Insular dwarfism in horses from the Aegean Sea and the Japanese archipelago.

The horse (Equus caballus) varieties from Skyros and Rhodes islands (Greece) in the Aegean archipelago are extremely small, reaching shoulder heights of only about 1 m. Furthermore, the Japanese archipelago is home to eight small, native horse breeds. We investigated the evolutionary morphology and provided a review of historical documentations of these horses of cultural interest in Greece and Japan, thus providing a comparison of the independent evolution of small size in islands. We integrate cranial data from historical literature with data from newly gathered and curated skulls and analyse a measurement dataset featuring various domestic and mainland horse breeds and varieties. We use non-invasive imaging to study and measure 3D models of the bony labyrinth, housing the inner ear, and the braincase endocast. When considering the effects of allometry by regressing each PC1 scores (for each set of measurements) with the cranial geometric mean from linear measurements as a body size proxy, we show that size explains a large amount of the shape variation in horse crania, the bony labyrinths and brain endocasts. We found high intrabreed variation in all the analysed datasets. Globally, there are at least 30 distinct horse populations on islands, offering the chance to further study processes of convergence in morphological divergence and evaluate the effect of drift and the environment. Supplementary Information: The online version contains supplementary material available at 10.1007/s42991-024-00408-4.

Eco-geographic and sexual variation of the ribcage in Homo sapiens.

Up to now, Allen and Bergmann's rules have been studied in modern humans by analyzing differences in limb length, height, or body mass. However, there are no publications studying the effects of latitude in the 3D configuration of the ribcage. To assess this issue, we digitally reconstructed the ribcages of a balanced sample of 109 adult individuals of global distribution. Shape and size of the ribcage was quantified using geometric morphometrics. Our results show that the ribcage belonging to tropical individuals is smaller and slenderer compared to others living in higher latitudes, which is in line with Allen and Bergmann's rules and suggests an allometric relationship between size and shape. Although sexual dimorphism was observed in the whole sample, significant differences were only found in tropical populations. Our proposal is that, apart from potential sexual selection, avoiding heat loss might be the limiting factor for sexual dimorphism in cold-adapted populations.

Tip-to-base conduit widening remains consistent across cambial age and climates in Fagus sylvatica L.

Water transport, mechanical support and storage are the vital functions provided by the xylem. These functions are carried out by different cells, exhibiting significant anatomical variation not only within species but also within individual trees. In this study, we used a comprehensive dataset to investigate the consistency of predicted hydraulic vessel diameter widening values in relation to the distance from the tree apex, represented by the relationship Dh ∝ Lß (where Dh is the hydraulic vessel diameter, L the distance from the stem apex and ß the scaling exponent). Our analysis involved ten Fagus sylvatica L. trees sampled at two distinct sites in the Italian Apennines. Our results strongly emphasize that vessel diameter follows a predictable pattern with the distance from the stem apex, and ß ~ 0.20 remains consistent across cambial age and climates. This finding supports the hypothesis that trees do not alter their axial configuration represented by scaling of vessel diameter to compensate for hydraulic limitations imposed by tree height during growth. The study further indicates that within-tree variability significantly contributes to the overall variance of the vessel diameter-stem length exponent. Understanding the factors that contribute to the intraindividual variability in the widening exponent is essential, particularly in relation to interspecific responses and adaptations to drought stress.

Interplay between drought and plant viruses co-infecting melon plants.

Drought affects crops directly, and indirectly by affecting the activity of insect pests and the transmitted pathogens. Here, we established an experiment with well-watered or water-stressed melon plants, later single infected with either cucumber mosaic virus (CMV: non-persistent), or cucurbit aphid-borne yellow virus (CABYV: persistent), or both CMV and CABYV, and mock-inoculated control. We tested whether i) the relation between CMV and CABYV is additive, and ii) the relationship between water stress and virus infection is antagonistic, i.e., water stress primes plants for enhanced tolerance to virus infection. Water stress increased leaf greenness and temperature, and reduced leaf water potential, shoot biomass, stem dimensions, rate of flowering, CABYV symptom severity, and marketable fruit yield. Virus infection reduced leaf water potential transiently in single infected plants and persistently until harvest in double-infected plants. Double-virus infection caused the largest and synergistic reduction of marketable fruit yield. The relationship between water regime and virus treatment was additive in 12 out of 15 traits at harvest, with interactions for leaf water content, leaf:stem ratio, and fruit set. We conclude that both virus-virus relations in double infection and virus-drought relations cannot be generalized because they vary with virus, trait, and plant ontogeny.

