auriculiformis > E. citriodora. Taller species at a given DBH had slender stems and narrower crowns. Smaller leaf δ18O and more efficient xylem water transport were observed in two taller tree species, A. auriculiformis and E. citriodora. Smaller SLA, higher leaf TC and larger leaf area indicated more carbon allocation to leaves of S. superba. The variations in architectural and functional traits with tree allometry among tree species may provide a more complete understanding of species-specific growth strategies in this subtropical region.

The many faces of synapsid cranial allometry

AbstractPrevious studies of cranial shape have established a consistent interspecific allometric pattern relating the relative lengths of the face and braincase regions of the skull within multiple families of mammals. In this interspecific allometry, the facial region of the skull is proportionally longer than the braincase in larger species. The regularity and broad taxonomic occurrence of this allometric pattern suggests that it may have an origin near the base of crown Mammalia, or even deeper in the synapsid or amniote forerunners of mammals. To investigate the possible origins of this allometric pattern, we used geometric morphometric techniques to analyze cranial shape in 194 species of nonmammalian synapsids, which constitute a set of successive outgroups to Mammalia. We recovered a much greater diversity of allometric patterns within nonmammalian synapsids than has been observed in mammals, including several instances similar to the mammalian pattern. However, we found no evidence of the mammalian pattern within Therocephalia and nonmammalian Cynodontia, the synapsids most closely related to mammals. This suggests that the mammalian allometric pattern arose somewhere within Mammaliaformes, rather than within nonmammalian synapsids. Further investigation using an ontogenetic series of the anomodontDiictodon felicepsshows that the pattern of interspecific allometry within anomodonts parallels the ontogenetic trajectory ofDiictodon.This indicates that in at least some synapsids, allometric patterns associated with ontogeny may provide a “path of least resistance” for interspecific variation, a mechanism that we suggest produces the interspecific allometric pattern observed in mammals.

Evolutionary negative allometry of orthopteran hind femur length is a general phenomenon

1. Body size influences the way that organisms both perform their locomotor activities and perceive their environment. Allometry of insect legs with respect to body size is affected by many factors such as ontogenetic constraints and natural selection. Negative allometry, positive allometry, or isometry could result from different mechanisms influencing leg length and locomotion performance. 2. We tested three main models of hind femur length allometry (natural selection for jumping performance, ontogenetic constraint, and the size-grain model) in Orthoptera, a Polyneopteran order with large size range and high habitat and lifestyle diversification. We used a data set of 1549 species including members of both suborders, Ensifera and Caelifera, and many representative families using a Linear Mixed Model approach, and Reduced Major Axis and Ordinary Least Squares regression to explore evolutionary interspecific allometry in this order. 3. Our results showed a generalized trend of negative allometry (leg length decreases with body size increase) at the ordinal, subordinal, and familial levels, contrary to the expectations of the size-grain model and supporting our main hypothesis of a common ancestral developmental pattern regulating leg negative allometry in Orthoptera. 4. The conservation of a common hind leg allometric pattern in Orthoptera provides a basic framework to study adaptation of hind limbs to different habitats and lifestyles, and has important implications for the analysis of ecogeographic and evolutionary rules.

The natural selection of metabolism and mass selects allometric transitions from prokaryotes to mammals

The exponents of inter-specific allometries for several life history (metabolism, lifespan, reproductive rate, survival) and ecological (population density, home range) traits may evolve from the spatial dimensionality (d) of the intra-specific interactive competition that selects net assimilated energy into mass, with 1∕4 exponents being the two-dimensional (2D) case of the more general 1∕2d (Witting, 1995). While the exponents for mass-specific metabolism cluster around the predicted -1/4 and -1/6 in terrestrial and pelagic vertebrates, the allometries of mobile organisms are more diverse than the prediction. An exponent around zero has been reported for protists and protozoa (Makarieva et al., 2005, 2008), and the exponent appears to be strongly positive in prokaryotes with a value of about 5/6 (DeLong et al., 2010).I show that the natural selection of metabolism and mass is sufficient to explain exponents for mass-specific metabolism that decline from 5/6 over zero to −1∕6 in 3D, and from 3/4 over zero to −1∕4 in 2D. These results suggest that mass-specific metabolism is selected as the pace of the resource handling that generates net energy for self-replication and the selection of mass, with the decline in the metabolic exponent following from a decline in the importance of mass-specific metabolism for the selection of mass. The body mass variation in prokaryotes is found to be selected from primary variation in mass-specific metabolism, while the variation in multicellular animals is selected from primary variation in the handling and/or densities of the underlying resources, with protists and protozoa being selected as an intermediate lifeform.

