RESUMEN
Hair is a natural polymeric composite primarily composed of tight macrobundles of keratin proteins, which are highly responsive to external stimuli, similarly to the hydrogels and other natural fibrous gel systems like collagen and fibrin.Hair and its appearance play a significant role in human society. As a highly complex biocomposite system, it has been traditionally challenging to characterize and thus develop personal care products. Over the last few decades, a significant societal paradigm shift occurred among those with curly hair, accepting the natural morphological shape of their curls and styling their hair according to its innate, distinct, and unique material properties, which has given rise to the development of new hair classification systems, beyond the traditional and highly limited race-based distinction (Caucasian, Mongolian, and African). L'Oréal developed a hair typing taxonomy based on quantitative geometric parameters among the four key patternsâstraight, wavy, curly, and kinky, but it fails to capture the complex diversity of curly and kinky hair. Acclaimed celebrity hair stylist Andre Walker developed a classification system that is the existing gold standard for classifying curly and kinky hair, but it relies upon qualitative classification measures, making the system vague and ambiguous of phenotypic differences. The goal of this research is to use quantitative methods to identify new geometric parameters more representative of curly and kinky hair curl patterns, therefore providing more information on the kinds of personal care products that will resonate best with them and thus maximize desired appearance and health, and to correlate these new parameters with its mechanical properties. This was accomplished by identifying new geometric and mechanical parameters from several types of human hair samples.Geometric properties were measured using scanning electron microscopy (SEM), photogrammetry, and optical microscopy. Mechanical properties were measured under tensile extension using a texture analyzer (TA) and a dynamic mechanical analyzer (DMA), which bears similarity to the common act of brushing or combing. Both instruments measure force as a function of applied displacement, thus allowing the relationship between stress and applied stretch ratio to be measured as a hair strand uncurls and stretches to the point of fracture. From the resulting data, correlations were made between fiber geometry and mechanical performance. This data will be used to draw more conclusions on the contribution that fiber morphology has on hair fiber mechanics and will promote cultural inclusion among researchers and consumers possessing curly and kinky hair.
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Cabello , Humanos , Cabello/anatomía & histología , Fenotipo , Microscopía Electrónica de RastreoRESUMEN
Body condition is an important proxy for the overall health and energetic status of fishes. The classically used Fulton's condition factor requires length and mass measurements, but mass can be difficult to obtain in large species. Girth measurements can replace mass for wild pelagic sharks. However, girth-calculated condition has not been validated against Fulton's condition factor intraspecifically, across ontogeny or reproduction, or in a controlled setting. We used the epaulette shark (Hemiscyllium ocellatum), because they are amenable to captive reproduction, to track fine-scale body condition changes across life stages, oviparous reproduction and between condition indices. We measured four girths, total length and mass of 16 captive epaulette sharks across 1 year and tracked female reproduction daily. We also collected length and mass data from an additional 72 wild-caught sharks and 155 sharks from five previous studies and two public aquaria to examine the relationship between length and mass for this species. Even though data were derived from a variety of sources, a predictable length-mass relationship (R2 = 0.990) was achievable, indicating that combining data from a variety of sources could help overcome knowledge gaps regarding basic life history characteristics. We also found that condition factor decreased during early life stages, then increased again into adulthood, with predictable changes across the female reproductive cycle. Finally, we determined that both Fulton's and girth condition analyses were comparable. Outcomes from this study uniquely provide body condition changes across the complete life history, including fine-scale female reproductive stages, and validate the use of girths as a nonlethal whole-organism energetic assessment for fishes.
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Rasgos de la Historia de Vida , Tiburones , Femenino , Animales , ReproducciónRESUMEN
Evolutionary reversals, including re-evolution of lost structures, are commonly found in phylogenetic studies. However, we lack an understanding of how these reversals happen mechanistically. A snake-like body form has evolved many times in vertebrates, and occasionally a quadrupedal form has re-evolved, including in Brachymeles lizards. We use body form and locomotion data for species ranging from snake-like to quadrupedal to address how a quadrupedal form could re-evolve. We show that large, quadrupedal species are faster at burying and surface locomotion than snake-like species, indicating a lack of expected performance trade-off between these modes of locomotion. Species with limbs use them while burying, suggesting that limbs are useful for burying in wet, packed substrates. Palaeoclimatological data suggest that Brachymeles originally evolved a snake-like form under a drier climate probably with looser soil in which it was easier to dig. The quadrupedal clade evolved as the climate became humid, where limbs and large size facilitated fossorial locomotion in packed soils.