Carpal variability and asymmetry in limb reduced Western lesser sirens (Siren nettingi).

Trait functionality can act as a constraint on morphological development. Traits that become vestigialized can exhibit unstable developmental patterns such as fluctuating asymmetry (FA) and variation in populations. We use clearing and staining along with morphometric analyzes to compare FA and allometry of limbs in Western lesser sirens (Siren nettingi) to Ouachita dusky salamanders (Desmognathus brimleyorum). Our results describe new carpal phenotypes and carpal asymmetry in our sample of S. nettingi. However, we found no significant evidence of limb length asymmetry in S. nettingi. The degree of relative limb asymmetry correlates inversely with body size in both of our samples. This work provides strong evidence of increased mesopodal variation within a population of S. nettingi. Our work provides a basis for further study of a broader range of morphological traits across salamanders.

Sexually dimorphic eye size in dragonfishes, a response to a bioluminescent signalling gap.

Deep-sea fishes must overcome extremely large nearest-neighbour distances and darkness to find mates. Sexual dimorphism in the size of luminescent structures in many deep-sea taxa, including dragonfishes (family Stomiidae), indicates reproductive behaviours may be mediated by visual signalling. This presents a paradox: if male photophores are larger, females may find males at shorter distances than males find females. Solutions to this gap may include females closing this gap or by males gathering more photons with a larger eye. We examine the eye size of two species of dragonfishes (Malacosteus niger and Phostomias guernei) for sexual dimorphism and employ a model of detection distance to evaluate the potential for such dimorphism to bridge the detection gap. This model incorporates the flux of sexually dimorphic postorbital photophores and eye lens size to predict detection distances. In both species, we found a significant visual detection gap in which females find males before males find females and that male lens size is larger, marking the second known case of size dimorphism in the actinopterygian visual system. Our results indicate the larger eye affords males a significant improvement in detection distance. We conclude that this dimorphic phenotype may have evolved to close the detection gap.

Beyond CREA: Evolutionary patterns of non-allometric shape variation and divergence in a highly allometric clade of murine rodents.

The shared functions of the skull are thought to result in common evolutionary patterns in mammalian cranial shape. Craniofacial evolutionary allometry (CREA) is a particularly prominent pattern where larger species display proportionally elongate facial skeletons and smaller braincases. It was recently proposed that CREA arises from biomechanical effects of cranial scaling when diets are similar. Thus, deviations from CREA should occur with changes in cranial biomechanics, for example due to dietary change. Here, we test this using 3D geometric morphometric analysis in a dataset of Australian murine crania, which are highly allometric. We contrast allometric and non-allometric variation in the cranium by comparing evolutionary mode, allometry, ordinations, as well as allometry, integration, and modularity in functional modules. We found evidence of stabilising selection in allometry-containing and size-free shape, and substantial non-allometric variation aligned with dietary specialisation in parallel with CREA. Integration among cranial modules was higher, and modularity lower, with size included, but integration between rostrum and cranial vault, which are involved in the CREA pattern, dropped dramatically after size removal. Our results thus support the hypothesis that CREA is a composite arising from selection on cranial function, with substantial non-allometric shape variation occurring alongside CREA where dietary specialisation impacts selection on gnawing function. This emphasises the need to research mammalian cranial evolution in the context of allometric and non-allometric selection on biomechanical function.

Induced Power Scaling Alone Cannot Explain Griffenfly Gigantism.

