What Amphibians Can Teach Us About the Evolution of Parental Care.

Parenting is considered a key evolutionary innovation that contributed to the diversification and expansion of vertebrates. However, we know little about how such diversity evolved. Amphibians are an ideal group in which to identify the ecological factors that have facilitated or constrained the evolution of different forms of parental care. Among, but also within, the three amphibian orders-Anura, Caudata, and Gymnophiona-there is a high level of variation in habitat use, fertilization mode, mating systems, and parental sex roles. Recent work using broad phylogenetic, experimental, and physiological approaches has helped to uncover the factors that have selected for the evolution of care and transitions between different forms of parenting. Here, we highlight the exceptional diversity of amphibian parental care, emphasize the unique opportunities this group offers for addressing key questions about the evolution of parenting, and give insights into promising novel directions of research.

Anatomical insights into fish terrestrial locomotion: A study of barred mudskipper (Periophthalmus argentilineatus) fins based on µCT 3D reconstructions.

Mudskippers are a group of extant ray-finned fishes with an amphibious lifestyle and serve as exemplars for understanding the evolution of amphibious capabilities in teleosts. A comprehensive anatomical profile of both the soft and hard tissues within their propulsive fins is essential for advancing our understanding of terrestrial locomotor adaptations in fish. Despite the ecological significance of mudskippers, detailed data on their musculoskeletal anatomy remains limited. In the present research, we utilized contrast-enhanced high-resolution microcomputed tomography (µCT) imaging to investigate the barred mudskipper, Periophthalmus argentilineatus. This technique enabled detailed reconstruction and quantification of the morphological details of the pectoral, pelvic, and caudal fins of this terrestrial mudskipper, facilitating comparison with its aquatic relatives. Our findings reveal that P. argentilineatus has undergone complex musculoskeletal adaptations for terrestrial movement, including an increase in muscle complexity and muscle volume, as well as the development of specialized structures like aponeuroses for pectoral fin extension. Skeletal modifications are also evident, with features such as a reinforced shoulder-pelvic joint and thickened fin rays. These evolutionary modifications suggest biomechanically advanced fins capable of overcoming the gravitational challenges of terrestrial habitats, indicating a strong selective advantage for these features in land-based environments. The unique musculoskeletal modifications in the fins of mudskippers like P. argentilineatus, compared with their aquatic counterparts, mark a critical evolutionary shift toward terrestrial adaptations. This study not only sheds light on the specific anatomical changes facilitating this transition but also offers broader insights into the early evolutionary mechanisms of terrestrial locomotion, potentially mirroring the transformative journey from aquatic to terrestrial life in the lineage leading to tetrapods.

Terrestriality across the primate order: A review and analysis of ground use in primates.

Terrestriality is relatively rare in the predominantly arboreal primate order. How frequently, and when, terrestriality appears in primate evolution, and the factors that influence this behavior, are not well understood. To investigate this, we compiled data describing terrestriality in 515 extant nonhuman primate taxa. We describe the geographic and phylogenetic distribution of terrestriality, including an ancestral state reconstruction estimating the frequency and timing of evolutionary transitions to terrestriality. We review hypotheses concerning the evolution of primate terrestriality and test these using data we collected pertaining to characteristics including body mass and diet, and ecological factors including forest structure, food availability, weather, and predation pressure. Using Bayesian analyses, we find body mass and normalized difference vegetation index are the most reliable predictors of terrestriality. When considering subsets of taxa, we find ecological factors such as forest height and rainfall, and not body mass, are the most reliable predictors of terrestriality for platyrrhines and lemurs.

Male-biased stone tool use by wild white-faced capuchins (Cebus capucinus imitator).