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Evolución Biológica , Clima , Lagartos/anatomía & histología , Animales , Locomoción , FilogeniaRESUMEN
This study describes and illustrates the jaws, teeth, and tooth microstructure of the Prickly Dogfish Oxynotus bruniensis. Detailed accounts of the dental morphology of O. bruniensis are rare and have not addressed the tissue arrangement or microstructure of the teeth. These features are documented and discussed in the contexts of interspecific comparisons with other elasmobranchs and the dietary specialization of O. bruniensis. The overall tooth morphology of O. bruniensis is similar to those of other closely related members in the order Squaliformes, as is the tissue arrangement, or histotype. Oxynotus bruniensis exhibits a simplified enameloid microstructure, which we compare with previously documented enameloid microstructures of other elasmobranchs. Though subtle interspecific differences in dental characters are documented, neither overall tooth morphology nor histotype and microstructure are unique to O. bruniensis. We conclude that in the case of O. bruniensis, dietary specialization is facilitated by behavioral rather than morphological specialization.
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Cazón/anatomía & histología , Diente/anatomía & histología , Animales , Dieta , Femenino , Maxilares/anatomía & histología , Maxilares/diagnóstico por imagen , Masculino , Diente/diagnóstico por imagen , Microtomografía por Rayos XRESUMEN
The skin surface structure of squamate reptiles varies greatly among species, likely because it plays a key role in a range of tasks, such as camouflage, locomotion, self-cleaning, mitigation of water loss and protection from physical damage. Although we have foundational knowledge about squamate skin morphology, we still know remarkably little about how intraspecific variation in skin surface structure translates to functional variation. This gap in our understanding can be in part traced back to: (i) our lack of knowledge on how body size determines skin surface structure; and (ii) the lack of means to perform high-throughput and detailed analysis of the three-dimensional (3D) anatomy of reptilian skin surfaces in a non-destructive manner. To fill this gap, we explored the possibilities of a new imaging technique, termed gel-based stereo-profilometry, to visualize and quantify the 3D topography of reptilian skin surface structure. Using this novel approach, we investigated intra-specific and intra-individual variation in the skin surface morphology of a focal lizard species, Anolis cristatellus. We assessed how various characteristics of surface topography (roughness, skew and kurtosis) and scale morphology (area, height, width and shape) scale with body size across different body regions. Based on an ontogenetic series of A. cristatellus males, we show that skin roughness increases with body size. Skin patches on the ventral body region of lizards were rougher than on the dorsum, but this was a consequence of ventral scales being larger than dorsal scales. Dorsal surface skew and kurtosis varied with body size, but surfaces on the ventral skin showed no such relationship. Scale size scaled isometrically with body size, and while ventral scales differed in shape from dorsal scales, scale shape did not change with ontogeny. Overall, this study demonstrates that gel-based stereo-profilometry is a promising method to rapidly assess the 3D surface structure of reptilian skin at the microscopic level. Additionally, our findings of the explanatory power of body size on skin surface diversity provide a foundation for future studies to disentangle the relationships among morphological, functional and ecological diversity in squamate reptile skin surfaces.
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Escamas de Animales/anatomía & histología , Biometría/métodos , Lagartos/anatomía & histología , Escamas de Animales/crecimiento & desarrollo , Animales , Tamaño Corporal , Imagenología Tridimensional , Lagartos/crecimiento & desarrollo , MasculinoRESUMEN
The ability of an animal to run fast has important consequences on its survival capacity and overall fitness. Previous studies have documented how variation in the morphology of the limbs is related to variation in locomotor performance. Although these studies have suggested direct relations between sprint speed and hindlimb morphology, few quantitative data exist. Consequently, it remains unclear whether selection acts in limb segment lengths, overall muscle mass or muscle architecture (e.g. muscle fiber length and cross-sectional area). Here, we investigate whether muscle architecture (mass, fiber length and physiological cross-sectional area), hindlimb segment dimensions, or both, explain variation in sprint speed across 14 species of Anolis lizards. Moreover, we test whether similar relationships exist between morphology and performance for both sexes, which may not be the case given the known differences in locomotor behavior and habitat use. Our results show that the main driver of sprint speed is the variation in femur length for both males and females. Our results further show sexual dimorphism in the traits studied and, moreover, show differences in the traits that predict maximal sprint speed in males and females. For example, snout vent length and overall muscle mass are also good predictors of sprint speed in males, whereas no relationships between muscle mass and sprint speed was observed in females. Only a few significant relationships were found between muscle architecture (fiber length, cross-sectional area) and sprint speed in male anoles, suggesting that overall muscles size, rather than muscle architecture, appears to be under selection.