Paleozoic skies were ruled by extinct odonatopteran insects called 'griffenflies', some with wingspans three times that of the largest extant dragonflies and ten times that of common extant dragonflies. Previous studies suggested that flight was possible for larger fliers because of higher atmospheric oxygen levels that would have increased air density. We use actuator disk theory to evaluate this hypothesis. Actuator disk theory gives similar estimates of induced power as has been estimated for micro-air vehicles based on insect flight. We calculate that for a given mass of griffenfly, and assuming isometry, a higher density atmosphere would only have reduced the induced power required to hover by 11%, which would have supported a flyer 3% larger in linear dimensions. Steady level forward flight would have further reduced induced power but could only account for a flier 5% larger in linear dimensions. Further accounting for the higher power available due to high oxygen air, and assuming isometry, we calculate that the largest flyer hovering would have been only 1.19 times longer than extant dragonflies. We also consider known allometry in dragonflies and estimated allometry in extinct griffenflies. But such allometry only increases flyer size to 1.22 times longer while hovering. We also consider profile and parasite power, but both would have been higher in denser air and thus would not have enhanced the flyability of larger griffenflies. The largest meganeurid griffenflies might have adjusted flight behaviors to reduce power required. Alternatively, the scaling of flight muscle power may have been sufficient to support the power demands of large griffenflies. In literature estimates, mass-specific power output scales as mass0.24 in extant dragonflies. We need only more conservatively assume that mass-specific muscle power scales with mass0, when combined with higher oxygen concentrations and induced power reductions in higher density air to explain griffenflies 3.4 times larger than extant odonates. Experimental measurement of flight muscle power scaling in odonates is necessary to test this hypothesis.

Extreme range in adult body size reveals hidden trade-offs among sexually selected traits.

Sexually selected weapons used to monopolize mating opportunities are predicted to trade-off with traits used in competition for fertilization. Yet, the limited size range typically found among adults of a species often precludes clear comparisons between population-level and individual-level relative trait investment. The jousting weevil, Brentus anchorago (Coleoptera: Brentidae), varies more than 26-fold in body mass, which is among the most extreme adult body size ranges of any solitary terrestrial species. We reveal a trade-off at a population level: hypermetric scaling in male weapons (slope = 1.59) and a closely mirrored reversal in allocation to postcopulatory traits (slope = 0.54). Yet, at the individual level, we find the opposite pattern; males that invest relatively more in weapons for their size class also invest more in postcopulatory traits. Across 36 dung beetle and 41 brentine weevil species, we find the allometric slope explains more trait variation at larger body size ranges; in brentines, population-level scaling patterns become more detectable in species with a larger range in adult body size. Our findings reveal that population-level allometries and individual-level trade-offs can both be important in shaping relative trait allocation; we highlight that the adult body size range is rarely examined but may be integral to gaining a deeper understanding of trade-offs in reproductive allocation.

Small- to medium-sized mammals show greater morphological disparity in cervical than lumbar vertebrae across different terrestrial modes of locomotion.

During mammalian terrestrial locomotion, body flexibility facilitated by the vertebral column is expected to be correlated with observed modes of locomotion, known as gait (e.g., sprawl, trot, hop, bound, gallop). In small- to medium-sized mammals (average weight up to 5 kg), the relationship between locomotive mode and vertebral morphology is largely unexplored. Here we studied the vertebral column from 46 small- to medium-sized mammals. Nine vertebrae across cervical, thoracic, and lumbar regions were chosen to represent the whole vertebral column. Vertebra shape was analysed using three-dimensional geometric morphometrics with the phylogenetic comparative method. We also applied the multi-block method, which can consider all vertebrae as a single structure for analysis. We calculated morphological disparity, phylogenetic signal, and evaluated the effects of allometry and gait on vertebral shape. We also investigated the pattern of integration in the column. We found the cervical vertebrae show the highest degree of morphological disparity, and the first thoracic vertebra shows the highest phylogenetic signal. A significant effect of gait type on vertebrae shape was found, with the lumbar vertebrae having the strongest correlation; but this effect was not significant after taking phylogeny into account. On the other hand, allometry has a significant effect on all vertebrae regardless of the contribution from phylogeny. The regions showed differing degrees of integration, with cervical vertebrae most strongly correlated. With these results, we have revealed novel information that cannot be captured from study of a single vertebra alone: although the lumbar vertebrae are the most correlated with gait, the cervical vertebrae are more morphologically diverse and drive the diversity among species when considering whole column shape.

Spotting mistakes: Reappraisal of Spotted Drum Stellifer punctatissimus (Meek & Hildebrand, 1925) (Teleostei: Sciaenidae) reveals species misidentification trends and suggests latitudinal sexual dimorphism.