Tool-using primates often show sex differences in both the frequency and efficiency of tool use. In species with sex-biased dispersal, such within-group variation likely shapes patterns of cultural transmission of tool-use traditions between groups. On the Panamanian islands of Jicarón and Coiba, a population of white-faced capuchins (Cebus capucinus imitator)-some of which engage in habitual stone tool use-provide an opportunity to test hypotheses about why such sex-biases arise. On Jicarón, we have only observed males engaging in stone tool use, whereas on Coiba, both sexes are known to use tools. Using 5 years of camera trap data, we provide evidence that this variation likely reflects a sex difference in tool use rather than a sampling artifact, and then test hypotheses about the factors driving this pattern. Differences in physical ability or risk-aversion, and competition over access to anvils do not account for the sex-differences in tool-use we observe. Our data show that adult females are physically capable of stone tool use: adult females on Coiba and juveniles on Jicarón smaller than adult females regularly engage in tool use. Females also have ample opportunity to use tools: the sexes are equally terrestrial, and competition over anvils is low. Finally, females rarely scrounge on left-over food items either during or after tool-using events, suggesting they are not being provisioned by males. Although it remains unclear why adult white-faced capuchin females on Jicarón do not use stone-tools, our results illustrate that such sex biases in socially learned behaviors can arise even in the absence of obvious physical, environmental, and social constraints. This suggests that a much more nuanced understanding of the differences in social structure, diet, and dispersal patterns are needed to explain why sex-biases in tool use arise in some populations but not in others.

Dental topographic proxies for ecological characteristics in carnivoran mammals.

Form-function relationships in mammalian feeding systems are active topics of research in evolutionary biology. This is due principally to their fundamental importance for understanding dietary adaptations in extinct taxa and macro-evolutionary patterns of morphological transformations through changing environments. We hypothesize that three-dimensional dental topographic metrics represent stronger predictors for dietary and other ecological variables than do linear measurements. To test this hypothesis, we measured three dental topographic metrics: Relief Index (RFI), Dirichlet Normal Energy (DNE), and Orientation Patch Count Rotated (OPCR) in 57 extant carnivoran species. Premolar and molar dental topographic indices were regressed against activity, diet breadth, habitat breadth, terrestriality, and trophic level variables within a phylogenetic framework. The results of this study showed significant correlations between RFI and the ecological variables diet breadth and trophic level. Weaker correlations are documented between OPCR and activity and between DNE and trophic level. Our results suggest that cusp height is strongly reflective of dietary ecology in carnivorans as a whole, and represents a proxy mainly for different degrees of hypercarnivory observed within this group of predatory mammals.

Calcaneal shape variation in humans, nonhuman primates, and early hominins.

The foot has played a prominent role in evaluating early hominin locomotion. The calcaneus, in particular, plays an important role in weight-bearing. Although the calcanei of early hominins have been previously scrutinized, a three-dimensional analysis of the entire calcaneal shape has not been conducted. Here, we investigate the relationship between external calcaneal shape and locomotion in modern Homo sapiens (n = 130), Gorilla (n = 86), Pan (n = 112), Pongo (n = 31), Papio (n = 28), and hylobatids (Hylobates, Symphalangus; n = 32). We use these results to place the calcanei attributed to Australopithecus sediba, A. africanus, A. afarensis, H. naledi, and Homo habilis/Paranthropus boisei into a locomotor context. Calcanei were scanned using either surface scanning or micro-CT and their external shape analyzed using a three-dimensional geometric morphometric sliding semilandmark analysis. Blomberg's K statistic was used to estimate phylogenetic signal in the shape data. Shape variation was summarized using a principal components analysis. Procrustes distances between all taxa as well as distances between each fossil and the average of each taxon were calculated. Blomberg's K statistic was small (K = 0.1651), indicating weak phylogenetic effects, suggesting variation is driven by factors other than phylogeny (e.g., locomotion or body size). Modern humans have a large calcaneus relative to body size and display a uniquely convex cuboid facet, facilitating a rigid midfoot for bipedalism. More arboreal great apes display relatively deeper cuboid facet pivot regions for increased midfoot mobility. Australopithecus afarensis demonstrates the most human-like calcaneus, consistent with obligate bipedalism. Homo naledi is primarily modern human-like, but with some intermediate traits, suggesting a different form of bipedalism than modern humans. Australopithecus africanus, A. sediba, and H. habilis/P. boisei calcanei all possess unique combinations of human and nonhuman ape-like morphologies, suggesting a combination of bipedal and arboreal behaviors.