Modeling the physiology of the aquatic temnospondyl <i>Archegosaurus decheni</i> from the early Permian of Germany

Abstract. Physiological aspects like heat balance, gas exchange, osmoregulation, and digestion of the early Permian aquatic temnospondyl Archegosaurus decheni, which lived in a tropical freshwater lake, are assessed based on osteological correlates of physiologically relevant soft-tissue organs and by physiological estimations analogous to air-breathing fishes. Body mass (M) of an adult Archegosaurus with an overall body length of more than 1 m is estimated as 7 kg using graphic double integration. Standard metabolic rate (SMR) at 20 °C (12 kJ h−1) and active metabolic rate (AMR) at 25 °C (47 kJ h−1) were estimated according to the interspecific allometry of metabolic rate (measured as oxygen consumption) of all fish (VO2 = 4. 8 M0. 88) and form the basis for most of the subsequent estimations. Archegosaurus is interpreted as a facultative air breather that got O2 from the internal gills at rest in well-aerated water but relied on its lungs for O2 uptake in times of activity and hypoxia. The bulk of CO2 was always eliminated via the gills. Our estimations suggest that if Archegosaurus did not have gills and released 100 % CO2 from its lungs, it would have to breathe much more frequently to release enough CO2 relative to the lung ventilation required for just O2 uptake. Estimations of absorption and assimilation in the digestive tract of Archegosaurus suggest that an adult had to eat about six middle-sized specimens of the acanthodian fish Acanthodes (ca. 8 cm body length) per day to meet its energy demands. Archegosaurus is regarded as an ammonotelic animal that excreted ammonia (NH3) directly to the water through the gills and the skin, and these diffusional routes dominated nitrogen excretion by the kidneys as urine. Osmotic influx of water through the gills had to be compensated for by production of dilute, hypoosmotic urine by the kidneys. Whereas Archegosaurus has long been regarded as a salamander-like animal, there is evidence that its physiology was more fish- than tetrapod-like in many respects.

Sex differences in gait utilization and energy metabolism during terrestrial locomotion in two varieties of chicken (Gallus gallus domesticus) selected for different body size.

ABSTRACTIn leghorn chickens (Gallus gallus domesticus) of standard breed (large) and bantam (small) varieties, artificial selection has led to females being permanently gravid and sexual selection has led to male-biased size dimorphism. Using respirometry, videography and morphological measurements, sex and variety differences in metabolic cost of locomotion, gait utilisation and maximum sustainable speed (Umax) were investigated during treadmill locomotion. Males were capable of greater Umax than females and used a grounded running gait at high speeds, which was only observed in a few bantam females and no standard breed females. Body mass accounted for variation in the incremental increase in metabolic power with speed between the varieties, but not the sexes. For the first time in an avian species, a greater mass-specific incremental cost of locomotion, and minimum measured cost of transport (CoTmin) were found in males than in females. Furthermore, in both varieties, the female CoTmin was lower than predicted from interspecific allometry. Even when compared at equivalent speeds (using Froude number), CoT decreased more rapidly in females than in males. These trends were common to both varieties despite a more upright limb in females than in males in the standard breed, and a lack of dimorphism in posture in the bantam variety. Females may possess compensatory adaptations for metabolic efficiency during gravidity (e.g. in muscle specialization/posture/kinematics). Furthermore, the elevated power at faster speeds in males may be linked to their muscle properties being suited to inter-male aggressive combat.

Intraspecific scaling of the minimum metabolic cost of transport in leghorn chickens (Gallus gallus domesticus): links with limb kinematics, morphometrics and posture.