A major part of the described species is understudied, falling into the Linnean shortfall. This is a major concern for cryptic species, which require integrative approaches to better evaluate their diversity. We conducted morphological analyses using specimens of Stellifer punctatissimus, S. gomezi, and S. menezesi to reassess their taxonomical identity. We evaluated the allometric and sexual components of the morphology of the Stellifer punctatissimus species complex, and tested and discussed species hypotheses. The combined evidence of our work and previous studies agrees with the current morphological hypothesis of three species, as opposed to the two-lineage molecular hypothesis. However, as cryptic species, they overlap in most their traits, especially females and juveniles. Previously unaccounted variation of allometric and sexually dimorphic characters in this species complex revealed a confounding effect that might explain past and current taxonomic errors. Taxonomical practice of using body depth as a diagnostic character had led to juveniles and females being, respectively, described as a different species or wrongfully identified. Hence, taxonomical studies demand better assessment of allometric and sexual dimorphism components. Herein, we present new characters in a key to the Atlantic species of Stellifer, which disclose size and sexual variation unnoticed in previous studies. The contrasting growth patterns among these species could imply distinct habitat use. As a result, it could be suggested that such species are under different threats, which highlights the need of differential management and conservation strategies.

A temporal allocation of amino acid resources ensures fitness and body allometry in Drosophila.

Organisms have evolved strategies to store resources and overcome periods of low or no nutrient access, including transient shortages or longer non-feeding developmental transitions. Holometabolous insects like Drosophila represent an attractive model to study resource allocation during development because they alternate feeding and non-feeding periods. Amino acids are essential components for tissue growth and renewal, but the strategies used for their storage remain largely unexplored. Here, we characterize the molecular mechanisms for the temporal production, accumulation, and use of specific storage proteins called hexamerins, and demonstrate their role in ensuring tissue formation and adult fitness. Moreover, we show that preventing hexamerin stores enhances the growth of early-developing organs while compromising the emergence of late-forming ones, consequently altering body allometry.

Individual energetics scale up to community coexistence: Movement, metabolism and biodiversity dynamics in fragmented landscapes.

Unravelling the intricate mechanisms that govern community coexistence remains a daunting challenge, particularly amidst ongoing environmental change. Individual physiology and metabolism are often studied to understand the response of individual animals to environmental change. However, this perspective is currently largely lacking in community ecology. We argue that the integration of individual metabolism into community theory can offer new insights into coexistence. We present the first individual-based metabolic community model for a terrestrial mammal community to simulate energy dynamics and home range behaviour in different environments. Using this model, we investigate how ecologically similar species coexist and maintain their energy balance under food competition. Only if individuals of different species are able to balance their incoming and outgoing energy over the long-term will they be able to coexist. After thoroughly testing and validating the model against real-world patterns such as of home range dynamics and field metabolic rates, we applied it as a case study to scenarios of habitat fragmentation - a widely discussed topic in biodiversity research. First, comparing single-species simulations with community simulations, we find that the effect of habitat fragmentation on populations is strongly context-dependent. While populations of species living alone in the landscape were mostly positively affected by fragmentation, the diversity of a community of species was highest under medium fragmentation scenarios. Under medium fragmentation, energy balance and reproductive investment were also most similar among species. We therefore suggest that similarity in energy balance among species promotes coexistence. We argue that energetics should be part of community ecology theory, as the relative energetic status and reproductive investment can reveal why and under what environmental conditions coexistence is likely to occur. As a result, landscapes can potentially be protected and designed to maximize coexistence. The metabolic community model presented here can be a promising tool to investigate other scenarios of environmental change or other species communities to further disentangle global change effects and preserve biodiversity.

Experimental evolution under predation reduces body size in dung flies but courtship displays persist in males (Diptera: Sepsidae).

Exaggerated sexual traits, such as ornaments and courtship displays, are crucial for mate acquisition in many species and are often subject to directional runaway selection. However, in the face of high predation risk, natural selection can result in a reduction of conspicuous precopulatory displays to avoid detection by potential predators. Sexual selection may then favour increased investment in inconspicuous postcopulatory traits. Here, we investigated the transgenerational effects of predation on precopulatory male courtship and postcopulatory sexual traits (testes size, sperm length) in a dung fly, Sepsis punctum (Sepsidae). Behavioural assays prior to selection document a marked decrease in male courtship displays in the presence of a predator, the Asian Ant Mantis (Odontomantis planiceps). However, after ten generations of experimental evolution, flies exhibited a marked increase in courtship, both in the absence and presence of a predator. Additionally, under sustained predation pressure, male and female body size decreased but male postcopulatory traits were not significantly affected. These results suggest that precopulatory courtship can be under strong sexual selection even in the face of predation pressure. Larger flies were more susceptible to predation, and there could be canalisation of postcopulatory traits that are crucial for fertilisation.