Gorilla calcaneal morphological variation and ecological divergence.

OBJECTIVES: The primate foot has been extensively investigated because of its role in weight-bearing; however, the calcaneus has been relatively understudied. Here we examine entire gorilla calcaneal external shape to understand its relationship with locomotor behavior. MATERIALS AND METHODS: Calcanei of Gorilla gorilla gorilla (n = 43), Gorilla beringei graueri (n = 20), and Gorilla beringei beringei (n = 15) were surface or micro-CT scanned. External shape was analyzed through a three-dimensional geometric morphometric sliding semilandmark analysis. Semilandmarks were slid relative to an updated Procrustes average in order to minimize the bending energy of the thin plate spline interpolation function. Shape variation was summarized using principal components analysis of shape coordinates. Procrustes distances between taxa averages were calculated and resampling statistics run to test pairwise differences. Linear measures were collected and regressed against estimated body mass. RESULTS: All three taxa exhibit statistically different morphologies (p < .001 for pairwise comparisons). G. g. gorilla demonstrates an anteroposteriorly elongated calcaneus with a deeper cuboid pivot region and mediolaterally flatter posterior talar facet. G. b. beringei possesses the flattest cuboid and most medially-angled posterior talar facets. G. b. graueri demonstrates intermediate articular facet morphology, a medially-angled tuberosity, and an elongated peroneal trochlea. DISCUSSION: Articular facet differences separate gorillas along a locomotor gradient. G. g. gorilla is adapted for arboreality with greater joint mobility, while G. b. beringei is adapted for more stereotypical loads associated with terrestriality. G. b. graueri's unique posterolateral morphology may be due to a secondary transition to greater arboreality from a more terrestrial ancestor.

Gliding Dragons and Flying Squirrels: Diversifying versus Stabilizing Selection on Morphology following the Evolution of an Innovation.

Evolutionary innovations and ecological competition are factors often cited as drivers of adaptive diversification. Yet many innovations result in stabilizing rather than diversifying selection on morphology, and morphological disparity among coexisting species can reflect competitive exclusion (species sorting) rather than sympatric adaptive divergence (character displacement). We studied the innovation of gliding in dragons (Agamidae) and squirrels (Sciuridae) and its effect on subsequent body size diversification. We found that gliding either had no impact (squirrels) or resulted in strong stabilizing selection on body size (dragons). Despite this constraining effect in dragons, sympatric gliders exhibit greater size disparity compared with allopatric gliders, a pattern consistent with, although not exclusively explained by, ecological competition changing the adaptive landscape of body size evolution to induce character displacement. These results show that innovations do not necessarily instigate further differentiation among species, as is so often assumed, and suggest that competition can be a powerful force generating morphological divergence among coexisting species, even in the face of strong stabilizing selection.

Massive increase in visual range preceded the origin of terrestrial vertebrates.

The evolution of terrestrial vertebrates, starting around 385 million years ago, is an iconic moment in evolution that brings to mind images of fish transforming into four-legged animals. Here, we show that this radical change in body shape was preceded by an equally dramatic change in sensory abilities akin to transitioning from seeing over short distances in a dense fog to seeing over long distances on a clear day. Measurements of eye sockets and simulations of their evolution show that eyes nearly tripled in size just before vertebrates began living on land. Computational simulations of these animal's visual ecology show that for viewing objects through water, the increase in eye size provided a negligible increase in performance. However, when viewing objects through air, the increase in eye size provided a large increase in performance. The jump in eye size was, therefore, unlikely to have arisen for seeing through water and instead points to an unexpected hybrid of seeing through air while still primarily inhabiting water. Our results and several anatomical innovations arising at the same time suggest lifestyle similarity to crocodiles. The consequent combination of the increase in eye size and vision through air would have conferred a 1 million-fold increase in the amount of space within which objects could be seen. The "buena vista" hypothesis that our data suggest is that seeing opportunities from afar played a role in the subsequent evolution of fully terrestrial limbs as well as the emergence of elaborated action sequences through planning circuits in the nervous system.

