Maximum trunk tip force assessment related to trunk position and prehensile 'fingers' implication in African savannah elephants.

African elephants have a wide range of abilities using their trunk. As a muscular hydrostat, and thanks to the two finger-like processes at its tip, this proboscis can both precisely grasp and exert considerable force by wrapping. Yet few studies have attempted to quantify its distal grasping force. Thus, using a device equipped with force sensors and an automatic reward system, the trunk tip pinch force has been quantified in five captive female African savanna elephants. Results showed that the maximum pinch force of the trunk was 86.4 N, which may suggest that this part of the trunk is mainly dedicated to precision grasping. We also highlighted for the first time a difference in force between the two fingers of the trunk, with the dorsal finger predominantly stronger than the ventral finger. Finally, we showed that the position of the trunk, particularly the torsion, influences its force and distribution between the two trunk fingers. All these results are discussed in the light of the trunk's anatomy, and open up new avenues for evolutionary reflection and soft robot grippers.

Effect of the habitat and tusks on trunk grasping techniques in African savannah elephants.

Among tetrapods, grasping is an essential function involved in many vital behaviours. The selective pressures that led to this function were widely investigated in species with prehensile hands and feet. Previous studies namely highlighted a strong effect of item properties but also of the species habitat on manual grasping behaviour. African savannah elephants (Loxodonta africana) are known to display various prehensile abilities and use their trunk in a large diversity of habitats. Composed of muscles and without a rigid structure, the trunk is a muscular hydrostat with great freedom of movement. This multitasking organ is particularly recruited for grasping food items while foraging. Yet, the diet of African savannah elephants varies widely between groups living in different habitats. Moreover, they have tusks alongside the trunk which can assist in grasping behaviours, and their tusk morphologies are known to vary considerably between groups. Therefore, in this study, we investigate the food grasping techniques used by the trunk of two elephant groups that live in different habitats: an arid study site in Etosha National Park in Namibia, and an area with consistent water presence in Kruger National Park in South Africa. We characterised the tusks profiles and compared the grasping techniques and their frequencies of use for different foods. Our results show differences in food-grasping techniques between the two groups. These differences are related to the food item property and tusk profile discrepancies highlighted between the two groups. We suggest that habitat heterogeneity, particularly aridity gaps, may induce these differences. This may reveal an optimisation of grasping types depending on habitat, food size and accessibility, as well as tusk profiles.

Show Me Your Teeth And I Will Tell You What You Eat: Differences in Tooth Enamel in Snakes with Different Diets.

Teeth are composed of the hardest tissues in the vertebrate body and have been studied extensively to infer diet in vertebrates. The morphology and structure of enamel is thought to reflect feeding ecology. Snakes have a diversified diet, some species feed on armored lizards, others on soft invertebrates. Yet, little is known about how tooth enamel, and specifically its thickness, is impacted by diet. In this study, we first describe the different patterns of enamel distribution and thickness in snakes. Then, we investigate the link between prey hardness and enamel thickness and morphology by comparing the dentary teeth of 63 species of snakes. We observed that the enamel is deposited asymmetrically at the antero-labial side of the tooth. Both enamel coverage and thickness vary a lot in snakes, from species with thin enamel, only at the tip of the tooth to a full facet covered with enamel. There variations are related with prey hardness: snakes feeding on hard prey have a thicker enamel and a lager enamel coverage while species. Snakes feeding on softer prey have a thin enamel layer confined to the tip of the tooth.

Armed to the teeth: The underestimated diversity in tooth shape in snakes and its relation to feeding behavior and diet.

The structure, composition, and shape of teeth have been related to dietary specialization in many vertebrate species, but comparative studies on snakes' teeth are lacking. Yet, snakes have diverse dietary habits that may impact the shape of their teeth. We hypothesize that prey properties, such as hardness and shape, as well as feeding behavior, such as aquatic or arboreal predation, or holding vigorous prey, impose constraints on the evolution of tooth shape in snakes. We compared the morphology of the dentary teeth of 63 species that cover the phylogenetic and dietary diversity of snakes, using 3D geometric morphometrics and linear measurements. Our results show that prey hardness, foraging substrate, and the main feeding mechanical challenge are important drivers of tooth shape, size, and curvature. Overall, long, slender, curved teeth with a thin layer of hard tissue are observed in species that need to maintain a grip on their prey. Short, stout, less curved teeth are associated with species that undergo high or repeated loads. Our study demonstrates the diversity of tooth morphology in snakes and the need to investigate its underlying functional implications to better understand the evolution of teeth in vertebrates.