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Miembro Posterior/anatomía & histología , Lagartos/anatomía & histología , Lagartos/fisiología , Locomoción , Animales , Femenino , Masculino , Factores Sexuales , Especificidad de la EspecieRESUMEN
Organismal functions are size-dependent whenever body surfaces supply body volumes. Larger organisms can develop strongly folded internal surfaces for enhanced diffusion, but in many cases areas cannot be folded so that their enlargement is constrained by anatomy, presenting a problem for larger animals. Here, we study the allometry of adhesive pad area in 225 climbing animal species, covering more than seven orders of magnitude in weight. Across all taxa, adhesive pad area showed extreme positive allometry and scaled with weight, implying a 200-fold increase of relative pad area from mites to geckos. However, allometric scaling coefficients for pad area systematically decreased with taxonomic level and were close to isometry when evolutionary history was accounted for, indicating that the substantial anatomical changes required to achieve this increase in relative pad area are limited by phylogenetic constraints. Using a comparative phylogenetic approach, we found that the departure from isometry is almost exclusively caused by large differences in size-corrected pad area between arthropods and vertebrates. To mitigate the expected decrease of weight-specific adhesion within closely related taxa where pad area scaled close to isometry, data for several taxa suggest that the pads' adhesive strength increased for larger animals. The combination of adjustments in relative pad area for distantly related taxa and changes in adhesive strength for closely related groups helps explain how climbing with adhesive pads has evolved in animals varying over seven orders of magnitude in body weight. Our results illustrate the size limits of adhesion-based climbing, with profound implications for large-scale bio-inspired adhesives.
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Adhesividad , Movimiento , AnimalesRESUMEN
This study examines the feeding behaviour and kinematics of three sub-adult sand tiger sharks Carcharias taurus on display at Mystic Aquarium (Mystic, Connecticut, USA). Using high-speed video data from 52 bites, we identify kinematic variables associated with the expansive and compressive phases of the bite. The mean bite duration from the onset of the expansive phase to the conclusion of the compressive phase is mean (± SE) 0.14 ± 0.01 s and across the 10 fastest bites of each individual, the maximum performance average is 0.13 ± 0.01 s. Values of maximum performance do not vary significantly among individuals. When compared with kinematic bite data from species studied previously, these results indicate that body size is not the only determinant factor of bite duration. This study also provides detailed descriptions of feeding behaviours in C. taurus and presents documentation of tooth loss both prior to and during feeding, suggesting that there are multiple mechanisms of tooth loss and use in C. taurus. Finally, we discuss the behavioural and ecological components of prey capture in C. taurus and suggest points of consideration to facilitate interspecific comparisons of prey capture performance in ram-feeding, macrophagous elasmobranchs.
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Conducta Alimentaria , Tiburones/fisiología , Animales , Fenómenos BiomecánicosRESUMEN
Booralana nickorum sp. nov. is described from the deep-water slope of the Exuma Sound, The Bahamas, from depths of 540 to 560 metres. It is the fourth species to be assigned to the genus and the second species described from the Western North Atlantic. The species can be distinguished from Booralana tricarinata Camp and Heard, 1988 and the other species by the sub-triangular pleotelson and the uropodal exopod of mature males being far longer than endopod, with both rami extending well beyond the posterior margin of the pleotelson. Additionally, pleopods 3 and 4 lack a prominent angle at midpoint of ramus.