Positional behavior and substrate use in wild adult bearded capuchin monkeys (Sapajus libidinosus).

Natural selection for positional behavior (posture and locomotion) has at least partially driven the evolution of anatomical form and function in the order Primates. Examination of bipedal behaviors associated with daily activity patterns, foraging, and terrestrial habitat use in nonhuman primates, particularly those that adopt bipedal postures and use bipedal locomotion, allows us to refine hypotheses concerning the evolution of bipedalism in humans. This study describes the positional behavior of wild bearded capuchins (Sapajus libidinosus), a species that is known for its use of terrestrial substrates and its habitual use of stones as tools. Here, we test the association of terrestrial substrate use with bipedal posture and locomotion, and the influence of sex (which co-varies with body mass in adults of this species) on positional behavior and substrate use. Behavior and location of 16 wild adult bearded capuchins from two groups were sampled systematically at 15 s intervals for 2 min periods for 1 year (10,244 samples). Despite their different body masses, adult males (average 3.5 kg) and females (average 2.1 kg) in this study did not differ substantially in their positional behaviors, postures, or use of substrates for particular activities. The monkeys used terrestrial substrates in 27% of samples. Bipedal postures and behaviors, while not a prominent feature of their behavior, occurred in different forms on the two substrates. The monkeys crouched bipedally in trees, but did not use other bipedal postures in trees. While on terrestrial substrates, they also crouched bipedally but occasionally stood upright and moved bipedally with orthograde posture. Bearded capuchin monkeys' behavior supports the suggestion from anatomical analysis that S. libidinosus is morphologically better adapted than its congeners to adopt orthograde postures.

A Short Cut to Mushrooms - Red Langur (Presbytis rubicunda) Consumption of Terrestrial Fungus.

We report the first in-depth evidence of targeted mushroom foraging in an Asian colobine. Using direct observations (2010-2018) and camera traps (2008-2018) in the Sebangau Forest, Central Kalimantan, Indonesian Borneo, we show how adult female red langurs (Presbytis rubicunda) are regularly descending to the ground to consume mushrooms. We recorded 82 counts (0.36% of all focal observations, n = 25,502) of the focal adult langur on the ground from direct observations of habituated groups, and 80 independent images/videos of red langurs on the ground were obtained from the camera traps representing 1.12% of total images (n = 7,145). Mushroom consumption took place in 4 families, representing 0.04% of total focal behaviour observations and 24.3% of total time feeding on the ground. From the camera trap photos, red langurs are spending 20% of time on the ground feeding. We speculate that mushrooms could be a supplementary food for adult female langurs as there is an increase in consumption in April and November.

Locomotor Kinematics of Two Semi-Wild Macaque Species (Macaca assamensis and Macaca arctoides) in Thailand.

This project aimed to investigate primate locomotor kinematics noninvasively in the wild. Semi-wild Assamese and stump-tailed macaques were selected for the study, which was performed in Thailand. We investigated their locomotor kinematics and its relationship to habitat use. The macaques' positional behavior was recorded with two video cameras, and kinematic parameters were estimated during terrestrial quadrupedal locomotion, using the markerless method. The data analyzed so far revealed that stump-tailed macaques walk with longer, less frequent strides than Assamese macaques. Although stump-tailed macaques present a smaller angular excursion of the shoulder joint than Assamese macaques, they exhibited a relatively large shoulder girdle motion and anteroposterior translation of the shoulder, which increased their stride length. Additionally, stump-tailed macaques exhibited a digitigrade gait and elbow extension, suggesting a good adaptation to terrestrial locomotion. Assamese macaques, on the other hand, exhibited a gait that did not seem optimized for terrestrial locomotion, using the hands in a palmigrade posture and frequently flexing the fingers at varying degrees. The kinematic characteristics of the two species studied is consistent with previous field observations reporting that Assamese macaques are highly arboreal, whereas stump-tailed macaques are more terrestrial.