ABSTRACTThe minimum metabolic cost of transport (CoTmin; J kg−1 m−1) scales negatively with increasing body mass (∝Mb−1/3) across species from a wide range of taxa associated with marked differences in body plan. At the intraspecific level, or between closely related species, however, CoTmin does not always scale with Mb. Similarity in physiology, dynamics of movement, skeletal geometry and posture between closely related individuals is thought to be responsible for this phenomenon, despite the fact that energetic, kinematic and morphometric data are rarely collected together. We examined the relationship between these integrated components of locomotion in leghorn chickens (Gallus gallus domesticus) selectively bred for large and bantam (miniature) varieties. Interspecific allometry predicts a CoTmin ∼16% greater in bantams compared with the larger variety. However, despite 38% and 23% differences in Mb and leg length, respectively, the two varieties shared an identical walking CoTmin, independent of speed and equal to the allometric prediction derived from interspecific data for the larger variety. Furthermore, the two varieties moved with dynamic similarity and shared geometrically similar appendicular and axial skeletons. Hip height, however, did not scale geometrically and the smaller variety had more erect limbs, contrary to interspecific scaling trends. The lower than predicted CoTmin in bantams for their Mb was associated with both the more erect posture and a lower cost per stride (J kg−1 stride−1). Therefore, our findings are consistent with the notion that a more erect limb is associated with a lower CoTmin and with the previous assumption that similarity in skeletal shape, inherently linked to walking dynamics, is associated with similarity in CoTmin.

The Biology of Gliding in Flying Lizards (Genus Draco) and their Fossil and Extant Analogs

The flying lizards of the genus Draco are among the most remarkable and successful clades of gliding vertebrates. Here, we evaluate the evolution of gliding in Draco and other lizards, describe the suite of morphological innovations that characterize Draco, discuss the ecological context of gliding in this genus, describe functions of their patagial membranes that are not related to gliding, and summarize the interspecific allometry of the Draco gliding apparatus, as well as the corresponding consequences for their now empirically quantified gliding performance. Several fossil reptilian lineages had morphologies similar to that of modern Draco, with patagial membranes supported by elongated ribs or rib-like dermal structures. Using Draco's snout-vent length/mass relationships, we provide improved estimates of wing loading for three of these fossil gliders (Icarosaurus seifkeri, Kuehneosaurus sp., Coelurosauravus elivensis) and then estimate absolute gliding performance for each taxon by extrapolating from Draco's wing loading/glide performance relationship. We find that I. seifkeri likely represented the best nonflapping terrestrial vertebrate glider yet described, whereas the larger Kuehneosaurus and Coelurosauravus probably required high descent velocities to achieve sufficient lift for gliding, with commensurately greater height loss with each glide.

Mammalian Metabolic Allometry: Do Intraspecific Variation, Phylogeny, and Regression Models Matter?

Power scaling relationships between body mass and organismal traits are fundamental to biology. Compilations of mammalian masses and basal metabolic rates date back over a century and are used both to support and to assail the universal quarter-power scaling invoked by the metabolic theory of ecology. However, the slope of this interspecific allometry is typically estimated without accounting for intraspecific variation in body mass or phylogenetic constraints on metabolism. We returned to the original literature and culled nearly all unique measurements of body mass and basal metabolism for 695 mammal species and (1) phylogenetically corrected the data using the fullest available phylogeny, (2) applied several different regression analyses, (3) resampled regressions by drawing randomly selected species from each of the polytomies in the phylogenetic hypothesis at each iteration, and (4) ran these same analyses independently on separate clades. Overall, 95% confidence intervals of slope estimates frequently did not include 0.75, and clade-specific slopes varied from 0.5 to 0.85, depending on the clade and regression model. Our approach reveals that the choice of analytical model has a systematic influence on the estimated allometry, but irrespective of the model applied, we find little support for a universal metabolic rate-body mass scaling relationship.