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Isópodos , Masculino , Animales , Bahamas , CrustáceosRESUMEN
Extant salamanders are used as modern analogs of early digit-bearing tetrapods due to general similarities in morphology and ecology but the study species have been primarily terrestrial and relatively small when the earliest digit-bearing tetrapods were aquatic and an order of magnitude larger. Thus, we created a 3D computational model of underwater walking in extant Japanese giant salamanders (Andrias japonicus) using 3D photogrammetry and open-access graphics software (Blender) to broaden the range of testable hypotheses about the incipient stages of terrestrial locomotion. Our 3D model and software protocol represent the initial stages of an open-access pipeline that could serve as a "one-stop-shop" for studying locomotor function, from creating 3D models to analyzing the mechanics of locomotor gaits. While other pipelines generally require multiple software programs to accomplish the different steps in creating and analyzing computational models of locomotion, our protocol is built entirely within Blender and fully customizable with its Python scripting so users can devote more time to creating and analyzing models instead of navigating the learning curves of several software programs. The main value of our approach is that key kinematic variables (e.g., speed, stride length, elbow flexion) can be easily altered on the 3D model, allowing scientists to test hypotheses about locomotor function and conduct manipulative experiments (e.g., lengthening bones) that are difficult to perform in vivo. The accurate 3D meshes (and animations) generated through photogrammetry also provide exciting opportunities to expand the abundance and diversity of 3D digital animals available for researchers, educators, artists, conservation biologists, etc. to maximize societal impacts.
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Many clades contain ecologically and phenotypically similar species across continents, yet the processes generating this similarity are largely unstudied, leaving fundamental questions unanswered. Is similarity in morphology and performance across assemblages caused by evolutionary convergence or by biogeographic dispersal of evolutionarily conserved ecotypes? Does convergence to new ecological conditions erase evidence of past adaptation? Here, we analyse ecology, morphology and performance in frog assemblages from three continents (Asia, Australia and South America), assessing the importance of dispersal and convergent evolution in explaining similarity across regions. We find three striking results. First, species using the same microhabitat type are highly similar in morphology and performance across both clades and continents. Second, some species on different continents owe their similarity to dispersal and evolutionary conservatism (rather than evolutionary convergence), even over vast temporal and spatial scales. Third, in one case, an ecologically specialized ancestor radiated into diverse ecotypes that have converged with those on other continents, largely erasing traces of past adaptation to their ancestral ecology. Overall, our study highlights the roles of both evolutionary conservatism and convergence in explaining similarity in species traits over large spatial and temporal scales and demonstrates a statistical framework for addressing these questions in other systems.
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Anuros/fisiología , Evolución Biológica , Adaptación Fisiológica , Animales , Asia , Australia , Ecosistema , Filogenia , América del SurRESUMEN
As anthropogenic activities are increasing the frequency and severity of droughts, understanding whether and how fast populations can adapt to sudden changes in their hydric environment is critically important. Here, we capitalize on the introduction of the Cuban brown anole lizard (Anolis sagrei) in North America to assess the contemporary evolution of a widespread terrestrial vertebrate to an abrupt climatic niche shift. We characterized hydric balance in 30 populations along a large climatic gradient. We found that while evaporative and cutaneous water loss varied widely, there was no climatic cline, as would be expected under adaptation. Furthermore, the skin of lizards from more arid environments was covered with smaller scales, a condition thought to limit water conservation and thus be maladaptive. In contrast to environmental conditions, genome-averaged ancestry was a significant predictor of water loss. This was reinforced by our genome-wide association analyses, which indicated a significant ancestry-specific effect for water loss at one locus. Thus, our study indicates that the water balance of invasive brown anoles is dictated by an environment-independent introduction and hybridization history and highlights genetic interactions or genetic correlations as factors that might forestall adaptation. Alternative water conservation strategies, including behavioral mitigation, may influence the brown anole invasion success and require future examination.
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Lagartos , Animales , Lagartos/genética , Estudio de Asociación del Genoma Completo , Aclimatación , Adaptación Fisiológica , AguaRESUMEN
Jumping is a common form of locomotion for many arboreal animals. Many species of the arboreal lizard genus Anolis occupy habitats in which they must jump to and from unsteady perches, e.g. narrow branches, vines, grass and leaves. Anoles therefore often use compliant perches that could alter jump performance. In this study we conducted a small survey of the compliance of perches used by the arboreal green anole Anolis carolinensis in the wild (N=54 perches) and then, using perches within the range of compliances used by this species, investigated how perch compliance (flexibility) affects the key jumping variables jump distance, takeoff duration, takeoff angle, takeoff speed and landing angle in A. carolinensis in the laboratory (N=11). We observed that lizards lost contact with compliant horizontal perches prior to perch recoil, and increased perch compliance resulted in decreased jump distance and takeoff speed, likely because of the loss of kinetic energy to the flexion of the perch. However, the most striking effect of perch compliance was an unexpected one; perch recoil following takeoff resulted in the lizards being struck on the tail by the perch, even on the narrowest perches. This interaction between the perch and the tail significantly altered body positioning during flight and landing. These results suggest that although the use of compliant perches in the wild is common for this species, jumping from these perches is potentially costly and may affect survival and behavior, particularly in the largest individuals.