Flexibility in positional behavior, strata use, and substrate utilization among Bale monkeys (Chlorocebus djamdjamensis) in response to habitat fragmentation and degradation.

Studies of the effects of habitat fragmentation and degradation on primate positional behavior, strata use, and substrate utilization offer valuable insights into the behavioral and ecological flexibility of primates whose habitats have undergone extensive anthropogenic disturbance. In this study, we evaluated how positional behavior, strata use, and substrate utilization differed between Bale monkeys (Chlorocebus djamdjamensis)-bamboo-eating cercopithecids endemic to the southern Ethiopian Highlands-occupying continuous versus fragmented forests. Bale monkeys in forest fragments (where bamboo had been degraded or eradicated) spent significantly more time on the ground and in understory strata whereas those in continuous forest spent significantly more time in the middle and upper strata. Bale monkeys in forest fragments also spent significantly more time walking and galloping and significantly less time climbing than those in continuous forest. Our results suggest that, unlike the primarily arboreal Bale monkeys in continuous forest, Bale monkeys in forest fragments should be characterized as semi-terrestrial. In response to habitat disturbance in fragments, we observed a greater emphasis on terrestrial foraging and travel among Bale monkeys in these human altered habitats, which may put them at greater risk of predation and conflict with nearby human populations. Bale monkeys in fragments exhibit flexibility in their positional behavioral repertoire and their degree of terrestriality is more similar to their sister taxa in Chlorocebus than to Bale monkeys in continuous forest. These findings suggest that habitat alteration may compel Bale monkeys to exhibit semi-terrestrial behaviors crucial for their persistence in human-modified habitats. Our results contribute to a growing body of literature on primate behavioral responses to anthropogenic modification of their habitats and provide information that can contribute to the design of appropriate conservation management plans.

Terrestriality of Wild Sapajus cay (Illiger, 1815) as Revealed by Camera Traps.

Although primarily arboreal, the capuchin monkey (Sapajus cay) descends to the ground for several reasons. We used terrestrial records obtained by camera-trapping surveys to investigate seasonality in the terrestriality of capuchin monkeys, reasons to descend to the ground, and periods of the day when terrestriality was more pronounced. We carried out the study in the Urucum Massif, Brazilian Pantanal. We obtained data from 2 camera-trapping surveys carried out in the dry and rainy seasons. Terrestrial behaviours were categorized, and terrestrial activity patterns were described using a kernel density approach. We observed a seasonal effect on the frequency of terrestrial behaviours of capuchin monkeys, who used the ground more during the dry season. We identified 6 different types of terrestrial behaviour, but travelling (33.3%), foraging (23.3%), and drinking water (23.3%) were the most frequently observed. All records occurred during the day, with 2 peaks in terrestrial activity. Seasonal terrestriality was mainly linked to ecological needs during periods of scarce food and water.

Being fat and smart: A comparative analysis of the fat-brain trade-off in mammals.

Humans stand out among non-aquatic mammals by having both an extremely large brain and a relatively large amount of body fat. To understand the evolution of this human peculiarity we report a phylogenetic comparative study of 120 mammalian species, including 30 primates, using seasonal variation in adult body mass as a proxy of the tendency to store fat. Species that rely on storing fat to survive lean periods are expected to be less active because of higher costs of locomotion and have increased predation risk due to reduced agility. Because a fat-storage strategy reduces the net cognitive benefit of a large brain without reducing its cost, such species should be less likely to evolve a larger brain than non-fat-storing species. We therefore predict that the two strategies to buffer food shortages (storing body fat and cognitive flexibility) are compensatory, and therefore predict negative co-evolution between relative brain size and seasonal variation in body mass. This trade-off is expected to be stronger in predominantly arboreal species than in more terrestrial ones, as the cost of transporting additional adipose depots is higher for climbing than for horizontal locomotion. We did, indeed, find a significant negative correlation between brain size and coefficient of variation (CV) in body mass in both sexes for the subsample of arboreal species, both in all mammals and within primates. In predominantly terrestrial species, in contrast, this correlation was not significant. We therefore suggest that the adoption of habitually terrestrial locomotor habits, accompanied by a reduced reliance on climbing, has allowed for a primate of our body size the unique human combination of unusually large brains and unusually large adipose depots.