Comparative morpho-functional analysis of the humerus and ulna in three Western European moles species of the genus Talpa, including the newly described T. aquitania.

The forelimb is involved in many behaviours including locomotion. Notably, the humero-ulnar articulation, implicated in the elbow joint, is of particular importance for both mobility and stability. Functional constraints, induced in part by environmental plasticity, are thought to drive an important part of the bone shape as bone directly responds and remodels in response to both muscle and external forces. In this context, the study of subterranean moles is of particular interest. These moles occupy a hard and heavy medium in comparison with air or water, requiring a powerful body structure to shear and shift the soil. Their general morphology is therefore adapted to digging and to their subterranean lifestyle. The various morpho-functional patterns, which drive diverse abilities according to the environment, are likely targets of natural selection and it is, therefore, useful to understand the relationships between the bone shape and their function. Here, we quantify, through 3D geometric morphometric methods, the interspecific variability in the morphology of the ulna and humerus of three Talpa species, including the new species Talpa aquitania, to infer their potential consequence in species digging performance. We also quantify shape covariation and morphological integration between the humerus and the ulna to test whether these bones evolve as a uniform functional unit or as more or less independent modules. Our results show that interspecific anatomical differences in the humerus and ulna exist among the three species. Shape changes are mostly located at the level of joints and muscle attachments. As the species tend to live in allopatry and the fossorial lifestyle induces strong ecological constraints, interspecific variations could be explained by the properties of the environment in which they live, such as the compactness of the soil. Our results also show that the humerus and ulna are highly integrated. The covariation between the humerus and ulna in moles is dominated by variation in the attachment areas and particularly of the attachment areas of shoulder muscles concerning the humerus, which affect the mechanical force deployed during locomotion and digging. This study also highlights that in the new species, T. aquitania, variations in anatomical structure (general shape and joints) exist and are related to the locality of collect of the individuals.

Measuring the force of the tip of the elephants trunk.

Forces that animals can exert is of great interest in biology. Regarding the elephant trunk, the maximum mass that an elephant can lift with its trunk is known, but the pinching force of the trunk tip is unknown. We here present an original system to measure this force for an organ much coveted in soft robotics for example. It consists of:•A wooden box protecting the elephants and the measuring system with no protruding parts except the sensor. This box has an opening at the back to fill it with apples and a trap door at the front to release the apples.•Two load cells protruding from the box connected to an electronic system controlled by an Arduino card that records and sends the pinch force via Wi-Fi to a laptop while releasing a reward apple.•Depending on the threshold chosen, the elephant must pinch harder than the previous time to release the next apple. The repetition of this action allows us to approach the maximum force it can achieve.The system, tested on elephants at the Beauval Zooparc (France), has demonstrated that it is effective in measuring the pinching force of the tip of the trunk.

The relationship between distal trunk morphology and object grasping in the African savannah elephant (Loxodonta africana).

Background: During reach-to-grasp movements, the human hand is preshaped depending on the properties of the object. Preshaping may result from learning, morphology, or motor control variability and can confer a selective advantage on that individual or species. This preshaping ability is known in several mammals (i.e., primates, carnivores and rodents). However, apart from the tongue preshaping of lizards and chameleons, little is known about preshaping of other grasping appendages. In particular, the elephant trunk, a muscular hydrostat, has impressive grasping skills and thus is commonly called a hand. Data on elephant trunk grasping strategies are scarce, and nothing is known about whether elephants preshape their trunk tip according to the properties of their food. Methods: To determine the influence of food sizes and shapes on the form of the trunk tip, we investigated the morphology of the distal part of the trunk during grasping movements. The influence of food item form on trunk tip shape was quantified in six female African savannah elephants (Loxodonta africana). Three food item types were presented to the elephants (elongated, flat, and cubic), as well as three different sizes of cubic items. A total of 107 ± 10 grips per individual were video recorded, and the related trunk tip shapes were recorded with a 2D geometric morphometric approach. Results: Half of the individuals adjusted the shape of the distal part of their trunk according to the object type. Of the three elephants that did not preshape their trunk tip, one was blind and another was subadult. Discussion and perspectives: We found that elephants preshaped their trunk tip, similar to the preshaping of other species' hands or paws during reach-to-grasp movements. This preshaping may be influenced by visual feedback and individual learning. To confirm these results, this study could be replicated with a larger sample of elephants.