Size matters in metabolic scaling: Critical role of the thermodynamic efficiency of ATP synthesis and its dependence on mitochondrial H+ leak across mammalian species.

In this last article of the trilogy, the unified biothermokinetic theory of ATP synthesis developed in the previous two papers is applied to a major problem in comparative physiology, biochemistry, and ecology-that of metabolic scaling as a function of body mass across species. A clear distinction is made between intraspecific and interspecific relationships in energy metabolism, clearing up confusion that had existed from the very beginning since Kleiber first proposed his mouse-to-elephant rule almost a century ago. It is shown that the overall mass exponent of basal/standard metabolic rate in the allometric relationship [Formula: see text] is composed of two parts, one emerging from the relative intraspecific constancy of the slope (b), and the other (b') arising from the interspecific variation of the mass coefficient, a(M) with body size. Quantitative analysis is shown to reveal the hidden underlying relationship followed by the interspecific mass coefficient, a(M)=P0M0.10, and a universal value of P0=3.23 watts, W is derived from empirical data on mammals from mouse to cattle. The above relationship is shown to be understood only within an evolutionary biological context, and provides a physiological explanation for Cope's rule. The analysis also helps in fundamentally understanding how variability and a diversity of scaling exponents arises in allometric relations in biology and ecology. Next, a molecular-level understanding of the scaling of metabolism across mammalian species is shown to be obtained by consideration of the thermodynamic efficiency of ATP synthesis η, taking mitochondrial proton leak as a major determinant of basal metabolic rate in biosystems. An iterative solution is obtained by solving the mathematical equations of the biothermokinetic ATP theory, and the key thermodynamic parameters, e.g. the degree of coupling q, the operative P/O ratio, and the metabolic efficiency of ATP synthesis η are quantitatively evaluated for mammals from rat to cattle. Increases in η (by ∼15%) over a 2000-fold body size range from rat to cattle, primarily arising from an ∼3-fold decrease in the mitochondrial H+ leak rate are quantified by the unified ATP theory. Biochemical and mechanistic consequences for the interpretation of basal metabolism, and the various molecular implications arising are discussed in detail. The results are extended to maximum metabolic rate, and interpreted mathematically as a limiting case of the general ATP theory. The limitations of the analysis are pointed out. In sum, a comprehensive quantitative analysis based on the unified biothermokinetic theory of ATP synthesis is shown to solve a central problem in biology, physiology, and ecology on the scaling of energy metabolism with body size.

Coping with environmental degradation: Physiological and morphological adjustments of wild mangrove fish to decades of aquaculture-induced nutrient enrichment.

The impact of eutrophication on wild fish individuals is rarely reported. We compared physiological and morphological traits of Siganus lineatus chronically exposed to aquaculture-induced eutrophication in the wild with individuals living at a control site. Eutrophication at the impacted site was confirmed by elevated organic matter (up to 150 % higher), phytoplankton (up to 7 times higher), and reduced oxygen (up to 60 % lower). Physiological and morphological traits of S. lineatus differed significantly between the two sites. Fish from the impacted site exhibited elevated hypoxia tolerance, increased gill surface area, shorter oxygen diffusion distances, and altered blood oxygen-carrying capacity. Elevated blood lactate and scope for anaerobic ATP production were observed, suggesting enhanced survival below critical oxygen levels. A significant 8.5 % increase in metabolic costs and altered allometric scaling, related to environmental degradation, were recorded. Our study underscores eutrophication's profound impact at the organism-level and the importance to mitigate it.

Bacterial sepsis triggers stronger transcriptomic immune responses in larger primates.