Determinants of terrestrial feeding in an arboreal primate: The case of the southern bamboo lemur (Hapalemur meridionalis).

OBJECTIVES: The proximate and ultimate determinants that may have prompted some primates to shift from an arboreal to terrestrial feeding niche, whether due to environmental change, seasonality, and/or predation pressure, are poorly understood. Within a fragmented littoral forest in southeast Madagascar, an arboreal strepsirrhine population spends a large proportion of time on the ground, thus we aimed to identify which factors influence terrestrial feeding. METHODS: From January to December 2013, we conducted 103 full-day focal follows on three social groups of southern bamboo lemurs H. meridionalis. We continuously recorded feeding time on all arboreal and terrestrial items, as well as whether the focal individual was under the canopy or exposed, and the distance to their nearest conspecific neighbor. All observed food items were collected and analyzed for macronutrient content. Daily climatic variables (temperature, precipitation), resource seasonality, daily path length (DPL), along with dietary and predation risk proxies, were used as fixed effects in a linear mixed model, with the daily proportion of terrestrial feeding as the dependent variable. RESULTS: Our model indicated that daily terrestrial feeding increased at cooler temperature, was associated with reduced DPL, and the intake of dietary metabolizable energy increased as terrestrial feeding increased. All other fixed effects were not significant predictors. DISCUSSION: Our study provides a window into the ultimate determinants of niche expansion: ancestral primates, in absence of their primary resources, may have initially descended to the ground in peripheral population range areas where the benefits (e g., nutritional pay-off) out-weighed the costs.

Why do orangutans leave the trees? Terrestrial behavior among wild Bornean orangutans (Pongo pygmaeus wurmbii) at Tuanan, Central Kalimantan.

Orangutans (genus Pongo) are the largest arboreal mammals, but Bornean orangutans (P. pygmaeus spp.) also spend time on the ground. Here, we investigate ground use among orangutans using 32,000 hr of direct focal animal observations from a well-habituated wild population of Bornean orangutans (P. p. wurmbii) living in a closed-canopy swamp forest at Tuanan, Central Kalimantan, Indonesia. Ground use did not change with increasing observation time of well-habituated individuals, suggesting it was not an artifact of observer presence. Flanged males spent the most time on the ground (ca. 5% of active time), weaned immatures the least (around 1%). Females and immatures descended mainly to feed, especially on termites, whereas flanged males traveled more while on the ground. Flanged males may travel more inconspicuously, and perhaps also faster, when moving on the ground. In addition, orangutans engaged in ground-specific behavior, including drinking from and bathing in swamp pools. Supplementary records from 20 ground-level camera traps, totaling 3986 trap days, confirmed the observed age-sex biases in ground use at Tuanan. We conclude that ground use is a natural part of the Bornean orangutan behavioral repertoire, however it remains unclear to what extent food scarcity and canopy structure explain population differences in ground use.

Evolution, ecology and physiology of amphibious killifishes (Cyprinodontiformes).