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Marcha , Lagartos/fisiología , Locomoción , Animales , Fenómenos Biomecánicos , Ambiente , Femenino , Florida , Masculino , Propiedades de Superficie , Grabación en VideoRESUMEN
The study of biological form is a vital goal of evolutionary biology and functional morphology. We review an emerging set of methods that allow scientists to create and study accurate 3D models of living organisms and animate those models for biomechanical and fluid dynamic analyses. The methods for creating such models include 3D photogrammetry, laser and CT scanning, and 3D software. New multi-camera devices can be used to create accurate 3D models of living animals in the wild and captivity. New websites and virtual reality/augmented reality devices now enable the visualization and sharing of these data. We provide examples of these approaches for animals ranging from large whales to lizards and show applications for several areas: Natural history collections; body condition/scaling, bioinspired robotics, computational fluids dynamics (CFD), machine learning, and education. We provide two datasets to demonstrate the efficacy of CFD and machine learning approaches and conclude with a prospectus.
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Locomotor performance is a crucial determinant of organismal fitness but is often impaired in certain circumstances, such as increased mass (loading) resulting from feeding or gravidity. Although the effects of loading have been studied extensively for striding locomotion, its effects on jumping are poorly understood. Jumping is a mode of locomotion that is widely used across animal taxa. It demands large amounts of power over a short time interval and, consequently, may be affected by loading to a greater extent than other modes of locomotion. We placed artificial loads equal to 30% body mass on individuals of the species Anolis carolinensis to simulate the mass gain following the consumption of a large meal. We investigated the effects of loading on jump performance (maximum jump distance and accuracy), kinematics and power output. Loading caused a significant 18% decline in maximum jump distance and a significant 10% decline in takeoff speed. In other words, the presence of the load caused the lizards to take shorter and slower jumps, whereas takeoff angle and takeoff duration were not affected. By contrast, jump accuracy was unaffected by loading, although accuracy declined when lizards jumped to farther perches. Finally, mass-specific power output did not increase significantly when lizards jumped with loads, suggesting that the ability to produce mechanical power may be a key limiting factor for maximum jump performance. Our results suggest that mass gain after a large meal can pose a significant locomotor challenge and also imply a tradeoff between fulfilling energy requirement and moving efficiently in the environment.
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Marcha , Lagartos/fisiología , Locomoción , Animales , Fenómenos Biomecánicos , Femenino , Masculino , Grabación en VideoRESUMEN
Although rapid adaptive changes in morphology on ecological time scales are now well documented in natural populations, the effects of such changes on whole-organism performance capacity and the consequences on ecological dynamics at the population level are often unclear. Here we show how lizards have rapidly evolved differences in head morphology, bite strength, and digestive tract structure after experimental introduction into a novel environment. Despite the short time scale ( approximately 36 years) since this introduction, these changes in morphology and performance parallel those typically documented among species and even families of lizards in both the type and extent of their specialization. Moreover, these changes have occurred side-by-side with dramatic changes in population density and social structure, providing a compelling example of how the invasion of a novel habitat can evolutionarily drive multiple aspects of the phenotype.
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Biodiversidad , Evolución Biológica , Dieta , Lagartos/fisiología , Animales , Fuerza de la Mordida , Ciego/fisiología , Ambiente , Femenino , Cabeza/anatomía & histología , Masculino , Datos de Secuencia Molecular , Fenotipo , Dinámica Poblacional , Conducta Predatoria , Estaciones del Año , Factores de TiempoRESUMEN
Physical principles and laws determine the set of possible organismal phenotypes. Constraints arising from development, the environment, and evolutionary history then yield workable, integrated phenotypes. We propose a theoretical and practical framework that considers the role of changing environments. This 'ecomechanical approach' integrates functional organismal traits with the ecological variables. This approach informs our ability to predict species shifts in survival and distribution and provides critical insights into phenotypic diversity. We outline how to use the ecomechanical paradigm using drag-induced bending in trees as an example. Our approach can be incorporated into existing research and help build interdisciplinary bridges. Finally, we identify key factors needed for mass data collection, analysis, and the dissemination of models relevant to this framework.