Interspecific allometry of morphological traits among trematode parasites: selection and constraints

Developmental constraints and selective pressures interact to determine the strength of allometric scaling relationships between body size and the size of morphological traits among related species. Different traits are expected to relate to body size with different scaling exponents, depending on how their function changes disproportionately with increasing body size. For trematodes parasitic in vertebrate guts, the risk of being dislodged should increase disproportionately with body size, whereas basic physiological functions are more likely to increase in proportion to changes in body size. Allometric scaling exponents for attachment structures should thus be higher than those for other structures and should be higher for trematode families using endothermic hosts than for those using ectotherms, given the feeding and digestive characteristics of these hosts. These predictions are tested with data on 363 species from 13 trematode families. Sizes of four morphological structures were investigated, two associated with attachment (oral and ventral suckers) and the other two with feeding and reproduction (pharynx and cirrus sac). The scaling exponents obtained were generally low, the majority falling between 0.2 and 0.5. There were no consistent differences within families between the magnitude of scaling exponents for different structures. Also, there was no difference in the values of scaling exponents between families exploiting endothermic hosts and those using ectotherms. There were strong correlations across families between the values of the scaling exponents for the oral sucker, the ventral sucker and the pharynx: in families where the size of one trait increases relatively steeply as a function of body size, the same is generally true of the other traits. These results suggest either that developmental constraints link several morphological features independently of their specific roles or that similar selection pressures operate on different structures, leading to covariation of scaling exponents. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 533‐540.

Bias in interspecific allometry: examples from morphological scaling in varanid lizards

The traditional approach to allometric analysis entails the fitting of a straight line to logarithmic transformations of the data, after which parameters in a two-parameter allometric equation are estimated by back-transformation to the original scale. We re-examined published data for dimensions of the limbs in 22 species of varanid lizards to illustrate the biases that can be introduced into allometric analyses by applying the aforementioned protocol. Statistical models fit to the original data by linear and nonlinear regression conformed better with underlying assumptions than did models obtained by back-transformation from logarithms, and the former generally were better than the latter for describing limb dimensions over the full range in body size. Allometric exponents estimated by the traditional method therefore were based on inappropriate and inaccurate statistical models and, consequently, were biased and misleading. Investigators can avoid problems such as these by performing preliminary graphical and statistical analyses on data in their original scale and by validating the fitted model. Logarithmic transformations should be used sparingly and only for cause.

Quantitative estimates of visual performance features in fossil birds

Eyeball structures such as the lens diameter (LD) and axial length are generally assumed to be highly correlated with optically meaningful parameters. However, these optical constraints on eyeball macroanatomy have never been tested explicitly. Tradeoffs between benefits of improved visual performance and cost of adaptation from an increase of tissue production predict that when eyeball size increases, optical parameters such as posterior nodal distance and maximum entrance pupil diameter should increase isometrically with eyeball axial length and LD, respectively. Here I show quantitatively that the interspecific allometry of the avian eye largely follows this predicted isometry. Additionally, I elaborate a method to estimate optically significant eyeball soft-tissue dimensions from scleral ring and orbit morphology based on analyses of interspecific allometry in Aves. The stringent correlations between avian eyeball morphology and optical function render this system ideal for the analysis of form-function relationships and allow for an accurate estimate of optically significant eyeball soft-tissue dimensions such as diameter, axial length, and LD in fossil species.

Ant colony size and the scaling of reproductive effort

1 Reproductive effort typically scales as mass0·75 in unitary organisms, but less is known about such scaling in colonial organisms. 2 I compiled data on worker and reproductive number at maturity for 65 ant species and found an interspecific allometry (alate number = worker number0·73) whose exponent was significantly < 1, even after a phylogenetic correction. 3 When I analyzed 15 species for which biomass data were available, I found an interspecific isometry (alate biomass = worker biomass0·89) whose exponent was not significantly different from 1. Analysis of maximum species biomass values, rather than averages, strengthened this isometry, yielding a slope b = 1·01 that was also not distinguishable from 1. 4 Species with larger colony size at reproduction tended to couple investment in proportionately fewer alates with investment in larger male and female alates. 5 This comparative analysis suggests a trade-off between alate size and number, and provides a framework for studying the diversity of colony life histories and the mechanisms generating allometries.