The order Cyprinodontiformes contains an exceptional diversity of amphibious taxa, including at least 34 species from six families. These cyprinodontiforms often inhabit intertidal or ephemeral habitats characterized by low dissolved oxygen or otherwise poor water quality, conditions that have been hypothesized to drive the evolution of terrestriality. Most of the amphibious species are found in the Rivulidae, Nothobranchiidae and Fundulidae. It is currently unclear whether the pattern of amphibiousness observed in the Cyprinodontiformes is the result of repeated, independent evolutions, or stems from an amphibious common ancestor. Amphibious cyprinodontiforms leave water for a variety of reasons: some species emerse only briefly, to escape predation or capture prey, while others occupy ephemeral habitats by living for months at a time out of water. Fishes able to tolerate months of emersion must maintain respiratory gas exchange, nitrogen excretion and water and salt balance, but to date knowledge of the mechanisms that facilitate homeostasis on land is largely restricted to model species. This review synthesizes the available literature describing amphibious lifestyles in cyprinodontiforms, compares the behavioural and physiological strategies used to exploit the terrestrial environment and suggests directions and ideas for future research.

Ecological divergence and talar morphology in gorillas.

Gorillas occupy a variety of habitats from the west coast to eastern central Africa. These habitats differ considerably in altitude, which has a pronounced effect on forest ecology. Although all gorillas are obligate terrestrial knuckle-walking quadrupeds, those that live in lowland habitats eat fruits and climb more often than do those living in highland habitats. Here we test the hypothesis that gorilla talus morphology falls along a morphocline that tracks locomotor function related to a more inverted or everted foot set. This proposed morphocline predicts that gorillas living in lowland habitats may have a talocrural joint configured to facilitate a more medially oriented foot during climbing, suggesting that they may be more adaptively committed to arboreality than gorillas living in highland habitats. To quantify the relative set of the foot in gorillas, we chose two three-dimensional measurements of the talocrural joint: mediolateral curvature of the trochlea and relative surface area of the lateral malleolus. Our results show that, in comparison to their eastern counterparts, western gorillas have talar features that reflect a more medially directed sole of the foot. This morphology likely facilitates foot placement in a wider range of positions and minimization of shearing stresses across the joint when the foot is loaded on more curved or vertically oriented substrates as occurs during climbing and other arboreal behaviors. In contrast, eastern gorilla talar morphology is consistent with habitual placement of the foot with the sole directed more inferiorly, suggesting more effective loading during plantigrade push-off on terrestrial substrates.

Trabecular bone variation in the gorilla calcaneus.

OBJECTIVES: Calcaneal external shape differs among nonhuman primates relative to locomotion. Such relationships between whole-bone calcaneal trabecular structure and locomotion, however, have yet to be studied. Here we analyze calcaneal trabecular architecture in Gorilla gorilla gorilla, Gorilla beringei beringei, and G. b. graueri to investigate general trends and fine-grained differences among gorilla taxa relative to locomotion. MATERIALS AND METHODS: Calcanei were micro-CT scanned. A three-dimensional geometric morphometric sliding semilandmark analysis was carried out and the final landmark configurations used to position 156 volumes of interest. Trabecular thickness (Tb.Th), trabecular spacing (Tb.Sp), and bone volume fraction (BV/TV) were calculated using the BoneJ plugin for ImageJ and MATLAB. Non-parametric MANOVAs were run to test for significant differences among taxa in parameter raw values and z-scores. Parameter distributions were visualized using color maps and summarized using principal components analysis. RESULTS: There are no significant differences in raw BV/TV or Tb.Th among gorillas, however G. b. beringei significantly differs in z-scores for both parameters (p = <0.0271). All three taxa exhibit relatively lower BV/TV and Tb.Th in the posterior half of the calcaneus. This gradation is exacerbated in G. b. beringei. G. b. graueri significantly differs from other taxa in Tb.Sp z-scores (p < 0.001) indicating a different spacing distribution. DISCUSSION: Relatively higher Tb.Th and BV/TV in the anterior calcaneus among gorillas likely reflects higher forces associated with body mass (transmitted through the subtalar joint) relative to forces transferred through the posterior calcaneus. The different Tb.Sp pattern in G. b. graueri may reflect proposed differences in foot positioning during locomotion.