Masticatory system integration in a commensal canid: interrelationships between bones, muscles and bite force in the red fox.

The jaw system in canids is essential for defence and prey acquisition. However, how it varies in wild species in comparison with domestic species remains poorly understood, yet is of interest in terms of understanding the impact of artificial selection. Here, we explored the variability and interrelationships between the upper and lower jaws, muscle architecture and bite force in the red fox (Vulpes vulpes). We performed dissections and used 3D geometric morphometric approaches to quantify jaw shape in 68 foxes. We used a static lever model and bite force estimates were compared with in vivo measurements of 10 silver foxes. Our results show strong relationships exist between cranial and mandible shape, and between cranial or mandible shape on the one hand and muscles or estimated bite force on the other hand, confirming the strong integration of the bony and muscular components of the jaw system. These strong relationships are strongly driven by size. The functional links between shape and estimated bite force are stronger for the mandible, which probably reflects its greater specialisation towards biting. We then compared our results with data previously obtained for dogs (Canis lupus familiaris) to investigate the effect of domestication. Foxes and dogs differ in skull shape and muscle physiological cross-sectional area (PCSA). They show a similar amount of morphological variation in muscle PCSA, but foxes show lower variation in cranial and mandible shape. Interestingly, the patterns of covariation are not stronger in foxes than in dogs, suggesting that domestication did not lead to a disruption of the functional links of the jaw system.

Bite force and its relationship to jaw shape in domestic dogs.

Previous studies based on two-dimensional methods have suggested that the great morphological variability of cranial shape in domestic dogs has impacted bite performance. Here, we used a three-dimensional biomechanical model based on dissection data to estimate the bite force of 47 dogs of various breeds at several bite points and gape angles. In vivo bite force for three Belgian shepherd dogs was used to validate our model. We then used three-dimensional geometric morphometrics to investigate the drivers of bite force variation and to describe the relationships between the overall shape of the jaws and bite force. The model output shows that bite force is rather variable in dogs and that dogs bite harder on the molar teeth and at lower gape angles. Half of the bite force is determined by the temporal muscle. Bite force also increased with size, and brachycephalic dogs showed higher bite forces for their size than mesocephalic dogs. We obtained significant covariation between the shape of the upper or lower jaw and absolute or residual bite force. Our results demonstrate that domestication has not resulted in a disruption of the functional links in the jaw system in dogs and that mandible shape is a good predictor of bite force.

Contrasting genomic and phenotypic outcomes of hybridization between pairs of mimetic butterfly taxa across a suture zone.

Hybrid zones, whereby divergent lineages come into contact and eventually hybridize, can provide insights on the mechanisms involved in population differentiation and reproductive isolation, and ultimately speciation. Suture zones offer the opportunity to compare these processes across multiple species. In this paper we use reduced-complexity genomic data to compare the genetic and phenotypic structure and hybridization patterns of two mimetic butterfly species, Ithomia salapia and Oleria onega (Nymphalidae: Ithomiini), each consisting of a pair of lineages differentiated for their wing colour pattern and that come into contact in the Andean foothills of Peru. Despite similarities in their life history, we highlight major differences, both at the genomic and phenotypic level, between the two species. These differences include the presence of hybrids, variations in wing phenotype, and genomic patterns of introgression and differentiation. In I. salapia, the two lineages appear to hybridize only rarely, whereas in O. onega the hybrids are not only more common, but also genetically and phenotypically more variable. We also detected loci statistically associated with wing colour pattern variation, but in both species these loci were not over-represented among the candidate barrier loci, suggesting that traits other than wing colour pattern may be important for reproductive isolation. Our results contrast with the genomic patterns observed between hybridizing lineages in the mimetic Heliconius butterflies, and call for a broader investigation into the genomics of speciation in Ithomiini - the largest radiation of mimetic butterflies.