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Evolución Biológica , Ecosistema , Fenotipo , ÁrbolesRESUMEN
Three-dimensional (3D) modeling techniques have been increasingly utilized across disciplines for the visualization and analysis of complex structures. We employ 3D-digital photogrammetry for understanding the scaling of the body axis of 12 species of scincid lizards in the genus Brachymeles. These skinks represent a diverse radiation which shows tremendous variation in body size and degree of axial elongation. Because of the complex nature of the body axis, 3D-methods are important for understanding how the body axis evolves. 3D-digital photogrammetry presents a flexible, inexpensive, and portable system for the reconstruction of biological forms. As body size increased among species, the cross-sectional area and circumference of the head and other portions of the body axis increased isometrically, which indicates that species of differing sizes possess proportionally similar head and body shapes. These results suggest that there are no substantial head and body shape changes with body size among the sampled species, but further comparative studies with larger sample sizes and functional studies of size and morphology effects on burrowing or above-ground locomotion are needed.
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Tipificación del Cuerpo , Tamaño Corporal , Imagenología Tridimensional , Lagartos/anatomía & histología , Fotogrametría , Animales , Análisis de los Mínimos Cuadrados , Filogenia , Análisis de RegresiónRESUMEN
Geckos are a highly diverse group of lizards, with more than 1,700 species that exhibit a wide range of behaviors, ecologies, and sizes. However, no study has examined links between habitat use and body shape in pad-bearing geckos. We set out to answer a basic question using a data set of pad-bearing geckos (112 species, 103 pad-bearing, 9 padless, 42 genera): Do geckos that occur in different habitats also differ in body shape? Overall, we found that body shape was surprisingly similar among our sample of pad-bearing species, with the exception of the genus Uroplatus, which was clearly distinct from other geckos due to its depressed body and long limbs. However, the padless geckos differed in body shape from the pad-bearing geckos by having longer arms and legs and less rotund bodies. We found that about half of the pad-bearing species primarily inhabit trees, with the other half, divided approximately equally among rocks, the ground, and mixed habitats. We found no significant links between habitat use and body shape, nor any propensity for larger species to occupy different habitats than smaller species. Padless species tend to inhabit rock and ground substrates. Our results indicate that pad-bearing geckos have a relatively uniform body form, which contrasts with to their diversity in color, size, and behavior. Indeed, our data show that the general gecko body shape is suitable for a wide range of habitats, ranging from arboreal to terrestrial. This pattern is a departure from other ecomorphological studies and suggests that geckos may not easily fit into the mold of adaptive radiation, as suggested by prior studies.
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Ecosistema , Lagartos/anatomía & histología , Animales , Tamaño Corporal , Filogenia , Análisis de Componente Principal , Especificidad de la EspecieRESUMEN
Sexual selection can lead to rapid evolution of sexual traits and striking morphological diversity across taxa. In populations where competition for mates is intense, males sometimes evolve distinct behavioral strategies along with morphological differences that help them secure mating opportunities. Strong postcopulatory selection and differential resource allocation across male strategy type can result in strategy-specific differences in sexual traits, such as sperm morphology, ejaculate components, and testis size. Some polymorphic species also have strategy-specific genital morphology. Thus far, among vertebrates, this has only been observed in fish. Here, we present the first morphological description of the intromittant copulatory organ, the hemipenis, of the three mating types of the side-blotched lizard, Uta stansburiana, from a population that exhibits alternative mating strategies. We found that the isometrically scaling hemipenis was shortest in the nonterritorial (yellow) morph that sneaks copulations with other males' mates. Although the hemipenes were generally the same shape across morphs, the usurping territorial (orange) morph had a significantly wider apical horn than the nonterritorial sneaker morph. Sneaker males also had smaller relative body masses than both the mate-guarding (blue) morph and the usurper morph, and shorter tibia than the usurper morph. This study using a small sample of males suggests that strong sexual selection may drive genital trait differentiation across morphs within populations of terrestrial vertebrates.