Empirical data relating body mass to immune defence against infections remain limited. Although the metabolic theory of ecology predicts that larger organisms would have weaker immune responses, recent studies have suggested that the opposite may be true. These discoveries have led to the safety factor hypothesis, which proposes that larger organisms have evolved stronger immune defences because they carry greater risks of exposure to pathogens and parasites. In this study, we simulated sepsis by exposing blood from nine primate species to a bacterial lipopolysaccharide (LPS), measured the relative expression of immune and other genes using RNAseq, and fitted phylogenetic models to determine how gene expression was related to body mass. In contrast to non-immune-annotated genes, we discovered hypermetric scaling in the LPS-induced expression of innate immune genes, such that large primates had a disproportionately greater increase in gene expression of immune genes compared to small primates. Hypermetric immune gene expression appears to support the safety factor hypothesis, though this pattern may represent a balanced evolutionary mechanism to compensate for lower per-transcript immunological effectiveness. This study contributes to the growing body of immune allometry research, highlighting its importance in understanding the complex interplay between body size and immunity over evolutionary timescales.

Advancing Fine Branch Biomass Estimation with Lidar and Structural Models.

BACKGROUND AND AIMS: Lidar is a promising tool for fast and accurate measurements of trees. There are several approaches to estimate aboveground woody biomass using lidar point clouds. One of the most widely used methods involves fitting geometric primitives (e.g. cylinders) to the point cloud, thereby reconstructing both the geometry and topology of the tree. However, current algorithms are not suited for accurate estimation of the volume of finer branches, because of the unreliable point dispersions from e.g. beam footprint compared to the structure diameter. METHODS: We propose a new method that couples point cloud-based skeletonization and multi-linear statistical modelling based on structural data to make a model (structural model) that accurately estimates the aboveground woody biomass of trees from high-quality lidar point clouds, including finer branches. The structural model was tested at segment, axis, and branch level, and compared to a cylinder fitting algorithm and to the pipe model theory. KEY RESULTS: The model accurately predicted the biomass with 1.6% nRMSE at the segment scale from a k-fold cross-validation. It also gave satisfactory results when up-scaled to the branch level with a significantly lower error (13% nRMSE) and bias (-5%) compared to conventional cylinder fitting to the point cloud (nRMSE: 92%, bias: 82%), or using the pipe model theory (nRMSE: 31%, bias: -27%).The model was then applied to the whole-tree scale and showed that the sampled trees had more than 1.7km of structures on average and that 96% of that length was coming from the twigs (i.e. <5 cm diameter). Our results showed that neglecting twigs can lead to a significant underestimation of tree aboveground woody biomass (-21%). CONCLUSIONS: The structural model approach is an effective method that allows a more accurate estimation of the volumes of smaller branches from lidar point clouds. This method is versatile but requires manual measurements on branches for calibration. Nevertheless, once the model is calibrated, it can provide unbiased and large-scale estimations of tree structure volumes, making it an excellent choice for accurate 3D reconstruction of trees and estimating standing biomass.

An extended multiplicative error model of allometry: Incorporating systematic components, non-normal distributions, and piecewise heteroscedasticity.

Allometry refers to the relationship between the size of a trait and that of the whole body of an organism. Pioneering observations by Otto Snell and further elucidation by D'Arcy Thompson set the stage for its integration into Huxley's explanation of constant relative growth that epitomizes through the formula of simple allometry. The traditional method to identify such a model conforms to a regression protocol fitted in the direct scales of data. It involves Huxley's formula-systematic part and a lognormally distributed multiplicative error term. In many instances of allometric examination, the predictive strength of this paradigm is unsuitable. Established approaches to improve fit enhance the complexity of the systematic relationship while keeping the go-along normality-borne error. These extensions followed Huxley's idea that considering a biphasic allometric pattern could be necessary. However, for present data composing 10 410 pairs of measurements of individual eelgrass leaf dry weight and area, a fit relying on a biphasic systematic term and multiplicative lognormal errors barely improved correspondence measure values while maintaining a heavy tails problem. Moreover, the biphasic form and multiplicative-lognormal-mixture errors did not provide complete fit dependability either. However, updating the outline of such an error term to allow heteroscedasticity to occur in a piecewise-like mode finally produced overall fit consistency. Our results demonstrate that when attempting to achieve fit quality improvement in a Huxley's model-based multiplicative error scheme, allowing for a complex allometry form for the systematic part, a non-normal distribution-driven error term and a composite of uneven patterns to describe the heteroscedastic outline could be essential.