Why has transparency evolved in aposematic butterflies? Insights from the largest radiation of aposematic butterflies, the Ithomiini.

Defended species are often conspicuous and this is thought to be an honest signal of defences, i.e. more toxic prey are more conspicuous. Neotropical butterflies of the large Ithomiini tribe numerically dominate communities of chemically defended butterflies and may thus drive the evolution of mimetic warning patterns. Although many species are brightly coloured, most are transparent to some degree. The evolution of transparency from a warning-coloured ancestor is puzzling as it is generally assumed to be involved in concealment. Here, we show that transparent Ithomiini species are indeed less detectable by avian predators (i.e. concealment). Surprisingly, transparent species are not any less unpalatable, and may in fact be more unpalatable than opaque species, the latter spanning a larger range of unpalatability. We put forth various hypotheses to explain the evolution of weak aposematic signals in these butterflies and other cryptic defended prey. Our study is an important step in determining the selective pressures and constraints that regulate the interaction between conspicuousness and unpalatability.

Does divergent selection predict the evolution of mate preference and reproductive isolation in the tropical butterfly genus Melinaea (Nymphalidae: Ithomiini)?

Many studies have shown that speciation can be facilitated when a trait under divergent selection also causes assortative mating. In Müllerian mimetic butterflies, a change in wing colour pattern can cause reproductive isolation. However, colour pattern divergence does not always lead to reproductive isolation. Understanding how divergent selection affects speciation requires identifying the mechanisms that promote mate preference and/or choosiness. This study addresses whether shifts in wing colour pattern drives mate preference and reproductive isolation in the tropical butterfly genus Melinaea (Nymphalidae: Ithomiini), and focuses on five taxa that form a speciation continuum, from subspecies to fully recognized species. Using genetic markers, wing colour pattern quantification, male pheromone characterization and behavioural assays of mating preference, we characterize the extent of genetic and phenotypic differentiation between taxa and compare it to the level of reproductive isolation. We show strong premating isolation between the closely related species M. satevis and M. marsaeus, in addition to genetic and phenotypic (colour pattern and pheromones) differentiation. By contrast, M. menophilus and M. marsaeus consist of pairs of subspecies that differ for colour pattern but that cannot be differentiated genetically. Pheromonal differentiation of subspecies was significant only for M. marsaeus, although most individuals were indistinguishable. Melinaea menophilus and M. marsaeus also differ in the strength of assortative mating, suggesting that mate preference has evolved only in M. marsaeus, consistent with selection against maladaptive offspring, as subspecific 'hybrids' of M. marsaeus have intermediate, non-mimetic colour patterns, unlike those of M. menophilus which display either parental phenotypes. We conclude that a shift in colour pattern per se is not sufficient for reproductive isolation, but rather, the evolution of assortative mating may be caused by selection against maladaptive intermediate phenotypes. This study suggests that mate preference and assortative mating evolve when adaptive, and that even in the early stages of divergence, reproductive isolation can be nearly complete due to mating preferences.

A new relictual and highly troglomorphic species of Tomoceridae (Collembola) from a deep Croatian cave.

Tritomurus velessp. n. (Tomoceridae) is described from a Croatian cave. It is characterized by troglomorphic features (absence of eyes, reduced pigmentation, slender claw, pointed tibiotarsal tenent hairs) that only compare, among Tomoceridae, to the microendemic species Tritomurus falcifer from the Pyrénées. Tritomurus veles also shares with Tritomurus falcifer the absence of macrochaetae on head, a presumably non-adaptive character that within Tomoceridae is unique to these two species. Both species have no known epigean relatives in their respective distribution areas and can be considered as relictual.